Marasmieae (Basidiomycetes - Tricholomataceae) || Marasmieae (Basidiomycetes - Tricholomataceae)

Organization for Flora Neotropica

Marasmieae (Basidiomycetes - Tricholomataceae)Author(s): Rolf SingerSource: Flora Neotropica, Vol. 17, Marasmieae (Basidiomycetes - Tricholomataceae) (Dec. 28,1976), pp. 1-347Published by: New York Botanical Garden Press on behalf of Organization for Flora NeotropicaStable URL: http://www.jstor.org/stable/4393709 .

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FLORA NEOTROPICA

Monograph No. 17

MARASMIEAE

(BASIDIOMYCETES - TRICHOLOMATACEAE)

by

Rolf Singer

-Vk\ L t*'11

TROPIC

of CANCER

--------------------------

FLORA NEOTROPICA/

TROPIC Of CAPtICORN /

Publishedfor Organization for Flora Neotropica

by THE NEW YORK BOTANICAL GARDEN

Bronx, New York 10458

1976

December 28, 1976

Published with the financial assistance of UNESCO.



Copyright ? 1976

THE NEW YORK BOTANICAL GARDEN

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



A MONOGRAPH OF THE NEOTROPICAL SPECIES OF THE MARASMIEAE (EXCEPTING THE OUDEMANSIELLINAE),

BASIDIOMYCETES - TRICHOLOMATACEAE

ROLF SINGER1

INTRODUCTION

Tricholomataceae tribe Marasmieae Fayod consists of several genera, some of which are particularly well represented by a large number of species in the neotropics and the

tropics in general. These genera have formerly been combined into one large genus Marasmius Fries which has often been confused with Marasmiellus and Collybia. Inso- far as some of these genera have no hymenophore but have a smooth hymenial surface, they have been termed "thelephoraceous" in the classical wide sense, but agreement has gradually been reached that most of these do not belong in the Thelephoraceae as this family is now understood, but rather in agaricoid "cyphellaceous" genera. Some of them are so closely related to Marasmius and Chaetocalathus that there can be no doubt that they are "reduced" forms of the Marasmieae differing only in smooth hymenial surface and, in some cases, "cyphelloid" habit. They have been recognized as genera different from Marasmius and Chaetocalathus in all those cases where additional characters separate them from either of these lamellate genera, aside from the absence of a hymenophore and/or the cyphelloid habit.

In the present monograph I have excluded from consideration all those fungi which do not belong in the subtribes Crinipellinae or Marasmiinae; thus, the subtribe Oudeman- siellinae has been omitted because in an earlier paper (Singer, 1964a) I have given a monograph of the South American species of Oudemansiella and Mycenella which, with very few additions or corrections, can serve as a monograph of the neotropical species of these genera. The third genus of subtribe Oudemansiellinae, viz Physocystidium, consists of a single species, P. cinnamomea (Dennis) Singer, which has been adequately treated by Dennis (1951a) and Singer (1962a, 1973), whereas the fourth one, Strobil- urus, is not found in the neotropics proper, but only in the north temperate zone.

This leaves the Crinipellinae (with pseudoamyloid hairs or appendages on sterile surfaces) and Marasmiinae (without such hairs). The morphological features have been amply discussed in previous papers, particularly my earlier monograph of Marasmius (Singer, 1965), Crinipellis and Chaetocalathus (Singer, 1942) and the cyphelloid Maras- miinae (Singer, 1960). Aside from general data on the morphology and chemical characters, the reader will find in these papers indications on the best way to collect and annotate specimens of these fungi. The delimitations of the various genera in- volved and the scope of all genera of the Marasmieae on a world-wide basis are discussed in Singer (1974).

While preliminary studies on many genera of Marasmieae have been published by Singer (1942, 1965) in the past for the South American continent 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.

1



2 Flora Neotropica

of the known species of the world flora, it has become obvious that there is a need for a new treatment including the numerous Central and North American species while omitting the exclusively temperate ones. At the same time there is a need to consider new taxa, discovered in regions where Marasmieae have been collected before - and especially in regions where mycologists have never been active until after 1965, such as the Cordilleras of Colombia and the Amazonas region of Ecuador. Including, then, the new discoveries of the period between 1965 and 1973 but excluding the Oudeman- siellinae, the Marasmieae as represented in the present monograph contain 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 Marasmieae are among the most important flor- istic elements of that vast region, followed by the Collybieae (compare my monograph of the neotropical species of Marasmiellus Singer 1973).

PHYTOGEOGRAPHY AND ECOLOGY

The species of Marasmieae, particularly the Marasmiinae, are mostly wood- and litter-inhabiting fungi, all of which are epigeous with exogenous spore production. None of them are ectomycorrhizal. Most are saprophytic, but several are decidedly, some even exclusively, parasitic (e g Crinipellis perniciosa (Stahel) Singer, C. siparunae Singer, Maras- mius viegasii Singer, M. crinisequi Muller ex Kalchbrenner), and it is these species which are of interest to the tropical phytopathologist. In contrast with the genus Marasmiellus and with Omphalina and Gerronema, lichenization has never been observed in the Marasmieae. Some of the saprophytic species seem to be specialized as far as the host material is concerned. While some species occur on a wide range of hosts, others are limited to conifers, Pteridophyta, Monocotyledones or Dicotyledones. Finally there is a rather large group of species which appear to be restricted to a single host genus or family. Some species are obviously restricted to high-alpine (andine) zones reaching up to abott 5000 m altitude; others are restricted to subxerophytic and xerophytic regions. There are also some species which are most common in destroyed vegetation or wherever ecosystems are in disequilibrium caused by human activities. The rest is restricted to the tropical and subtropical wet forest. Many of the latter appear to grow only in the subtropical-tropical montane 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 Panama south to Ecuador. The determining factors for such area restriction cannot, in most cases, be ascertained. Certainly, sometimes it is geographical isola- tion, sometimes restriction of the host species and sometimes climatic factors, particularly 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-inhabitants or inhabitants of dead fallen woody material, or else parasitizing or otherwise epiphytic on living plants.

The area of distribution, as can be seen from the preceding paragraph, varies from very restricted island-like areas within some of the major landscapes of the neotropics to panneotropical or even pantropical. Some of the pantropical species such as Maras- mius haematocephalus will occasionally occur in the temperate zone of North America,

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



Introduction 3

where they are introduced with plant material from the subtropical and tropical zones. Others frequently invade the greenhouses of the temperate zones e. g. Crinipellis siparunae. If I exclude the collections of adventitious species from the statistics, I may say that of the approximately 300 well defined species of Marasmius less than 10% are pantropical and as many as 229 have been recognized as occurring in the neotropics. This hligh percentage is not only due to the relatively large, perhaps exceptionally large rep- representation of Marasmius in the neotropics but, to a certain degree, also to less extensive collecting and relatively few monographic revisions (Ceylon, Zaire) in the paleotropics. A similar preponderance of neotropical species can be observed in other genera such as Gloiocephala, Crinipellis, Chaetocalathus. The genera Physocystidium, Manuripia, Epictwphus, Hymnenogloea have thus far been observed only in the neotropics and only Palaeocephala exclusively in the palaeotropics.

The statistics for the whole of the Marasmieae show that among the species of Marasmieae (without Oudemansiellinae) treated in the present monograph there are

species belonging in the following four groups; Group I. Species occurring exclusively in the subtropics and/or tropics of the Americas

or in parts of it: 274 species or 85.9%

Group II. Species occurring aside from an area in the neotropics also in some parts of the paleotropics, Oceania or Australia: 21 species or 6.6%

Group III. Species occurring aside from some parts of the neotropics also in the temperate 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 palaeotropics are gradually explored so that eventually

our knowledge of the Marasmieae flora of the paleotropics will match the knowledge we have now of the neotropical species, the percentage of the species belonging in group II will turn out to be larger than 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 percentages reported above will be substantially altered after future research, appear to be small.

MYCOSOCIOLOGY

Since ecologists - not so inexplicably, since it is difficult to obtain the collabora- tion of a competent mycologist - have, in spite of their important role, omitted the

fungi from consideration in 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 observations were not coordinated with a study of

phanerogamic or other cryptogamic vegetation. One might wish that scientists active in elaborating and 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 consideration to the fungi or protists.

Five localities with virgin, undisturbed vegetation of the Amazonas region whose

non-fungus vegetation can easily be checked, have been selected in order to see whether comparable aspects (i e individuals fruiting during the main rainy season) of the fruiting consociation are similar. These localities have been subdivided into (1-3) terra-firme vegetation of the Amazonian lowlands and (4-5) inundated or inundable



4 Flora Neotropica

forests just after the retreat of the flooding. In these localities extensive and intensive collecting of Marasmieae has taken place (other collections have been added in

brackets). The results are interesting insofar as surprisingly few species are common to the

five localities even within their grouping according to inundability. This may mean two things, one exclusive of the other: either Marasmieae (and obviously other agarics) are, as a group of organisms, not characteristic for 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-distribution and geographical iso- 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 phytogeographic and ecological data in the keys and species descriptions shows that many species are selective as far as area and association 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 spectacular aspect during its flowering period or with recognizable features (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 characteristic for the partic- ular forest type, are widely spaced, often with distances of over 1 km between individuals. If it is assumed then, given the dependence of the heterotrophic organisms on their hosts, that fungus species may be equally scattered in the forest and that even

repeated visits and very intensive collecting will not register every population in fruit-

ing - species cannot be detected when in the vegetative state - it becomes clear that the necessarily reduced collecting area and the fungi observed in it, cannot by any means be complete enough to produce fully comparable lists.

On the other hand, if the Marasmieae were, although widely scattered in the Amazonas Basin, nevertheless evenly distributed among 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 subtropical forest type in the neotropics where this fungus does not occur, and the same species has also been observed in the palaeotropics and, adventitiously, in the temperate zone of North America.

Somewhat more similarity between localities 1, 2 and 3, and 4 and 5 respectively would be apparent if we were able to add all collections from the Napo river region and compare them to all localities visited in the Beni, Mamore and Madre de Dios regions, or all collections of the lower Para and all gathered in inundated forests of the upper Amazonas and 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 deterioration to a variable degree; 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 particularly wide area including some cosmopolitan species and

species forming cicatrizing mycorrhizae. Consequently, we have to rely on the data given here or else we might obtain misleading results. 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 destroyed at an alarming rate not only in the Amazon region but also over the entire neotropics. This is also the reason why the thus far somewhat inconclusive results presented here are believed to merit publication.

As far as other regions of the neotropics are concerned, similar lists are now in preparation. The data presented here (Table I-III) as well as others referring to the mycosociological exploration of the neotropical plant communities suggest, if nothing else, that it is certainly worth the trouble to collect data on the fungus component of each association whenever ecological work on neotropical forest communities is under- taken. The present monograph will help to make identification of such collections easier.

TABLE I

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- Manuripia bifida 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]

geonematis] [Simocybe amazonica]



6 Flora Neotropica

TABLE II

Amazonas, Inundable river-side forest

4. March aspect in inundable forests along 5. May-June aspect of the inundable rain the lower Madre de Di6s up to Conquista. forest in the lower Amazonas. BRAZIL: Bolivia: Pando Para', Belem

Marasmius tereticeps Marasmius rotuloides M. guyanensis M. cladophyllus M. pandoanus M. leoninus M. dodecaphyllus [Marasmiellus stenophylloides] M. panerythrus [Marasmiellus defibulatus var strictior] Gloiocephala longifimbriata [Marasmiellus bisporiger] Crinipellis eggersii var flavipes [Marasmiellus aporposeptus] C. excentrica [Marasmiellus nigripes] [Marasmiellus pandoanus] [Hydropus paraensis] [Trogia cantharelloides] [Hydropus depauperatus] [Pluteus neophlebophorus] [Hydropus semimarginellus] [Inocybe matris-dei] [Gerronema strombodes] [Pterula palmicolal [Mycena aosma]

[Mycena biornata] [Lepiota xanthophylla] [Russula batistae]

TABLE III

6. Tropical rain forest, spring-summer aspect, 7. Tropical rain forest, summer aspect, Gulf Coast (Estaci6n Biologica de los Tuxtlas, Pacific Coast low hills region near Tuxtla de (nearly virgin) Gutierrez: Selva between Finca Sospiro and

El Pozo (nearly virgin)

Marasmius leveilleanus Marasmius chiapasensis Marasmius carpenterianus Marasmius atroincrustatus var inodorosus Marasmius floriceps Marasmius brunneocinctus Marasmius haematocephalus Marasmius dictoyledoneus Marasmius rubroflavus Marasmius xerampelinus Marasmius trinitatis Marasmius chrysoblepharis Marasmius neosessilis var montepiensis Marasmius haematocephalus [Many species of Agaricales, Aphyllophorales Marasmius longisporus

and Gasteromycetes, see Singer (to be pub- [Micromphale separatum] lished, none of them observed in area 7.1 [Mycena maculata var]

[Pluteus nitens] [Psathyrella crinipellis] [Favolaschia selloana]

CIRCUMSCRIPTION OF THE AREA

While I adhere in principle to the definition and limits of the area covered by the neotropics as accepted by the Organization for Flora Neotropica, it may be useful to point out here that the Marasmieae have a mycelium protected by the host tissue or substratum and the majority of species is found in spots with comparatively favor- able microclimatic conditions. Consequently, many basically tropical species pass over into the subtropical, a few even in the warm-temperate zone of both northern and southern hemisphere. Because of such a relatively unaltered and gradual transition of the flora, it was believed to be preferable to admit a species to treatment in Flora Neo- tropica even if it was found only in the subtropical zone - provided its precise area of distribution is insufficiently known - i e, occurrence in the intertropical region



Introduction 7

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

NOTES ON DESCRIPTIVE DATA AND ILLUSTRATIONS

In the descriptive data, colors are, unless otherwise indicated, those of Maerz & Paul (1930), designated either by name given in quotation marks or by number. Other terms are the ones commonly used in agaric taxonomy. Where definitions or discuss- ions of the terms are desirable, the reader is referred to Singer (1942, 1958b, 1965, 1974). The data on the types are given briefly only where precise data are indicated under "Material studied." Data on the known distribution can be found after description of the habitat or else they can be deduced from the material studied. The distribution can rarely be given in precise and detailed form so that a special paragraph on distribution is not included.

Many taxa have been illustrated in the papers cited in the literature; these have been cited at the end of each description and discussion in the descriptive part. The original illustrations of the present paper correspond to the following magnifications: carpophores and carpophore parts (pileus, etc) X 1/2; spores X 1000; other microscopical 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 number is indicated in the Index of Exsiccatae

Herbaria are quoted according to the Index Herbariorum (Holmgren & Keuken

1974) with the corresponding abbreviations.

ACKNOWLEDGMENTS

The present monograph has been prepared and edited with the help of two grants, GB 7477, GB 35471, from the National Science Foundation, Washington D.C.



8 Flora Neotropica

SYSTEMATIC TREATMENT

Tricholomataceae tribe Marasmieae Fayod, 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 appendiculis 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

carpophores cyphelliform.

TYPE GENUS. Crinipellis Patouillard.

Key to the Genera of Subtribe Crinipellinae

1. Long, conspicuous, mostly pseudoamyloid or else incrusted and thick-walled hairs present. 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. Crinipellis Patouillard, Jour. de Bot. 3: 336. 1889. TYPE SPECIES. C. stipitaria (Fries) Patouillard. Habit of the carpophores collybioid, marasmioid, or rarely pleurotoid, but always

with a well developed true stipe: pileus and stipe covered with thick-walled hair-like elements whose wall is partly or entirely pseudoamyloid; these hairs long and without

clamped septa, cyanophilous, rarely and slightly incrusted, most frequently without incrustation and smooth. Hymenophore always 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-walled and eventually sometimes septate, inamyloid, more rarely pseudoamyloid, cyanophilous, smooth; basidia without sideropliilous granulosity, generally 4-spored, sometimes somewhat deformed

(basidioles not sporulating); basidioles fusoid, rarely (in species with very voluminous

spores and basidia) more clavate-subcylindrical; stipe insititious, rarely with a distinct basal mycelium which is fibrillose or tomentose, or with white rhizomorphs; trama

inamyloid, clamp connections present. Saprophytic and parasitic on plants and plant debris, stems, roots, fruits, woody material, etc.

Development of the carpophores: unknown; the youngest carpophores seen have the hymenial primordium covered by the hairs of the margin of the pileus and the

apex of the stipe.

Key to the Sections and Subsections of 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) condition: 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.

Crinipellis sect Crinipellis

Pileus not brightly colored and not turning green or green-black with KOH or NaOH, and the hairs not greenish or greenish-melleous in 5% KOH; stipe generally central and

pigmented, insititious. The type is that of the genus.

Crinipellis sect Crinipellis subsect Macrosphaerigerae Singer, 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 9.5 u, 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 p1 broad, some hyaline or some brownish-melleous, strongly and entirely pseudoamy-



Crinipellis 11

loid, with the wall 1.4-2.8 , thick wall, not secondarily septate, very long, with rounded tips.

On living tree in subtropical forest MATERIAL STUDIED. Known only from the type locality: BRAZIL. Rio Grande

do Sul: Estacao Sao Salvador, A. Sehnem 1917, 21 Jan 1946 (LIL: C 8339, type).

Crinipellis sect Crinipellis subsect Stipitarinae Singer, Lilloa 8: 461. 1942. Spores much smaller or much more elongated than in subsection Macrosphaer-

igerae; (pleuro-) cystidia absent, or present only as cystidioles in old carpophores (or where the lamellae have been bruised in situ) or as cheilocystidia (scattered at a short distance from the extreme edge); if as cystidioles, these are degenerated basidioles.

TYPE SPECIES. C. stipitaria (Fries) Patouillard.

Key to the Species of Subsection Stipitarinae

1. Spores 9-14.5(-20) X 3.5-6.3,u, often curved Q 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 apical finger-like appendages; rarely entire pileus whitish, drying to gilvous (Hebeloma-sinapizoans-color). 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, ochraceous 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 subacute tips, minority with obtuse tips. 8. C. bisulcata.

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



12 Flora Neotropica

11. Pileus with or without a small papilla or umbo, and not or inconstantly 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. pseudostipitaria var pseudostipitaria. 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 entire.

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 subdistant, 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 appendages. 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 broader than 5.5,; lamellae white or cream to buff when fresh.

25. Cheilocystidia with mostly apical and rather numerous appendages; pileus not broader than 6 mm when dried (if the stipe is central, see "17" above; if the stipe is eccentric, see section Excentricae).

25. Cheilocystidia often branched but not only apically; pileus as above or 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 rarely up to 2) (see "19" above); if spores 9-11 X 6-7.5u and pileus somewhat larger at timies, compare. C. molfinoana p25.

26. Pileus about 18 mm broad, rather evenly pilose; spores 8.5-9.6 X 5.5-6.2u 23. C. dicotyledonum.

14. Spores up to 6.3 X 4.2tp. not larger. 11. C schini.

Stirps Carecomoeis

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

2. Crinipellis carecomoeis (Berkeley &Curtis) Singer, Lilloa 8: 487. 1942. Fig 1. Agaricus carecoimoeis Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 284. 1868. Collybia carecozmoeis (Berkeley & Curtis) Saccardo, Sylloge Fungorum 5: 232. 1887. TYPES: Wright, Cuba (FH, K) Pileus ochraceous brown to rusty brown in the center when fresh, margin uniform-

ly white or white-rimose between curly strands of hairs which are concolorous with the center, radially plicate-pectinate-pilose, center tomentose-pilose and 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 convex 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 margin in 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 assuming a pale orange buff shade, rather narrow to 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 ferrugineous below, but fading to whitish to dirty light brown or pale argillaceous and eventually unicolorously so, appressedly pilose 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 depression on the middle portion of the iner side or even sausage-shaped, hyaline, thin-walled, unicellular, overaged spores mostly yellowish, thick-walled, and sometimes with a cross-wall near 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; basidioles fusoid-subclavate; cystidia none, but basidioles in old specimens often remaining sterile and transformed into cystidioles which are ampullaceous to 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 branched nor crowned with finger-like appendages or 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-walled to very slightly thick-walled, or wall locally somewhat thickened (to 1.5 u), hyaline to melleous-hyaline. Hyphae of the regular hymenophoral trama with clamp connections, hyaline, inamyloid. Covering layers: hairs of the pileus 3.5-11 p broad, with acute or rounded tips, with stramineous to melleous or (near margin 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-septate hyphae which are broader than those of the trama proper (5-8 ,u broad), with hyaline, rather thin walls. Hairs of stipe much like the hairs of the margin of the pileus.

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 gregarious, 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 varieties subelata and litseae which may also be separate species 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 numerous and conspicuous cystidioles on the sides of the older lamellae. This may be a bisporous-parthenogenetic form of the same species, or possibly a related species; it was also the only one on Gramineae. The characters of this form have not been included in the general description of the species, but the specimen is cited under "Material studied."

3. Crinipellis gracilis 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 centrali inserta, 1-2 mm lato. Lamellis candidis, subangustis, subarcuatis, confertis, liberis vel subliberis. Stipite albo, mox atrocastaneo ad 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 subnullis vel nullis; cheilocystidiis subventricosis, excrescentiis nonnullis brevibus nodosis apicalibus instructis, hyalinis. Hyphis in stipite parallelis, filamentosis, crasse tunicatis, brunneolo-tunicatis, inamyloideis. Tegumentis: crinibus pilei 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 secundario subapicali interdum praesente. Crinibus stipitis brevibus vel longis (brevibus subulatis), crasse tunicatis (parietibus 2-2.8 p crassis), 7-8.5 p diametro, omnibus vel plurimis acutis.

KOH in pileo sicco colorem haud mutat. Ad folia Monocotyledonum (probabiliter Aracearum), gregatim, Pileus white with cinnamon-ochraceous center, minutely pilose with appressed,

somewhat curly hairs except for the naked papilla, but at the margin hairs 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 carpophores mature, 1-2 mm broad. Lamellae pure white, rather narrow, subarcuate, close, free or subfree. Stipe white, soon becoming chestnut-fuliginous from the base upwards and dried entirely 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-walled when 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 parallel with each other, filamentous, thick-walled, brown-walled, inamyloid. Covering layers: 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 subapical secondary 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. MATERIAL STUDIED. BRAZIL. Pernambuco: Mun. Sao Lourenco, Mata do Sao, Joao,

16 Jul 1960, Singer B 3412 (F, typus). This is similar to, though still smaller than C. careconzoeis, but has slightly

smaller spores, less pigmentation especially in youth, and a glabrous papilla; it grows on monocotyledonous dead leaves in the tropical forest.

4. Crinipellis urbica (Montagne) Dennis, Kew Bull. 1951: 409. 1951. Fig 2. Agaricus urbicus Montagne, Ann. Sci. Nat. 1: 94. 1854. Collibia urbica (Montagne) Saccardo, Sylloge Fungorum 5: 218. 1887. TYPE. Leprieur 989, 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, radially sulcate, convex, soon with depressed center 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; cheilocystidia 21-30 X 6-8.5 ,. either entire, or more often with apical appendages, these finger-like and obtuse, 3-7 X 1.3-3 4, all hyaline, main body ventricose to cylindric-subventricose. Hyphae of trama with clamp connections, inamyloid. Covering layers: 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-obtuse tips.

On sandy soil among grass, often on grass roots, often in dense groups. MATERIAL STUDIED.FRENCH GUYANA. Cayenne, Leprieur 989 (PC), typus. BRAZIL.

Rio de Janeiro: Cabo Frias, Caqueiral, 25 Jan 1967, Jezek 4 (BAFC). This species has relatively narrower spores, less pigmented pileus, less acute hairs

on the pileus, fewer simple and entire cheilocystidia and relatively shorter stipes in an



16 Flora Neotropica

average of a population and a strictly neotropical distribution, than C. stipitaria. It differs from C. pseudostipitaria var mesites in less pigmented surfaces, less close lamellae, slightly larger spores and the absence of a papilla.

5. Crinipellis subtomentosa (Peck) Singer, Lilloa 8: 463. 1942. Marasmius subtomentosus Peck, BulL Torrey Club 22: 487. 1895. Marasmiuzs tomentosuis Peck in Ellis Everhart, North American lFungi II 3403. 1903 (non

Quelet). Marasmiius dasypuis Spegazzini, Anal. Mus. Buenos Aires 19: 264. 1909. Marasmius nolaneiformis Murrill, Bull. Torrey Club 67: 149. 1940.

TYPE. Bartholomew, Kansas Fungi 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 appearing zonate, often becoming coarsely sulcate over the marginal portion, otherwise smooth, neither viscid nor radially long-

pilose, opaque, campanulate or convex, then applanate and often with depressed center, or with a low, obtuse umbo, 8-25(-35) mm broad. Lamellae pale cream to

grayish ochraceous or sordid flesh color ("ahnong", "aloma", even reaching "army br.", "tawny, birch" or between "fawn" and "Sandal wood" when old), not intervenose but often forked, irregularly intermixed, moderately broad (to 2.7 mm), distant or subdistant, rounded and very narrowly attached 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, immersed portion (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, inodorous, 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 narrower than the 4-spored ones; cystidia none (or some cheilocystidia 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, subcylindric to 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 gelatinous, thin-walled or in old carpophores partly somewhat thick-walled; hymenophoral trama regular, of filamentous, slightly interwoven hyphae; subhymenium of small ele-

ments, some rectangular and in chains, some more elongated. Covering layers: hairs of the pileus not very long in comparison with related species, usually not longer than about 200 pu, somewhat irregular in 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 stramineous in KOH; hypotrichium stramineous. Hairs of stipe similar.

On old grass roots and sometimes passing over to other plant debris, often sub-



Crinipellis 17

merged ones in sandy soil or earth, gregarious, fruiting from January until July, in North America perhaps also 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. mauretanica Maire (1928) is the same species, but this has longer hairs, intervenose lamellae, filiform

cheilocystidia and grows on "buried plant debris" in Africa. It is A. chortophilus Berkeley.

ILLUSTRATION. Singer (1942) fig 2 d.

6. Crinipellis megalospora Singer 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

margin but not white, when faded "spruce yellow" appressedly hairy on whitish ground with the disc "camels hair," appressedly pilose all over and in dried condition the white

background showing between the hairs, convex with applanate to depressed disc around which there usually is a circular furrow, not papillate, but margin strongly declivitous, 4-10 mm broad, mostly 5-7 mm. Lamellae white or whitish, rather narrow, close, subfree. Stipe ochraceous brown, light cinnamon or light grayish cinnamon when dry-faded, grayish cinnamon 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-aged spores thick-walled and 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 basidia or basidioles), 19-50 X 4-8.3 u, or broader where branched, ventricose to clavate and with forked apex or with up to 5

apical finger-like appendages, forks and main body also sometimes with lateral knobs or short branches or bulges, hyaline. Hyphae with clamp connections, hyaline, inamyloid. Covering layers: 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-obtuse or even rounded, with or without cross septa or at least one cross-

septum underneath the apex, but not consistently nor distinctly ladder-like, in KOH melleous to hyaline, some fulvous in ammonia, those of the extreme margin sometimes with some hook-like outgrowths or irregularities, otherwise those of the margin like those of the center. Hairs of the stipe similar.

On the base of tufts of grass and on grass roots in open places. South America, 1000-1200 m alt. Growing gregariously. Fruiting in the rainy season.

MATERIAL STUDIED. ARGENTINA. Tucuman, Anta Muerta, 26 Dec 1951, Singer T 1666 (LIL), holotype.

7. Crinipellis perpusilla (Spegazzini) Singer, Lilloa 25: 494. 1952. Lentinus perpusillus Spegazzini, Bol. Acad. Nac. Ci. Cordoba 11: 406. 1889. Crinipellis bamnbusae Patouillard, Bull. Soc. Mycol. France 24: 8. 1908. Marasmius bambusae (Patouillard) Saccardo & Trotter, Sylloge Fungorum 21: 113. 1912.

Type. Puiggari 1644, from Brazil. Pileus whitish to pale tan or subrufescent, densely covered with minute almost



18 Flora Neotropica

silky appressed hairs, with smooth or somewhat sulcate margin, convex then applanate, often umbonate, 1-2 mm broad. Lamellae white, subdistant, narrowly adnexed. Stipe whitish, faintly pilose-silky, but darker in 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-walled and hyaline, becoming somewhat 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-like apical branches or appendages or with short projections giving them the profile of a fir tree, but often forked (branches or appendages 7.5-17.5 j, long). Hyphae inamyloid, with clamp connections, hyaline. Covering layers: hairs of the

pileus (3-5)7 s, in diameter, thick-walled, with tapering but obtuse apex, more rarely broadly rounded at the tip, pseudoamyloid.

On Bambuseae, gregarious. MATERIAL STUDIED. BRAZIL: Sao Paulo, Campinas, Noack 34 (FH) (type of C. bam-

busae). Apiai, Oct 1881 (LPS), type. In this section, C. perpusilla and C gracilis are the smallest species of the Mono-

cotyledones-inhabiting forms, but they are easily distinguishable from each other, by the size and shape of the spores.

ILLUSTRATION. Singer (1942) fig 2 e.

8. Crinipellis bisulcata (Patouillard & Gaillard) Patouillard, Jour. de Bot. 3: 336. 1889.

Collybia bisulcata Patouillard & Gaillard, Bull. Soc. Mycol. France 4: 14. 1888.

Type. Gaillard 136 (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, apparently the deeper the umbilicus the darker the 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 additional concentric furrows further outwards, 8-13 mm broad. Lamellae initially probably white but concolorous with the margin of the pileus when dried, moderately broad, medium distant to rather distant, almost free or very narrowly adnexed. Stipe concolorous 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 peripheric line in the dried material.

Spores (6.3-)7-8.5 X 3-3.8 u Q=?2, hyaline, smooth, inamyloid. Hymenium: basidia 35-41 X 6.5-8 ,; cystidia none seen; cheilocystidia 27-56 X 4-7.5 ,, simple or with short protuberances up to 6.5 ,u long at the apex of the main body or all over, rarely with 2-3 longer branches up to 14 ,, never truly echinate of the Siccus-type. Hyphae with clamp connections. Covering layers: 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-stramineous in KOH, pseudoamyloid, sometimes septate below the apex by a secondary septum, not ladder-like.

On small stalks of Monocotyledones, apparently on grasses, fruiting from March to April.

MATERIAL STUDIED. VENEZUELA: Orinoco-region, Atures, A. Gaillard 136 (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

ILLUSTRATION. Patouillard and Gaillard (1888) P1 7, fig. 3.

9. Crinipellis pseudostipitaria Singer, Lilloa 8: 470. 1942, subsp pseudostipitaria var pseudostipitaria.

Crinipellis pseudostipitaria subsp occidentalis Singer, Lilloa 8: 470. 1942. TYPE Duss 492, from Guadeloupe (FH). Pileus rusty fibrillose-woolly with 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 appressed fibrillose-pilose covering, in dried material sometimes with concentric walls and furrows but these much less distinct and less constant than in C. bisulcata. hemispherical or almost conical when young, then convex eventually applanate, with depressed center, sometimes distinctly umbilicate, 3-13 mm broad. Lamellae white when fresh, close or subclose, rarely subdistant, sometimes some forked, subventricose, narrow to medium broad, attenuate and narrowly adnexed to subfree. Stipe concolorous with the pileus or more bister especially where the hairs have been rubbed off, spiny strigose from agglutinated hirsute hairs below, 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 several apical and/or lateral finger-like appendages 3-12 p long and sometimes spreading to 12 p in breadth. Hyphae with clamp connections, hyaline in KOH, inamyloid. Covering layers: hairs of the pileus 4-7 p

broad, light melleous, some hyaline, pseudoamyloid, with walls 1.2-2.5 ju thick, with rounded, more rarely attenuate-obtuse tips, without ladder-like secondary septa (only some hairs with scattered septal); hairs of stipe 5-7.7 p broad, with attenuate-obtuse tip, otherwise like those of the pileus.

On dead fallen grass debris and other monocotyledonous plants, sometimes passing over onto other herbaceous stems or small sticks rotting on the ground nearby, gregarious, fruiting in the rainy season. Known hosts: Panicurn maximum.

MATERIAL STUDIED: CUBA: Wright 855 (FH), probably belonging here. GUADELOUPE: Morne Hirondelle, Duss 1 765 (FH); Camp Sacal, Duss 1888 (FH, this and preceding collection are paratypes); Plateau des Rivieres, Duss 492 (FH, type). BOLIVIA. La Paz: Nor-Yungas,Charobamba, Singer B 720, 30 Jan 1956 (F).

9b. Crinipellis pseudostipitaria var mesites Singer, Lilloa 8: 471. 1942. Fig 4. TYPE. Duss 1877, from Martinique (FH). Pileus ferruginous-chestnut, drying to ochraceous buff with umber center, margin

sometimes pale stramineous, and fimbriate hairy-tomentose all over, with a small and rather low papilla, with almost sulcate margin, conic-convex to strongly convex, then more applanate, 5-10 mm broad. Lamellae white, rather narrow, close to almost subdistant, subfree. Spore print white. Stipe concolorous with pileus, tomentose-pilose to strigose-pilose, insititious, equal, 9-35 X 0.3-1.5 mm. Context white, inodorous.

Spores 7-10 X 3.7-6 ,, with rather variable Q (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 neighborhood of the edges, rarely simple and entire and then mostly clavate, frequently forked or with three apical branches, but also apically appendiculate with 3-5 branches making the cheilocystidium hand- or Ramaria-shaped, very rarely with strictly apical setula-like appendages (en brosse),



20 Flora Neotropica

hyaline. Hyphae hyaline with clamp connections, inamyloid. Covering layers: hairs of the pileus similar on margin and 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-obtuse or broadly rounded at the tip, rarely with an obtuse appendage at the tip; Hypotrichium consisting of hyaline clamped hyphae with rather numerous

septa. hairs of the stipe 5-7.7 p broad, with 1.8-2 p thick pseudoamyloid wall, with attenuate-obtuse, more rarely broadly rounded tip.

On dead grasses and Bambuseae, gregarious, fruiting in the rainy seasons. Known hosts: Andropogon sp; Chusquea sp.

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 intermediate between subsp pseudostipitaria and subsp orienltalis, the latter having spores broader than 5 p (none narrower) growing exclusively in the

Palaeotropics.

10. Crinipellis catamarcensis Singer, 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-hairy with undulating appressed pilose covering all over but near margin in dry condition the white trama of hypotrichium showing through among the strands of hairs at least when mature, with a circular elevation around the disc, or near the margin, showing a concentric circular slight depression, but at first

convex, later more repand and mostly obtusely low-umbonate, never abruptly papillate, about 21 mm broad. Lamellae cream color (e g 9 C 3 M&P), about 2 mm broad, close, free. Stipe nearly concolorous with the pileus, appearing fibrillose from numerous long hairs, attenuate towards the insititious base, about 18 X 3.5 mm. Context white, thin, inodorous.

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 pseudoamyloid ones present, not septate. Hymenium: basidia 26-27 X 6.5 u; cystidia none; cheilocystidia 20-25 X 6.5-7.5 ,, clavate or ventricose, not appendiculate or with 1 to several apical appendages 1-14p long and then often spreading above and broader than 7.5 u, rarely longer than 25 p (reaching 43 , in some cells), all hyaline. Hyphae of the regular hymenophoral trama inamyloid and with clamp connections. Covering layers: 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-obtuse ones

present, on the margin nearly 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 subxerophytic vegetation, 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. Crinipellis schini Singer, sp nov TYPE. Argentina, Catamarca, Singer T 1 793 (LIL) Pileo brunneo, appresse piloso, umbonato-papillato, 8-19 mm lato. Lamellis albis



Crinipellis 21

vel cremeis, liberis. Stipite subconcolori, appresse piloso, 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 putrescens Schini aliorumque Dicotyledonum in zona subtropicali subxerophytica montana. 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," appressedly pilose all over, convex, later applanate in the wide marginal zone, soom depressed or flattened around a large conspicuous papilla, this persistent, diameter 8-19 mm. Lamellae pure white to pale cream, subclose, broad, subventricose, free. Stipe almost concolorous with the pileus, appressedly pilose, 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 suprahilar depression, hyaline, many pseudoamyloid, smooth, Hymenium: basidia 27.5 X 6.2 p, 4-spored; cystidia none; cheilocystidia 15-34 X 6.8-8.2 ,clavate or cylindrical or 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. Covering layers: hairs of the pileus (middle between center and margin) varying from deep melleous to hyaline, thick-walled with or more frequently without secondary septa, and these if present not forming a ladder-like structure, 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.

MATERIAL STUDIED. ARGENTINA. Catamarca: Hill above the Estancia Santa Rosa near the Rfo Campo, 20 Jan 1952, Singer T 1793 (LIL, type); between Rio Chacras and Rio Pot- rero, 11 Feb 1952, Singer T 1855 (LIL).

The small pseudoamyloid spores remind one of C. catamarcensis, a species from the same region but which grows on Monocotyledones rather than on Dicotyedones and has a nonpapillate pileus and both absolutely and relatively shorter stipe. The lamellae are paler in C. schini.

12. Crinipellis atrobrunnea Patouillard, 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 surrounded by a series of concentric furrows or one circular depression, young convex, later more repand and more or less umbilicate, 9-17 mm broad. Lamellae pure white when fresh but tending to become brown in the herbarium, rather broad, moderately close or distant, with 12-29 lamellae + lamellulae, rounded-free or attenuate-subfree. Stipe concolorous or "Cochin" M&P, strongly hairy with strands of 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 branching with up to five usually more or less straight and 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 branched the branches more rarely curved or crooked, and these rarely subacute or acute, in dried material often strongly agglutinated, hyaline. Hyphae of the trama filamentous, inamyloid, nongelatinized with clamp connections. Covering layers: hairs of the pileus 4.3-10.5 broad, in KOH and ammonia stramineous-yellowish to deep melleous, the paler ones in dried material often showing (necro-)pigment precipitated in brown granules, with distant secondary septa or without them, but without ladder-like structure even near the mostly broadly rounded tip, very rarely some with such a structrue, with 0.7-5.5 p thick wall, slightly but distinctly pseudoamyloid.

On grass culms and grass roots, often densely gregarious with 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. MATERIAL STUDIED. ARGENTINA. Tucuman, garden of the Instituto Miguel Lillo,

Singer T 108 and 108a (LIL). NORTH VIETNAM. Hanoi, Bon 4362, 31 May 1890 (FH type): Cai Kinh massive: Bau Mau (Tonkin), Bouitan 341b (FH, authentic).

13. Crinipellis missionensis Singer 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), concentrically zonate and at the extreme margin almost white-pilose with deeper colored hairs further inside, the hairs here more appressedly and wavy- tomentose but in the zone immediately surrounding the papilla less hairy and often almost naked, radially rugose in the intermediate zone, the papilla central, deepest colored and nearly or quite glabrous, convex, the marginal region becoming applanate. with at first involute or strongly incurved margin, reaching about 40 mm wlhen fully mature. Lamellae whitish 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 weak.

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 irregular in shape, mostly short and narrowly mucronate, a minority of cheilocystidia laterally or apically branched or 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, hymenophoral layer not gelatinized, regular, hyaline. Covering layers; 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 narrower and 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 subtropical forest, fruiting in spring (November, December)

MATERIAL STUDIED. Known only from the type locality: ARGENTINA. Misiones: Cataratas del Iguaz6 27 Nov 1949, Singer & Digilio M 123 (LIL).

This is easily the largest of the neotropical species of this section, and in some aspects reminds one of Crinipellis zonata (Peck) Patouillard, from which it differs in

spore characters. It is close to the African Crinipellis ghanaensis Singer in Pegler but differs from

the latter in having broader spores. Both the African and South American species differ from C. zonata in the inamyloid spores, and in the absence of a characteristic odor.

14. Crinipellis herrerae Singer, sp nov Fig 5. TYPE: Singer M 8125, from Veracruz, Mexico. Pileo ferrugineo-brunneo, KOH ope haud virescente nec grisente, in disco tomen-

toso, circum discum piloso pilis centrum versus depressis, marginem versus sparse piloso, sulcato, convexo, centro depresso, obtuso, 9-19 mm lato. Lamellis subdistantibus, sub- latis, subliberis. Stipite hyaline- vel pallide brunneo-piloso, subaequali 8-16 X 1-1.5 mm. Sporis (7-)9-10 X (4-)4.5-5.3A,, ellipsoideis vel oblongo-subfusoideis. Basidiis tetrasporis. Cystidiis nullis. Cheilocystidiis numerosis, dimorphis. Hypotrichio ex elementis latis ferrugineo-incrustatis efformato. Pilis haud multiseptatis neque acutis, crasse tunicatis; eis stipitis similibus. Ad ramulos exiguos in silva pluviali, typus in Mexico: Veracruz, Estacion Biologica de las Tuxtlas, Singer M 8125 (F).

Pileus rusty brown but rather deep colored and almost unchanged on drying, rather thinly beset with tough, long, brownish hairs which are denser around the flat disc where the hairs appear depressed inwards, on the small disc region few or no hairs visible but here the surface tomentose (whereas in the marginal zone the surface is glabrous among the sparse hairs), sulcate on the margin, smooth in the center, convex, soon with depressed center, neither umbilicate nor papillate or umbonate, 9-19 mm broad. Lamellae brown when collected, subdistant, not intervenose, medium broad to broad, with white edge, narrowly adnexed to subfree. Stipe with numerous hyaline 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, subequal or tapering downwards, 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 margin of pileus 13-17 X 6-9 p, clavate to subventricose and broad, much like the cheilocystidia of most Inocybes, brownish hyaline, smooth and entire, thin-walled, with an occasional hook, appendage or bulge at the apex, (2) over most of the edge 28-38 X 6.5-9 u, with several apical finger-like appendages, and because of these apically spreading to about 20 ,, main body cylindrical or subventricose to clavate, often with ramifications from the base or middle and thus assuming the shape of a carpophore of Ranzaria, the branches and appendages usually with rounded tips, rarely acute, sometimes secondarily forked or secondarily laterally or apically appendiculate. Hyphae of the trama brownish-stramineous-hyaline, inamyloid, with clamp connections, thin-walled, relatively broad but filamentous. Covering layers: hairs of the pileus mostly more than 200 p long, not densely crowded, in KOH light



24 Flora Neotropica

melleous to tawny, without ladder-like secondary 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 H thick, pseudoamyloid; hypotrichium consisting of repent broad (5-15 ,), rusty- incrusted hyphae with clamp connections; hairs of the stipe very similar to those of the margin of the pileus, stiff, more variable and 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 ,u thick, 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, apparently of dicotyledonous trees or vines, gregarious,

fruiting in the rainy season. MATERIAL STUDIED. MEXICO Veracruz: 7 km south of Montepio, Estacion Biologica

de las Tuxtlas, Herrera, 21 June 1969, comm. Singer M 8125 (F, type). The species is named for Dr. Teofilo Herrera, noted Mexican mycologist, collector

of the species.

15. Crinipellis septotricha Singer, Lilloa 8: 468. 1942. TYPE. Duss 513 (FH), from Guadeloupe. Pileus rusty brown when fresh, but soon fading to avellaneous brown, pale brown,

fulvous-buff, more or less strongly pilose, the hairs in curly strands radially appressed, merely tomentose or somewhat less thickly pilose on the papilla, convex, but often with a concentric wall or depression around the disc, more rarely alnost terraced, with a moderately prominent, rarely inconspicuous papilla visible in both fresh and dried condition and located inside an umbilicus, or rising without an umbilicus, eventually the whole middle zone flattened or depressed or at least becoming so in dried material, 3-16 mm broad mostly 6-12 mm. Lamellae pallid or white, then mostly becoming brownish pallid especially in the edge zone, medium broad, subventricose, close or subclose to medium close, not subdistant nor distant, emarginate-free or 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 depression or applanation, with Q > 2, rarely and rather exceptionally very slightly below 2, but often = 2, smooth, hyaline in KOH, with thin, eventually slightly thickened 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, narrowly subventricose, subclavate, the apex sometimes forked or more rarely subcapitate, with or without (more often without) 1-5 apical finger-like appendages, 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-like outgrowths all over their length, making the gill-edge heteromorphous. Hyphae of the trama hyaline thin-walled, inamyloid, with clamp connections. Covering layers: hairs of the pileus (middle zone outside the central papilla) mostly light melleous-fuscidulous, some hyaline in KOH, with a dense ladder-like structure from 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 narrowed towards 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 appendages at 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 ,u diameter, with 1.5-2.7 , thick wall.

KOH on the dried pileus darker brown, 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 nigricans.

MATERIAL STUDIED. BERMUDA. B & JDodge (NY paratypes). PORTO RICO. Bruce Fink (FH), probably belonging here. CUBA. Wright 45, 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) described originally a "forma umbilicata ad int" differing from the

typical papillate form in the absence of a papilla in the umbilicus or on the disc, growing on smaller sticks 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 extratropical South American species which differs however in subglabrous to glabrous disc on the pileus and broader spores. It is Crinipellis mlolfinoalna (Speg.) Sing. which was described by Spegazzini as Marasmius molfinoanus Speg., Bol. Acad. Nac. Ci. Cordoba 29: 123, 1926, from dead roots of Fagara coco. Only the type (leg. Mlolfino 212, 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 appendages or diverticula; the hairs are about 8 u broad and have the ladder-like structure of C. septotricha, with attenuate-obtuse to frequently needle-sharp acute tips, at least those at the margin of the pileus. It comes from the Sierra Grande, provincia de Cordoba, at Yaci, February 1912. This species, certainly different from C. septotricha, is mentioned here because its host extends north into the subtropical zone, together with the "Monte"

-vegetation characteristic for the Sierras de Cordoba. ILLUSTRATION. Dennis (1951a), p1 20, fig 6.

16. Crinipellis podocarpi Singer in Singer & Digilio, Lilloa 25: 225, 1952. Fig 6. TYPE. Singer T 833 (LIL), Argentina. Pileus in the broad marginal zone light brown to ochraceous-ferrugineous (reach-

ing "Arab" M&P fading to "terrapin" M&P), the entire disc region darkerchestnut to

fuliginous or the disc proper, a deep chestnut to ferruginous-fuliginous dot surrounded

by a narrow zone which is light rusty brown fresh and grayish pallid dried, and this zone separated from the marginal zone by a deeper colored (deep brown when dried) concentric line, the extreme margin often stramineous-pallid, hairy all over, but the hairs long, often fasciculate in the broad marginal zone and frequently projecting beyond the margin, whereas in the disc zone the hairs are short and the surface appears more tomentose to almost subglabrous, with some erect bristle-like hairs occasionally showing on the papilla, the latter at first very prominent and reaching 1 mm in height, later becoming inconspicuous and appearing like a tiny umbo under a lens, convex with

applanate zone around the papilla or slightly depressed in the center in adult material, the marginal zone strongly declivous for a long time and often radially sulcate, even-

tually more repand but with circular circumference, 4-13 mm broad. Lamellae white or whitish, broader towards the stipe but still narrow to 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 broadened at 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 suprahilar applanation or 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 cylindrical main body and with four to nine erect or obliquely erect apical finger-like appendages 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. Covering layers: hairs of the pileus (marginal zone) long-filamentous or gradually tapering to a narrow-obtuse or 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 sparse to numerous secondary septa (but these rather distant from each other, extraordinarily weakly 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 A thick walls, often with hyaline incrustation but not pigment-incrusted.

On living and dead trunks and branches of Podocarpus parlatorei, often ascending up to 3.5 m, gregarious, fruiting in the summer rainy season.

MATERIAL STUDIED. ARGENTINA. Jujuy: 15 km NE of Posta de Lozano, 20 Feb 1966, Singer T 5309 (F); Tucuman: Taficillo, 6 Jan 1950, Singer T 833 (UIL, types).

This species is often parasitic on Podocarplus in the subtropical-montane to tropical-montane zone of the Selva Tucumano-Boliviana. It is apparently closest to C. stip- paria.

17. Crinipellis stupparia (Berkeley & Curtis) Patouillard, Essai Tax. Hym. 143. 1900. Fig 7.

Agaricus stupparius Berkeley & Curtis, Jour. Linn. Soc. 10: 285, 1869. Collybia stupparia Berkeley & Curtis) Saccardo, Sylloge Fungorum 5: 217. 1887. TYPE. Wright (K, FH), from Cuba. Pileus ochraceous brown in the broad marginal zone, fading to near "thrush" M&P,

with blackish brown disc which is often surrounded by a blackish line, sulcate over about one third of the marginal radius, strongly and often coarsely pilose-floccose with mostly fasciculate and agglutinated hairs which form more or less curly-undulate strands but gradually less hairy and more tomentose and eventually glabrescent around 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 depressed around the usually prominent papilla, 1.5-7 mm broad. Lamellae white, in herbarium often becoming brownish-wood-color, medium broad to rather narrow, medium close to subdistant (12-20 lamellae and lamellulae) free to narrowly adnexed. 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 peripheric line 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, subventricose or subcylindrical main body and with four to twelve finger-



Crinipellis 27

like or forked apical appendages which are either erect or obliquely upright rarely distributed on 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, arising at the same level, rarely the main body sending out a long lateral filamentous branch, branches and appendages often secondarily diverticulate by short wart- or knot-like protuberances, rarely the whole cheilocystidium entire and smooth (in some preparations no 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 appendages or branches are spreading out above, not thick-walled, inamyloid. Hyphae of the trama hyaline, inamyloid, with clamp connections. Cover- ing layers: hairs of the pileus long and straight or flexuous, 4-11 broad, the walls melleous-brown or melleous-fuscous to (more rarely) subhyaline, strongly pseudoamyloid and cyanophilous, walls 1.5-4s, thick, tapering at 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; hypotrichium consisting of rather broad, cylindric, sometimes branching, almost thin- to moderately thickish- walled hyphal cells which are inamyloid and melleous to brown pigmented, the pipnent intraparietal but also here and there incrusting the walls.

KOH on the dried pileus negative. On sticks, branches, small twigs and fallen leaves of the evergreen tough type,

always on dicotyledonous never on living host tissue, growing usually gregariously in shaded ravines and cloud forest vegetation but also in subtropical and tropical forest of the plains from Cuba to South Brazil.

MATERIAL STUDIED. CUBA. Wright (FH, type); Wright 15 (part) (FH). VENEZUELA. Dto,Federal: Sierra Costera, El Junquito, + 1800 m alt, Deinnis 1800, 10 Jun 1958 (K). COL- UMBIA. Valle: Mun. Call: Cerro La Horqueta, + 2000 -m alt, 2 May 1968, R. Singer B 6903 (F). BRAZIL. Rio Grande do Sul: Sao Leopoldo, ex Herbario Theissen, Fungi Austroamericani, Rick 213 (FH).

The Brazilian collection has been determined by Theissen as Marasmius thwaitesii Berk. & Br., but it is not identical with that species.

18. Crinipellis coroicae Singer, sp nov Fig 8. TYPE. Singer B 558 (F), from Bolivia. Pileo cupreo-ferrugineo, in siccis colore C. stipitariae gaudente, piloso, centro

tomentoso-piloso vel fibrilloso-tomentoso, convexo, dein applanato, obtuso vel centro

depresso, 9-11 mm lato. Lamellis albis, 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. Basidiis tetrasporis. Cystidiis nullis. Cheilocysti- diis 16-28 X 5-7.5 j,, ventricosis vel clavatis et duobus vel six appendiculis apicalibus coronatis. Hyphis inamyloideis, fibulatis. Pilis in pileo obtusis vel acutis, in stipite frequenter acutissimis.

Pileus copper-rust color, soon with whitish portion on the marginal part, with

curly pilose covering evenly all over, or more tomentose-pilose to fibrillose-tomentose in the center, there also often darker colored at least in dried condition, dried fading to the dull tan-straw color 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 subumbonate convex, then

applanate or with depressed center, 9-11 mm broad. Lamellae white, close, medium broad to narrow, free. Stipe concolorous with the pileus but with white apex, pilose



28 Flora Neotropica

but with glabrous apex, insititious, equal, 23-32 X 0.5-0.7 mm. Context white, inodorous.

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-like appendages which 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, appendages 2-13 p long and some secondarily forked. Hyphae hyaline, more or less interwoven in the trama, with clamp connections, inamyloid. Covering layers: hairs of the pileus acute to broadly rounded, most frequently broadly rounded or attenuate-obtuse, stramineous to melleous-brownish in KOH, psuedoamyloid, 3.5-6.2 Ip

broad, with walls 1-2 1 thick, without ladder-structure; hypotrichium consisting of rather thick-walled, mostly not swollen elements which are stramineous-hyaline or 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. herrerae in relatively longer stipe and in the lack of

the rusty incrustation of the hypotrichiumn it differs from the C. patoitillardii of the South Pacific in the lack of glabrous zones in the less differentiated disc region of the pileus, the more appendiculate cheilocystidia, and in the habitat. It differs from C. stupparia in size of the carpophores and shape of the spores, and from C co/iiiiiuta in the absence of pleurocystidia.

19. Crinipellis tenuipilosa 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&P fascicles of hairs in the marginal zone, center more sparsely pilose, on black

ground visible among the hairs under a lens in dried condition, whereas the ground color appearing under the same circumstances among the hairs of the marginal zone is white, but without any naked zones, nor distinctly concentrically zonate, conic-applanate, with a prominent papilla, not umbilicate, about 17 mm broad. Lamellae white, subdistant, almost broad, free. Stipe brown below, concolorous with the pileus in the middle portion, and white at the apex, flocculose-pilose below, appressedlv hairy in the middle portion, and subglabrous at the apex, subequal, or slightly tapering downwards, 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 narrow as 8.3 X 2.7 (Q=3), oblong or sub-fusoid, smooth, hyaline, inamyloid, thin- walled, some over-aged ones 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 several apical cylindrical appendages of 5.5-8.5 X 2 p1 over a clavate main body, entirely hyaline. Hyphae with clamp connections, inamyloid. Covering layers: hairs of the pileus agglutinated in 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 branches of dicotyledonous trees in sub-tropical forest, solitary, fruiting in the rainy season.

MATERIAL STUDIED. Only the type collection is known. ARGENTINA. Tucuman: Quebrada de Lules, 24 Jan 1951, Singer T 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 larger size and somewhat larger spores.

20. Crinipellis mexicana Singer, sp nov Fig 9. TYPES. Murrill 142 (NY), from Mexico. Pileo papilla penicillata gaudente, 2-5 mm lato. Lamellis cremeo-albis, subconfertis

vel mediocriter confertis. Stipite atrobrunneo in vegetis, centrali, curvato, 9-14 X 0.3-1 mm. Sporis 7-10 X 3-6.5 p; cystidiis raris, cheilocystidiis simillimis; cheilocystidiis integris, interdum ad apicem bifurcatis, haud multi-appendiculatis. Pilis pilei aut rotundato-obtusis aut 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, reaching a dull stramineous-tan, strongly pilose, radially curly-rivulose in marginal portion (about two thirds of radius), radially appressedly fibrillose and much less hairy in a disc zone but often with a brush-like tuft of bristle- like hairs immediately around or more often on the top of the central papilla, the whole disc region occasionally glabrescent, convex, then applanate around the umbo or papilla, later more repand all over with the zone around the umbo or papilla becoming depressed and 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. Lamellae cream 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 sparsely and weakly pilose all over, mostly more strongly hirsute-pilose below and weakly pilose and glabrescent above, 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 variable in size and shape but much more frequently with Q< 2 than with Q = 2 or > 2, hyaline, smooth, inamyloid, thin-walled, when over-aged more 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 heteromorphous or almost so, 10-50 X 5.5-7 p, mostly 20-28 X 5.5-6 p, characteristically entire and not apically appendiculate or only rarely with a forked apex or with tripartite (three finger-like appendages) apex, clavate to ventricose or subcylindrical, more rarely ampullaceous, but often constricted in the middle, rarely with an occasional lateral diverticulum or apically crooked, hyaline. Hyphae of the regular hymenophoral trama hyaline, inamyloid, with clamp connections, interwoven. Covering layers: hairs of the pileus long, 4-7.8 p broad, melleous to rusty brownish in KOH, some subhyaline, some flexuous, some with rather numerous thin-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-obtuse or rounded than acute.



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 margin of 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, 12-20 Dec1909, W. A. & Edna L.

Murrill 142 NY, type), 116 (FH, paratype). BOLIVIA. La Paz: Nor-Yungas: Capillaria 22 Feb 1956, Singer B 1323 (F, paratype).

Singer (1942) originally believed this species to be the same as C patouillardii Sing. which, in a restricted sense, i e excluding the neotropical material, occurs only in the South Pacific and grows on leaves, petioles etc. of Dicotyledones. The carpophores in that species are slightly larger in an average, the pleurocystidia slightly 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. Crinipellis foliicola Singer, Mycologia 47: 771. 1955. Fig 10. TYPE. Singer F 444a (F), from Florida. Pileus "Mummy Brown" or between "Mummy Brown" and "Prout's Brown"

Ridgway, fading to "Dresden Brown" and "Cinnamon Brown", sometimes more rusty brown when fresh and then becoming sordid stramineous when dried, the disc often darker but not blackish, wavy-appressedly pilose, but the disc-zone less hairy and appearing merely tomentose, more rarely evenly hairy all over, if tomentose appearing more

pallid-canescent when dried, except for the darker small umbo or papilla, in the marginal zone the white ground showing through the strands of hairs in dried material, one or two concentric furrows developing in dried material, and sometimes also visible in fresh material, convex-campanulate or convex, then more or less applanate with a flattened or umbilicate depressed zone around the umbo or papilla, the marginal zone often somewhat coarsely sulcate and occupying about two thirds of the radius or somewhat more, 5-7.5 mm broad. Lamellae white to cream color, close to medium close, with entire and at first always pure white edge, narrow to broad (up to 1.5 mm

broad), rounded free or attenuate free. Stipe concolorous with or slightly paler than the marginal portion 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-mature ones

usually thick-walled and 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 X 3.5-8(-8.5)p, obconic or clavate, cylindrical or ventricose, sometimes forked or with three branches, with, rarely without, finger- or sterigma-like apical appendages or hooks (1-7) which may again be somewhat diverticulate or forked, 1.5-6 p long, with rather thin wall, hyaline to yellow or melleous-hyaline, apparently at first all hyaline, making the edge heteromorphous. Hyphae hyaline, inamyloid, with

clamp connections; hymenophoral trama regular. Covering layers: hairs of the pileus in the marginal zone mostly long (about 350 ,), in the central zone often shorter, the latter more often rounded at the tip, the former variable and both rounded to attenuate-obtuse as well as acute tips observed, all rather pale melleous or more rarely sub-



Crinipellis 31

hyaline to hyaline in the apical portion, and there often with ladder-like or at least dense secondary septa, in the lower portion melleous to melleous-umber in 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; hypotrichium hyaline, pigment-less, more rarely pale melleous, multiseptate with rather broad (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 1 broad, with walls 2-3 p thick, tips rarely rounded, most frequently acute, with or without ladder-structure.

KOH on dried pileus deeper rusty brown, not greening or graying. On fallen leaves, rarely passing over to small dead woody twigs, in subtropical

and tropical-montane forests, gardens and plantations, always on Dicotyledones, solitary or gregarious. Known hosts: Smilax sp., Quercus sp., Coffea arabica.

MATERIAL STUDIED. U.S.A. Florida: Highlands Co., Highlands Hammock State Park, 2 Sep 1942, Singer F 444a (F, type), 30 Aug 1942, Singer F 444 F. paratype). MEXICO. Oaxaca: Sierra Mazateca: between Huautla de Jimenez and Teotitlan del Camino, km 20-21, 11 Jul 1969, Singer Al 8444 (F); Rancho del Cura 7 Jul 1969, Singer M 8288 (F); Road to Huautla de Jimenez, Cerro Verde, 6 Jul 1969, Singer M 8274 (F); Chiapas, 15 km W of San Cristobal de las Casas, 2400 m alt, 2 Aug 1969, Singer M 8820 (F). VENEZUELA. Dto.Federal: Caracas Botanical Garden 19 Jun 1958, Dennis 1071A (K).

This is very close to C. phyllophila and C. dipterocarpi, the latter from Asia, the former from Mexico. Crinipellis phyllophila differs in slightly larger carpophores, slightly broader spores, 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-spored typical form by transitions - are often difficult to distinguish from 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 somewhat incomplete with regard to the anatomy of the covering layers, and therefore, for the time being, I describe the two as different species.

22. Crinipellis phyllophila Singer, Sydowia 11: 358. 1958. Fig 11. TYPE. Singer M 1507 (MICH), from Mexico. Pileus in central ring-wall and on the ribs between the sulcations deep ferruginous,

rich brown, but soon fading to cinnamon tan, paler on the margin because the pale tan to whitish ground color between the more scattered hairs shows through, concolorous with the margin or with the ring-wall in the disc-zone which is neither glabrous nor shining, but likewise pilose, the central disc either flat or more rarely with a central papilla, a circular wall appearing around the disc-zone, and outside the wall surface coarsely sulcate over one third or one half of the radius, convex, very soon more applanate or even depressed in the disc-zone, later flat-convex to applanate over all, 6-11 mm broad. Lamellae pure white, not cream color when fresh, close to rather close, moderately broad or rather broad, free. Spore print pure white. Stipe in upper portion 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, somewhat longer than twice the breadth, unicellular even if overmature, hyaline, smooth, inamyloid, thin-walled, slightly thick-walled when over-mature. Hymenium: basidia 21-30 X 4.5-6 p, 4-spored; cystidia none (excepting some occasional cheilocystidia very close to the edge proper); cheilocystidia 21-30 X 4.5-6 P crowned by two to six erect appendages 2-5.5 X 1-1.2 ,, main body cla-



32 Flora Neotropica

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

clamp connections, not gelatinizing. Covering layers: hairs of the pileus (marginal zone) brown or melleous brownish, with the apical portion usually hyaline or subhyaline, filamentous but often somewhat flexuous, with needle-sharp or 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 secondary septa, pseudoamyloid.

KOH on the dried pileus not green or gray. On fallen leaves of evergreen Quercus in a remainder of original tropical-montane

mixed oak woods rotting in a thick layer of leafmold, fruiting in summer, gregarious. MATERIAL STUDIED. Known only from the type locality; MEXICO. Oaxaca: Huautla

de Jimenez, below the Rancho del Cura, about 1500 malt, 10 Jul 1957 Singer 3 1507 MICH, type), 12 Jul 1957, SingerM 1528 (MICH, paratype).

23. Crinipellis dicotyledonum Singer in Singer & Digilio, Lilloa 25: 221. 1952. TYPE. Singer T 1140 (LIL), from Argentina. Pileus ferruginous "Alamo" M&P, in the marginal zone partly lighter colored, evenly

pilose all over, eventually glabrescent especially in the marginal zone, convex and papillate, eventually becoming convex with depressed center and a papilla in the depression, about 18 mm broad. Lamellae not white but buff (10 C 6,M&P), subclose, medium broad, emar-

ginate-free. Stipe concolorous with the pileus, pilose, gradually and slightly attenuated towards the abruptly but slightly widened base which is insititious, about 19 mm long and 1.5 mm broad. Context white, thin, with an odor ofMicroniphale ofetidlum or of sauerkraut.

Spores 8.5-9.6 X 5.5-6.2 ,, ellipsoid, hyaline, inamyloid, thin-walled, the overmature ones mostly thick-walled, smooth. Hymenium: basidia 33-36 X 6.8-8.2 p, 4-spored; cystidia none (excepting some cheilocystidia off the edge proper); cheilocystidia densely crowded on the heteromorphous edge of the lamellae, (20-)27-44(-58) X 5.5-7.5 , either

simple and entire and then cylindric-fusoid, or subulate or ramified in the basal or middle

portion of the cell, and then the branches 4.8-5.5 pi broad, sometimes the main body forked or tripartite or somewhat nodose-uneven all over, with obtuse tips, rarely subacute

slightly opaque, constantly hyaline. Hyphae in the regular hymenophoral trama hyaline, with clamp connections, inamyloid. Covering layers: hairs of the pileus melleous-stramineous in KOH, pseudoamyloid, smooth, 5.5-7 p broad, very slightly and gradually attenuate towards the apex, with walls 1.3-2.5 uthick, with rounded tip, without secondary septa or with few distant ones.

On dead branches and small sticks of dicotyledonous woody plants in subtropical forest, gregarious, fruiting in the summer rainy season.

MATERIAL STUDIED. ARGENTINA. Tucuman: Rio de los Sosas 27 Jan 1951, Singer T 1140

(LIL, type).

Crinipellis sect Crinipellis subsect Heteromorphinae Singer, Lilloa 8: 509. 1942. Characters like those of subsection Stipitarinae, but pleurocystidia differentiated. TYPE SPECIES. Crinipellis minutula (Hennings) Patouillard.

Key to the Species of Subsection Heteromophinae

1. Spores strongly elongated, 9-17.5(-20) X 3.5-6.3st (see subsection Stipitarinae). 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 var junia.



Crinipellis 33

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

24a. Crinipellis commixta 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 dehydration soon 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 strands of appressed curly hairs of the wide marginal zone, and likewise appressedly tomentose-pilose on 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 rather obtuse papilla or small umbo, later depressed around the latter, and eventually applanate to concave retaining the papilla which becomes more evident when dried, 4-11 mm broad. Lamellae pure 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 appressedly tomentose-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, rarely ventricose, 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-like branches or apical appendages, some branching off at the middle, more at the apex of the main body and of very variable size and shape, hyaline to pale stramineous, some somewhat opaque because of slightly thickened wall, numerous and crowded, making the edge heteromorphous or almost so. Hyphae hyaline, with clamp connections, mainly filamentous, inamyloid. Hymenophoral trama regular. Covering layers: hairs of the pileus in the wide marginal zone long (450 s and longer), melleous-brownish in KOH, some hyaline, all pseudoamyloid, 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 rather numerous 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 gradually tapering to an acute, more rarely obtuse or rounded tip, with 1.5-2.2 p thick wall.

KOH on pileus not provoking a green or gray color reaction. On fallen woody twigs and branches and on roots, always on Dicotyledones,

mostly gregarious, fruiting in the rainy seasons (summer or winter). Known hosts: Acacia sp, Prosopis torquata, Dwialia breviflora.

MATERIAL STUDIED. ARGENTINA. Tucuman: San Pedro de 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), probably belonging here; 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. Crinipellis commixta var junia Singer, var nov TYPE. Singer T 2791, from Argentina.



34 Flora Neotropica

A varietate comiimixta differt sporis angustioribus. Typus varietatis a R. Singer Junio mense in Argentina, Tucuman, lectus et in LIL conservatus.

Pileus as in var comitixnta. Lamellae as 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, rather scattered; cheilocystidia as pleurocystidia or more frequently with one apical appendage or bifurcate at the apex, sometimes with irregularly located and sized branchlets, often ventricose in the middle or constricted. Hyphae and covering 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 subxerophytic vegetation elements, solitary, or in small groups.

MATERIAL STUDIED. Known only from the type locality. ARGENTINA. Tucuman: Aroyo Mista, 15 Jun 1955, Singer T 2791 (LIL, type).

In the type variety, I include a winter-fruiting form which may well be another distinguishable taxon, possibly on the subspecific level, in which the spores are partly pseudoamyloid when freshly dried, but relatively broad as in the summer-fruiting forms. The narrow-spored winter-fruiting form which is here described as var junitia differs insofar 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 considerably lower 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 necessary to gather more collections of the winter-fruiting forms so that the constancy of the diagnostic characters can be re-checked.

Crinipellis colninixta var junia may be considered a form intermediate between C. commixta and C n2inutula. It would seem that it is closer to C. conmmixta var commixta.

C. minutula (Henn.) Pat. (Len tinus minuutulus Henn.) is still smaller than C. conmmixta var junia and has still narrower spores, more frequently a more typical ladder structure and more often acute tips in the hairs of the pileus. It is a tropical African

species.

Crinipellis sect Excentricinae (Singer) Singer, stat nov Crinipellis sect Eu-Crinipellis subsect Excentricinae Singer, Lilloa 8: 507. 1942. Crinipellis sect Crinipellis subsect Excentricae [sic] Singer, Agaricales in modern Taxonomy,

2nd e. p 361. 1962. TYPE SPECIES. Crinipellis excentrica (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. Cheilocystidia either ventricose below with long and

relatively thin apical neck, or strongly apically branched-setulose. Surface of the pileus not greening or graying with KOH 5%;



Crinipellis 35

Key to the Species of Section Excentricinae

1. Pileus 7-12 mm broad, with an abrupt papilla, lamellae close or subclose. 25. C. excentrica. 1. Pileus 3-7 mm broad, without a distinct papilla, lamellae distant or subdistant.

2. Cheilocystidia ventricose with wart-like or short rodshaped appendages or setulae in the upper third. Spores 6-8.5 X 5.3-7.71; stipe appressedly villous-tomentose; pileus reniform. On myrtaceous branchlets. 26. C. myrti.

2. Cheilocystidia ventricose below, with a rather narrow and rather long apical neck and thus ampullaceous, the neck often sparsely diverticulate. Spores 8.2-10.5 X 4.5-5.5,; stipe with rather scattered erect hairs, hirsute; pileus circular. Not on Myrtaceae.

27. C. albipes.

25. Crinipellis excentrica (Patouillard & Gaillard) Patouillard, Jour. de Bot. 3: 336. 1889. Fig 13.

Collybia excentrica Patouillard & Gaillard, Bull. Soc. Mycol. France 4: 15. 1888. TYPE.. Gaillard 128 (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 marginal zone, silky-pilose to radially appressed strongly hairy in the wide marginal zone as well as in the center, but on the latter thinly silky and subglabrous or glabrescent, in the marginal region of the primordia with fluffy-woolly, not radially depressed hairs, in age sulcate over one quarter to three quarters of the radius, convex and strongly papillate when young, even later not depressed excepting in dried material, and even then maintaining a distinct, abrupt papilla, with reniform outline, not or rarely circular, 7-12 mm broad. Lamellae white, 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, insititious, solid, curved and eccentric in adult caps, equal or tapering up- 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 sterigmatoid outgrowths at the

apex, opaque and somewhat thick-walled, projecting slightly beyond the basidia, fusoid, clavate, or otherwise elongated, hyaline, numerous; cheilocystidia 19-31 X 3.5-8.5 A, the branches often spreading to up to 12 ,, most of them apical, some mediane,

with entire, bifurcate or sterigmatoid-setulose apex and the setulae sometimes second-

arily knotty-diverticulate, the branches also 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, appendages and branches up to 12 X 3 1 but usually much shorter and narrower.

Hyphae with clamp connections, inamyloid. Covering layers: hairs of the pileus hyaline, 2-5 p broad, with 1-2.5 u thick wall, with mostly rounded tip but some hairs with a thin-walled, inamyloid or scarcely pseudoamyloid appendage and showing up to 4

clampless cross-walls there, the appendage obtuse or acute, below the appendages (if these are present) pseudoamyloid in young specimen but in older ones gradually more and more weakly or moderately pseudoamyloid, sometimes wall appearing to thicken in 5% KOH.

KOH on dried pileus negative (no color change). On Bambuseae (and possibly other plants), dead branches, gregarious. MATERIAL STUDIED. VENEZUELA. Atures (Orinoco region), A. Gaillard 128 (FH,

type). BOLIVIA. Pando: Madre de Dios, Rfo Madre de Dios, near Santa Teresa, 28 May 1956 Singer B 2295 (F).

ILLUSTRATION. Patouillard (1900), fig 70, Singer (1942), fig 1 f, fig 3 d.



36 Flora Neotropica

26. Crinipellis myrti Patouillard in Patouillard & Lagerheim, Bull. Soc. Mycol. France 9: 125. 1893.

Marasmius myrti (Patouillard in Pat. & Lag. Saccardo, Sylloge Fungorum 11: 37. 1895. TYPE. Lagerhleim (FH), from Ecuador. Pileus pallid to pale brownish in dried condition, with appressed hairy squamules,

somewhat striate near margin, convex with incurved margin which is more villous than squamulose, with depressed center when adult, (dried), but apparently at first convex. 3-6 mm broad, in dried condition. Lamellae white or whitish, distant, intermixed, medium broad or rather narrow, almost free. Stipe dark chestnut brown when dried. with appressed silky 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-walled and 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. Covering layers: hairs of the pileus hyaline, the thinner ones often brownish, 4-8 , broad, with obtusely rounded, rarely acute tip, many with distinct ladder structure because of numerous and moderately crowded secondary septa. only slightly pseudoamyloid.

On fallen branches and twigs of Mylrtus sp. MATRRIAL STUDIED. Known only from the type locality. ECUADOR: Rfo

Pululahua, Lagerheim (FH, type).

27. Crinipellis albipes Singer, sp nov Fig 14. Types. Singer M1 8083 (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-ochraceous dot above the attachment of the stipe, finely and loosely wooly-tomentose all over, when wet shallowly sulculate over three quarters of the radius, dried smooth or nearly so, convex, eventually with depressed center, obtuse 6-7 mm broad. Lamellae white, subdistant, narrow, intermixed didymous, with 12-13 through-lamellae, not intervenose, adnexed to rarely subfree. Stipe white, finely long-pubescent from hyaline hairs from top to base,solid, curved, subequal, slightly eccentric, more rarely nearly central, central, when quite young, insititious, 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 suprahilar depression, rarely slightly bean-shaped in lateral view, 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, straight or flexuous, eccentrically inserted or central-apical (more commonly so), smooth or with scattered lateral or apical diverticula, sometimes at the apex bifurcate, hyaline, with rounded but sometimes narrowed 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-walled near subhymenium which appears to be subcellular. Hymenophoral trama regular. Covering layers: 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-obtuse tip (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. MATERIAL STUDIED. known only from the type locality. MEXICO. Veracruz:

Near Montepfo, Estacion Biologica de las Tuxtlas, 20 Jun 1969, Singer M 8083 (F, type).

Crinipellis sect Grisentinae (Singer) Singer, stat nov Crinipellis sect Eu-Crinipellis subsect Grisentinae Singer, Lilloa 8: 497. 1942. TYPE. species. Crinipellis mirabilis Singer. 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. Fresh pileus pink, red or purple; on dicotyledonous wood or leaves. 2. Spores 4-5 broad; stipe central.

3. Cheilocystidia crowned with several apical appendages; on wood. 33. C. rubida. 3. Cheilocystidia with few if any appendages; on leaves. 32. C. dusenii.

2. Spores 4.8-6.8u broad: stipe eccentric and not longer than the diameter of the pileus. 34. C austrorubida.

1. Fresh Pileus neither pink nor purple, but chestnut-brown to straw-ocher, rusty brown to cinnamon-tan; pleurocystidia moderately to quite numerous. 4. On wood or fallen leaves of Iresina, Solanum, Boehrneria, Metahybe and other dicoty-

ledonous vines, shrubs or trees. 5. Spores oblong, 3-3.5p broad: cheilocystidia crowned by numerous apical appen-

dages. 30. C. sapindacearum. 5. Spores ellipsoid, 4-6, broad, cheilocystidia as above but also many (mostly the

majority) entire or simply forked or with scattered branches. 31. C. tucumanensis. 4. On grass roots, grass leaves, culms, or on other Monocotyledones, particularly Bam-

buseae. 6. Pileus umbilicate, 6-12 mm broad; cheilocystidia entire or with rarely more than

three apical appendages or setulae; hairs partly deep green incrusted in KOH mounts. 28. C. alcalivirens.

6. Pileus papillate, the papilla often with an apical brush-like strand of hairs; 2-7 mm broad; cheilocystidia in their majority with several apical appendages; hairs tending to be greenish or greenish melleous in KOH mounts but not conspicuously incrusted. 29. C. trichialis.

28. Crinipellis alcalivirens Singer, sp nov Fig 15. TYPE. Singer B 3510 (F) from Colombia. Pileo ferrugineo-vel castaneo-brunneo, desiccatione sordide corii-colori et sub-

pallide crinito, 6-12 mm lato. Lamellis candidis, 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. Cheilocystidiis versiformibus. Pilis pilei Na OK vel KOH ope virescentribus, praesertim incrustationis viridis causa. Ad Bambuseas in Columbia, Singer B 3510 (F), typus.

Pileus rusty to chestnut brown ("Arab" to "Mohawk" M&P), drying to dull tan or a brown overlaid by almost pallid or quite pallid strands of 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-ascendant hairs, the large marginal zone with more or less appressed, and the extreme margin with fimbriate-projecting hairs, adult caps, convex, later with a shallow and small central depression, 6-12 mm broad. Lamellae pure white, narrow or rather narrow, close, mostly free, more rarely very narrowly adnexed. 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 remains pallid 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, narrowly ventricose, 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, similar to the pleurocystidia but more versiform, numerous and crowded, often more clavate, often apically forked, or with 1-3 large apical appendages, these, if present, irregularly shaped, obtuse or acute, thin to moderately thickish walled. Hyphae of the trama hyaline not gelatinized, inamyloid, with clamp connections. Hymenophoral trama regular, of mostly filamentous hyphae, Covering layers: hairs of the pileus long, 4-5 1 broad, light melleous brownish, but turning green in the pigmented portion from an incrusting mass of pigment which is deep green in alkali (NaOH, KOH, 5 or 10%), with 1.5-2.7 p thick walls, with the apex thinner-walled in 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 downwards below the first clamped septum, with thinner (0.2-0.4 p) wall and gradually narrowing downwards, generally subhyaline, not dark incrusted from pigment unless the latter is greenish in KOH; hairs of the stipe 7-8.8 , broad, hyaline, subacute, more rarely rounded at the tip, walls 2.5-3.5 p thick, generally without a typical ladder structure and 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 Bambuseae in wet locality. MATERIAL STUDIED.Known only from the type locality. COLOMBIA: Cundinamarca:

Salto de Tequendama, 24 Jul 1960, Singer B 3510 (F type).

29. Crinipellis trichialis(Leveill) Patouillard, Bull. Soc. Mycol. France 24: 8. 1908. Fig 16.

Agaricus trichialis Leveille, Ann. Sci. Nat. III, 5: 113. 1846. Naucoria ? trichialis (L6veille) Saccardo, Sylloge Fungorum 5: 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 darker and reaching fuliginous brown in the disc area, in the marginal half eventually some white ground color showing through the strands of hairs, occasionally the hairs of the marginal zone whitish also, with entire or fimbriate margin (from projecting hairs), in the center more tomentose or short-pilose than in the wide marginal region, and eventually glabrescent, but on the top of the central papilla often or always with a penicillate fascicle of bristle-like agglutinations of long hairs, more or less sulcate on the margin, campanulate-convex or 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 subascendant, soon horizontal, free. Stipe"russet brown," 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 suprahilar applanation, thin- then thick-walled (0.7 s) and if over-aged often developing a 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 amorphous incrustation; others are cystidiole-like and resemble larger basidioles, these often mucronate or with 1-2 apical prongs, hyaline, with mostly thickish wall, rarely solid. Cheilocystidia numerous but often with mixed-in basidioles or basidia, 13-48 X 5.5-8.3 ,, varying from being shaped like the pleurocystidia to more branched or appendiculate and then ventricose to clavate with one to several apical, finger-like appendages, others with lateral branchlets 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 subhyaline or hyaline, the pigmented ones often (but not all of them) turning olive 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-structure from 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-walled and less strongly pseudoamyloid, more often obtuse. Hairs of the stipe similar.

KOH on the surface of the dried pileus weakly or partially greenish, but not constantly so, sometimes merely deeper brown, not black-brown, nor distinctly green or gray.

On dead Monocotyledones, particularly on culms of Bambuseae (Chusquea, Anrn- dinaria), and on dead grass culms in subtropical and tropical forests. Gregarious fruiting in the rainy seasons.

MATERIAL STUDIED. VENEZUELA. Distrito Federal: Botanic Garden of Caracas, 19 Jun 1958, Deinnis 1071 (K, F). BRAZIL. SSo Paulo: Cipo, 1 Sep 1964, Skvortzov & Alim 71393, (NY). ARGENTINA. Tucuman; Las Lenguas, 20 Feb 1951, SingerT 1310 (LIL), Singer T 1291 (LIL), Sin- ger T 1291a (LIL). INDONESIA. Java: Hohnel (FH), Zollinger 2078 as "A. trichophorus" (FH, type).

The cystidia of this species are characteristic and generally numerous. In some specimens true cystidia are formed, whereas in another collection a second, cystidiole- like, type of pleurocystidium may be predominant. This species is closely related to the preceding one and should not be confused with such small species of section Crinipellis as C. perpusilla, C. stupparia, or C. mexicata or C. commixta.

30. Crinipellis sapindacearum Singer, sp nov TYPE. Brazil: Singer B 3377 (F). Pileo brunneolo, KOH ope nigricante, piloso, umbonato vel papillato, 3-5 mm lato.



40 Flora Neotropica

Lamellis confertis. Stipite 9-14 X 0.4-0.5 mm. Sporis 7.5-8.2 X 3-3.5 , cystidiisnumero- sis. Cheilocystidiis apicaliter appendiculatis. Hypotrichio viridi-incrustato. Ad folia coriacea 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-pilose with the papilla subglabrous or glabrescent, with fimbriate-ciliate margin, without a setiform fascicle of penicellate bristle-like agglutinated hairs on the papilla, convex, with rather low papilla, 3-5 mm broad. Lamellae whitish or pallid, becoming brown in the herbarium, close, moderately broad to narrow, subfree. Stipe concolorous with the pileus, with abundant curly hairs

projecting up to 0.5 mm, insititious, approximately three 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, inamyloid; cheilocystidia making the edge heteromorphous since they are crowded on the

edge and differentiated from the pleurocystidia, basidiomorphous but with numerous

apical erect-subdivergent obtuse appendages 2.7-7 X 1.2-2.5 p, the entire cell with the

appendages included (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. Covering layers: hairs of the pileus long, 3-7.5 u broad, hyaline to light brownish, with rounded or attenuate-obtuse tips, with walls 1-3.5 u thick, pseudoamyloid, the hairs of the middle of the broad marginal zone and those from the ciliate fibrils of the margin not differentiated from each other; hypotrichium characteristically green incrusted.

KOH on the dried pileus provoking a blackish reaction but with a greenish-melleous or greenish background (from the hypotrichium.)

On dead fallen coriaceous leaves of Sapindaceae and perhaps other dicotyledonous families in low tropical rain forest, gregarious. Known host: MetahYlbe sp.

MATERIAL STUDIED. Known from the type locality. BRAZIL. Paraibo: Jo5o Pessoa, 11 Jul 1960, Singer B 3377 (F, type).

This species is close to the South Pacific C. mirabilis Singer which has somewhat larger spores, more abnormal hairs, on the pileus, more conical pileus and grows on dead twigs. Crinipellis sapindacearunm should not be confused with C. foliicola.

31. Crinipellis tucumanensis Singer, sp nov TYPE. Singer T 2158, from Argentina. Characteribus C. patouillardii, C. commixtae, C. corticalis gaudens sed KOH op6

oliveoviridans: apice stipitis castaneo-rubido colore insignis; ad ramos et caules. Typus in LIL conservatur.

Pileus fulvous-brown between "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 reaching as deep (but not as reddish) as "Mandalay" (M&P) and fading there to "butterscotch," the primordia still entirely color of the adult disc-region, densely and strongly pilose all over excepting the zone on the sides of the papilla and around the papilla which is generally much less strongly hairy and tends to become glabrescent, often with striate-sulcate margin, 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 material the papilla less striking, and in old material often so indistinct that it can scarcely be distinguished, with long, agglutinated bristle-like erect 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. Lamellae white, not cream, remaining whitish in age, intermixed-tridymous, almost narrow to (more frequently) broad, 1.2-1.8 mm broad, close to subdistant, mostly subclose, mostly slightly ventricose, free, more rarely very narrowly adnexed. Stipe at first slightly paler than the pileus, later "kis kilim" to "cocoa" or "leaf mold" or 15 E/H 12, M&P, but with a particularly bright reddish chestnut 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 appressed rather dense hairs over most of the surface, but towards the base often with bristle-like hirsute 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-aged thick-walled, and averaging 8.2 X 5.5 A1, inamyloid, smooth. Hymenium: basidia 26-30 X 6.7-8.8 ,, 4-spored; cystidia moderately numerous, basidiomorphous, well differentiated from the basidia in phloxine-KOH preparations, 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-subcapitate or subclavate, very rarely crowned with more than two appendages, but more frequently subvesiculose below with a mucro, mostly somewhat opaque, but not thick-walled, sometimes many or all with a resinous melleous-hyaline incrustation or hyaline incrusted at the apex only. Hyphae of the trama hyaline, with clamp connections, inamyloid. Covering layers: hairs of the pileus between center and extreme margin long and brown, a few among them subhyaline to hyaline, in 5% KOH turning greenish or greenish melleous, 4-6 u broad, with acute, even needle-sharp tip 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 approaching a 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 subtropical forests and in shady wet ravines, fruiting in the summer rainy season (December until February). Known hosts: Iresina sp (Amaran- thaceae); Boehmeeria caudata (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 distinguished from C. patouillardii and C. conmmrita. However, the KOH reaction in C. tucumanensis is characteristic and the bright color at the apex of the stipe has not been observed in C. commixta.

The European Crinipellis corticalis (Desmazieres) Singer & Clemen9on is closely related and differs from C. tucumanensis Singer in darker KOH reaction, somewhat larger spores, more scattered fusoid-subampullaceous cystidia, and the host range (Caprifoliaceae and Oleaceae).

32. Crinipellis dusenii (P. Hennings) Singer in Pegler, Kew Bull. 21: 518. 1968. Fig 17.

Marasmnius dusenii 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 marginal zone, the browning starting at the center, finely pilose-fuzzy there, with a more appressedly pilose zone around a subglabrous central dot, the margin somewhat more pilose than the center, not zonate when fresh but slightly so when dried, sulcate, campanulate, then convex, often either umbonate or depressed in the center, 3-18 mm, mostly about 10 mm broad. Lamellae white, narrow, more rarely medium broad, intermixed with lamellulae, round- free or attenuate-subfree. Stipe umber or brown ("Wood Brown" Ridgway according to Pegler), but sometimes also purple or chestnut when fresh, pilose-hirsute, equal, insititious, 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 rather numerous, (15-)27-37(-39) X (5.5-)6.5-9.5(13.5) ,, clavate, sometimes with one or two appendages or 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, basidioles and 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. Covering layers: 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 subacute tip, smooth or vaguely incrusted in places, numerous, simple, rarely with a secondary septum in the upper portion of the hair, rarely laterally branched, no ladder-like septation. Similar hairs on the surface of the stipe.

KOH on pileus (dried) glaucous to greenish gray, then green-black. On fallen leaves of dicotyledonous trees, in Africa and South America. MATERIAL STUDIED. ECUADOR. Tungurahua, Rio Verde, 28 Apr 1973, Singer B 7170

(F). GHANA. Tofo, 8 Apr 1955, Holden GC 35 (K). ZAIRE. Binga, 16 May 1927, Goossens- Fontana 605 (BR); Yangambi, 18 May 1939, Louis 14862 (BR).

ILLUSTRATION. Pegler (1968), fig 6, la-lf.

33. Crinipellis rubida 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 radial ribs strigose clothed, almost squamulose, with incised-denticulate margin, convex, then applanate or depressed, with a slight papilla in the center, reaching 20 mm broad. Lamellae white, 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-aged ones thick-walled and 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-like appendages in the apical region, these erect or somewhat oblique. Hyphae inamyloid. Covering layers: hairs of the pileus (marginal zone) long, in bunches of subparallel hairs running radially on 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 Trinidad but, as mentioned before (Singer, 1955, p 396)this is a different species whichl has been named C trinitatis Dennis. This species, unknown to me, appears to belong in the section Iopodinae rather than the Grisentinae, and Dennis's description is repeated there.

34. Crinipellis austrorubida Singer, sp nov TYPE. Sinlger T 917, from Argentina. Pileo purpureo-rubro; crinibus epicuticularibus KOH ope olivacentibus. Typus in

LIL. Pileus purple red (between "canyon" and "rose ebony" M&P), deeper or paler

colored according to the density of the fibrils and hairs of the pileus), deeper colored toward the more densely pilose center, convex, 5-6 mm broad. Lamellae white but the interlamellar spaces pale purplish, distant (about 10 through-lamellae and 10 lamellulae), rather narrow, adnate. Stipe concolorous with the pileus below but white above, fibrillose-pilose below, glabrous at 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 narrowly cylindrical and capitate, etc., not making the edges heteromorphous in all parts. Hyphae inamyloid, not gelatinized, hyaline, filamentous, with clamp connections (so in pileus trama and in the regular hymenophoral trama). Cortical layer: epicutis of the pileus consisting of hairs which are mostly strongly pseudoamyloid, few weakly so, with rounded tips, some with an apical callus, with a partial pigment incrustation which is lilac in ammonia but becomes olive green in KOH, with thick (1-2.5 p) wall, with a diameter of about 3-8.5 ,; hlypotrichium of similarly incrusted, repent, relatively broad hyphae with numerous clamped septa, 4-16.5 p in diameter.

On fallen twigs in subtropical forest, 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 preceding species because of the

greening of the pigment in KOH and the smaller spores, but is externally similar to C. eggersii (section Iopodinae).

Crinipellis sect Iopodinae (Singer) Singer, stat nov Crinipellis sect Eti-Crinipellis subsect Iopodinae Singer, Lilloa 8: 498. 1942.

TYPE SPECIES Crinipellis iopus Singer. Pileus bright colored: red, carmin-purple, crimson, lilac, violet, not greening or

graying with KOH or NaOH, but sometimes becoming slowly slate blue with KOH.

Crinipellis sect 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 broader than 4.7 p, or else ellipsoid and then not more than 4 p broad, Q = 2 or larger.

TYPE SPECIES. Crinipellis iopus Singer. Only neotropical species of the subsection:

35. Crinipellis purpurea Singer, sp nov Fig 19. TYPE. Singer B 6151a (F), from Colombia. Pileo purpureo, KOH ope purpureo vel 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. Cheilocystidiis echinatis. Pilis variabilibus. Ad folia emortua delapsa Dicotyledonum. Typus in F conservatur.

Pileus purple red, with dark purple to purple-black center, about "Indian r." M&P in the larger marginal zone but reaching "maroon" M&P, fading to light reddish brown with fuliginous center or remaining purple when dried, appressedly radially pilose around a subglabrous central disc or papilla fibrillose-short-pilose, often slightly radially rimose towards the margin when adult, hairs irregular projecting over the margin and therefore often ciliate, convex, finally with depressed center, in the depression with a low circular elevation around a small umbilicus, or instead of the umbilicus with a small to medium more or less abrupt papilla, 6-8 mm broad. Lamellae white 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), eventually becoming deep umber brown from the base upwards or not changing with age, hispid-pilose all 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 suprahilar applanation or depression, smooth, hyaline, inamyloid, thin-walled, eventually sometimes thick-walled but 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, cylindrical or ventricose main body, echinate, sometimes only at the apex with finger-like, erect appendages, sometimes with many oblique lateral echiniform or spinulose appendages from about the middle portion, rarely from the base of the main body up to the apex, appendages 2-10 X 0.3-2(-2.5)p, with thin to very moderately thickened wall, hyaline, inamyloid; near the margin of the pileus often only the main body without appendages present. Hy- phae inamyloid, hyaline, non-gelatinized, with clamp connections. Hymenophoral trama regular, of elongated, interwoven hyphae. Covering layers: hairs of the pileus rather long, 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 purplish to hyaline-stramineous, in faded caps pale fuscous or pale brownish, fewer hyaline to fulvous brown, distinctly pseudoamyloid, but the upper portion often inamyloid, some with secondary septa, smooth; hypotrichium and spine-like agglutinations of smaller hairs 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 purplish black, not greening nor 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, sp.

MATERIAL STUDIED. COLUMBIA. Valle, Mun. Cali, Saladito, 1800 m alt, 15 Apr 1968, Singer B 6151a (F, type), BOLIVIA. La Paz: Nor-Yungas, Capellaria, 22 Feb 1956, Singer B 1318 (F), Coroico, 7 Feb 1956, Singer B 958 (F), 6 Feb 1956, Singer B 935 (F), 4 Feb 1956, Singer B 860 (F).

Crinipellis sect Iopodinae subsect Insignes Singer, subsect nov Pileo laete colorato, stipite breviore diametro triplici pilei, plerumque curvato vel

excentrico, insititio vel ad basin fibrilloso. Sporis eis subsectionis antecedentis similibus vel latioribus. Typus subsectionis: Crinipellis insignis Singer.

Key to the Species of Subsection Insignes

1. Saprophytes. 2. Spores 3.2-4,u broad. 38. C. sublividia. 2. Spores more than 4,.

3. Cheilocystidia scarce, inconspicuous, 3.7-5,u broad. 36. C. insignis. 3. Cheilocystidia making the edge of the lamellae heteromorphous. 37. C. eggersii.

1. Parasites. 4. Pileus crimson red, "Vandyke Red, Acajou Red," R., forming "krulloten" on Theo-

broma or growing from the base of living Vitex. Cheilocystidia simple or forked; spores up to 9(-1 1), or smaller. 5. On Theobroma; spores 7-9(-11) X 4-4.8(-5)u (if spores larger - see "3" above).

41. C. perniciosa. 5. On Vitex; spores 5-7 X 3-4,. 40. C. trinitatis.

4. Pileus lilac: on other trees, usually fruiting on living, seemingly healthy, eventually rotting cortex of standing branches of trees. Cheilocystidia mostly clavate, many or almost all with 2-6 short apical appendages; spores 9-14 X 4.5-8,. 39. C. siparunae.

36. Crinipellis insignis Singer, sp nov Fig 20. TYPE. Singer B 2320 (F), from Bolivia. Pileo purpureo vel purpureo-roseo, radiatim rugoso, minute fibrilloso-pubescente

17-19 mm lato. Lamellis albis, 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 ramulos ligneos emortuos. Typus in F conservatur.

Pileus purple ("India r. M&P") or purplish pink ("rose vale" M&P), the color concentrated in the hairs which are densest in the center and therefore the center darker than the margin, further towards the margin these appressed fine hairs forming a network, all appressed fibrillose-pubescent, on pallid or paler ground, radially sulcate along the through-lamellae but less so on the margin proper, 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 depressed center, 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 lamellulae connected by veins with the nearest through-lamella, sometimes forked, young lamellae without any intervenation, subfree, or very narrowly adnexed. Stipe yellow and purple colored, becoming very dark when dried, with scanty hyaline pubescence, generally seemingly glabrous when seen macroscopically, curved or straight, eccentric, stuffed, then hollow, equal or slightly ventricose, seemingly insititious, 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 4.5-6.5 1, ellipsoid to oblong, and sometimes somewhat curved when seen in profile, but without suprahilar depression, smooth, hyaline, inamyloid, 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 inconspicuous, 16-51 X 3.7-5 ,, fusoid or fusoid-subcapitate or wavy-filamentous, rarely subclavate, sometimes with slightly nodulose apex, hyaline. Hyphae of the trama of the pileus hyaline, inamyloid, with clamp connections. Hymenophoral trama regular. Hyphae of the stipe thick-walled, with clamp connections, appearing weakly pseudoamyloid 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 A broad walls 0.5-1 u thick, with rounded tips, weakly pseudoamyloid. Hypotrichium strongly developed around the hair-bases, but not all terminal hyphal cells giving rise to hairs and then with rounded tips, hyaline, thin to thick (1 u) wall, (5-)9-12 u, broad, strongly purple red incrusted (in NH40H).

KOH on the dried pileus causing darker (purple) spots, no greening or graying. On woody twigs in tropical forest. MATERIAL STUDIED. Known only from the type locality. BOLIVIA. Beni: Vaca

Diez. Riberalta, 29 Mar 1956, Singer B 2320 (F. type). This species is characterized by some primitive characteristics such as the scat-

tered inconspicuous cheilocystidia, the weakly pseudoamyloid hairs, the distant lamellae, and the slight development of the basal mycelium. Thus, it comes close to Marasmiellus with which, however, it has no demonstrable affinities. It is closest to the following species.

37. Crinipellis eggersii Patouillard in Patouillard & Lagerheim, Bull. Soc. Mycol. France 9: 125. 1893. Fig 21.

Maraslius eggersii (Patouillard in Patouillard & Lagerheim) Saccardo, Sylloge Fungorum 11: 37. 1895.

Marasmius vinosus Spegazzini, Anal. Mus. Nac. Buenos Aires 9: 264. 1909.

Type Eggers (FH), from Ecuador.

37a. Crinipellis eggersii var 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 recurved margin, depressed or umbilicate in the center, 3-32 mm broad. Lamellae white or gray ("cob- web" M&P), the ground of the lamellae often pale violet ("Hydrangea r." M&P) or up to half-breadth of lamellae concolorous with the pileus, with pallid edges at first, later sometimes part of edge faintly violet-purple under 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 separating, young without intervenation, old mostly more or less intervenose. Stipe concolorous with the pileus below and white at the apex, later deeper (8J 3, M&P), or chestnut to reddish black, finely and often sparsely villous-pruinate with subglabrous apex, mostly curved or oblique, often eccentric or becoming so, sometimes subcentral even in age but then shorter than pileus diameter, tapering upwards or 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 suprahilar depression, 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 numerous but not everywhere makinlg the edges heteromorphous, rising from hyphae running along 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, subcapitate or capitate apex, rarely filiform or subvesiculose, rarely with 1-2 short lateral or apical outgrowths, very rarely with several finger-like outgrowths, 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. Hymenophoral trama regular. Covering layers: 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 stramineous walls and at least in lower portion often violet or purplisli incrusted by the pigment, pseudoamyloid but sometimes 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 numerous clamp connections, strongly lilac or violet incrusted by the pigment when seen in NH4OH mount, the pigment tending to turn greenish in H2SO4 and KOH mounts.

KOH reaction on dried pileus negative or becoming slate blue. On dead branches and sticks, coriaceous petioles and rotting logs in subtropical

forests, tropical-montane forests, plantations and parks, always on Dicotyledones, fruiting in the rainy seasons.

MATERIAL STUDIED.ECUADOR. Guayas, Balao, 26 Jan 1892, Eggers (FH, type); Napo, Lago Agrio, 16 May 1973, Singer B 7504 (F). BOLIVIA. La Paz: Nor-Yungas: Rio Llolosa, 31 Jan 1956. Singer B 779 (BAFC); Rio Yariza, 16 Feb 1956, Singer B 1181 (BAFC). ARGEN- TINA. Tucuman, Quebrada de Lules, 18 Dec 1951, Singer T 1647 (LIL); Capital, Parque Roca, Spegazzini (LPS, type of Il vinosus Speg.).

One of the Bolivian specimens had 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 drying (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. Crinipellis eggersii var 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, circular to oval in outline, 4-10 mm broad. Lamellae white, often with purplish-livid ground, rather broad, 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-white at the apex, orange yellow, yellow or yellow brown at the base, with scattered hyaline hairs or above subglabrous, tomentose below when seen under a lens, subcentral to 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 partially ascendant on the lower part of the stipe and then tomentose-vellereous and 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 suprahilar depression, not curved, hyaline, inamyloid. Hymenium: basidia 18-28 X (7-)8-9 E; 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 (capitulum 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-walled with clamp connections, hyaline, inamyloid. Covering layers: hairs of the pileus scattered or in bunches, moderately long (35)60-200 X 5.5-12.5(-17)lp, irregularly shaped or filamentous, some subventricose or 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 intraparietal and incrusting pigment.

KOH on the dried pileus no color change, eventually slate blue or black, H2S04 likewise.

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-water level (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).

Crinipellis eggersii var flavipes 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 yellow pigment on the stipe. On the other hand, the following varieties appear to be quite constant.

37c. Crinipellis eggersii var epiphyllus Singer, var nov TYPE. Singer B 747, from Bolivia.

Carpophoris epiphyllis; multis cheilocystidiis appendiculatis. Typus in LIL. Pileus lilac, 3-5 mm broad. Lamellae white, distant. Stipe white, strigose to-

mentose near and at base but subinsititious, subglabrous above, 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 suprahilar depression but 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 numerous and making the edge heteromorphic in mature specimens, 27-58 X 2.8-10.3 ,, of two types, (1) ampullaceous with 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-or knot-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

trama hyaline, thin-walled, inamyloid, with clamp connections, but many secondary (clamp-less) septa present. Covering layer: epicutis of the pileus consisting of hairs which are hyaline or slightly fuscidulous inside or in the wall, some with short lateral excrescences, with obtuse tips, pseudoamyloud, accompanied by some shorter versiform epicuticular elements.

On dead fallen leaves and leaf petioles of Theobroma (cacao plantation) in the Yungas, Bolivia.

MATERIAL STUDIED. BOLIVIA. La Paz, NorYungas, Charobamba 30 Jan 1956, Singer B 747 (LIL).

37d. Crinipellis eggersii var 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 conservatus est.

Pileus violet to lilac ("Corinth," 47 J 1, 48 L 2, M&P), appressedly fibrillose and very finely pilose, glabrescent, in age often with a reticulate appressed fibrillosity which is lilac (on white ground), coarsely sulcate over 5/6 to 6/7 of the radius of mature specimens, convex, umbilicate, 13-27 mm broad. Lamellae white with pallid edges and mostly purplish-violet ground color or purplish violet in the upper half, rather narrow, then rather broad but not ventricose, distant (about 7 through-lamellae when mature), intervenose when mature, adnexed to adnate, often collariate. Stipe chestnut with white apex, macroscopically appearing glabrous but finely pilose with scattered hairs, the apex often glabrous, subcentral to somewhat eccentric, mostly curved, subequal or tapering upwards, 5-6 X 1-2 mm; basal mycelium present, consisting of minute white radiating fibrils. Context white, thin; odor none.

Spores 10-13 X 6-6.5 p ellipsoid, smooth, hyaline, inamyloid. Hymenium: basidia 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 cylindrical or ampullaceous to ampullaceous-ventricose and with a constriction underneath a clavate, subcapitate or capitate apex; apex, if ampullaceous, 3.5-8 p across, sometimes forked with two such narrowed apices, constriction, if present, 4-9 M wide, capitulum, if present, 7.5-12 j in diameter, sometimes with 1-5 lateral (rarely some apical) short-finger-like appendages, but the vast majority entire, sometimes rising from a vesiculose basal cell 10 1 diam, hyaline, thin-walled. Hyphae inamyloid, with clamp connections. Covering layers: epicutis of pileus consisting of sparse to rather numerous 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; hypotrichium consisting of numerous appressed hyphal 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 pseudoamyloid hairs.

KOH on dried pileus without reaction. On dead, fallen branches of dicotyledonous trees in hylaea. MATERIAL STUDIED. ECUADOR. Napo, Lago Agrio, 9 May 1973, SingerB 7316 (F) type. This variety appears to be intermediate between Crinipellis siparunae and Crini-

pellis eggersii. It differs from the first of these species in the predominantly entire, smooth cheilocystidia which are ampullaceous to subcapitate or capitate and in its habitat on dead wood (which does not necessarily mean that the mycelium can never be parasitic). It differs from var epiphylla in its habitat and color and from var eggersii in its color and presence of some appendiculate cheilocystidia.



50 Flora Neotropica

38. Crinipellis sublivida Murrill, North American Flora 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. Lamellae pale lilac, rather narrow, distant, very narrowly adnexed 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 irregular projections all over or only at the apex, more rarely ampullaceous or capitate, often forked, often asymmetric, walls thicker than those of the basidia, sometimes up to 1 jp thick. Hyphae thin-walled, with clamp connections. Covering layers: hairs of the pileus in minute tufts, often short or irregularly thickened, all pseudoamyloid and with rounded-obtuse tips. Hypotrichium of hyphae which are incrusted by a (now) brown pigment.

On dead branches, fruiting in January in virgin forest. MATERIAL STUDIED . Known only from the type locality. MEXICO. Veracruz:

Motzorongo near Cordoba, 15 Jan 1910, I'. A. & Edna L. Murrill 1047 (NY, type). The above description is based on a study of the type, published first in Singer

(1942). This species differs from the preceding one in the relatively longer stipe which

is less or not eccentric and has a distinctly tomentose basal mycelium and also in the narrower spores. It has not been re-collected in Mexico, probably because the virgin forests of Motzorongo no longer exist.

Crinipellis palmarum, from Africa, differs in having subclose lamellae, insititious stipe and slightly smaller spores.

39. Crinipellis siparunae Singer, Lilloa 8: 505. 1942. TYPE. Singer, from a greenhouse in U.S.S.R. Pileus lilac, brownish lilac, livid pink, at first densely covered by tufts of hairs

which later become sparser and 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 maintaining the original color, the outer two thirds of the radius costate-sulcate to vaguely sulcate in age, convex, umbilicate or umbonate or neither, but color and surface characters rather uniform, eventually flattened in part or all over, 6-22 mm broad. Lamellae white, narrow, later broad, reaching 4 mm broad in the largest carpophores, some forked or indistinctly rugulose-anastomosing in some specimens, in old specimens usually intervenose, medium close to distant, up to 14 through-lamellae present, adult tridymous with many lamellulae, rounded-adnexed but frequently soon separating with 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-squamulose, with less pilose-tomentose apex (which may be glabrescent), solid, mostly curved, 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, rarely ventricose or subcylindric, apically beset with rod-shaped or phialid-shaped to 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 similar elements without appendages visible and then main body usually 4-6.7 p broad, all hyaline, or some melleous-hyaline, inamyloid. Hyphae of the trama 5-10 p broad, filamentous, inamyloid, with

clamp connections. Hymenophoral trama regular, 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 generally also carpophores) on living trees, cortex of branches and twigs, often high above ground, in tropical and subtropical forests and in greenhouses. In the greenhouses fruiting in June. Known host: Siparuna sp.

MATERIAL STUDIED. CUBA. Wright 858, as Marasmius scalpturatus (ined.), (FH). BRAZIL. Rio Grande do Sul: Nova Petropolis, 1925, Rick 477 (FH). U.S.S.R.: Leningrad: Orangeries of the Botanical Garden, 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 larger than those of C. perniciosa.

It differs from Crinipellis eggersii and its varieties by growing generally on living host material, by apically multiappendiculate cheilocystidia which may or may not be intermixed with entire ones.

ILLUSTRATION. Singer (1942) fig 3e.

40. Crinipellis trinitatis 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 undulating and indistinctly striate margin, convex with a small umbilicus, then flattened and often depressed at the center, up to 25 mm broad. Lamellae "vinaceous buff" with reddish edges, rather close, didymous, adnate. Spore print pure white. Stipe "Acajou Red" (Ridgway), darker and browner below, downy, cylindric or slightly enlarged upwards, solid, (14-23 X 1.5-2.5 mm). Context "Vinaceous Buff" (Ridgway), dark brown on the base of the stipe, thin, thicker and browner at the disc of the pileus.

Spores 5-7 X 3-4 1, ellipsoid, inamyloid, [smooth]. Hymenium: cheilo-

cystidia 30 X 8 1, cylindrical or irregular and forked, thin-walled, red. Covering layers: hairs of the pileus about 300 p long, 5-6 broad, more or less hyaline, pseudoamyloid (deep red in Melzer's reagent) incrusted with light red granules which dissolve in 10% KOH solution to give a bluish purple tint, distantly septate, undulating, rather thin- walled: hypotrichium formed by polygonal thick-walled cells, walls red in Melzer's

reagent. On the bark of living trees, gregarious on base of Vitex sp. trunk. MATERIAL STUDIED. Known only from the type locality. TRINIDAD. Path to

Maracas waterfall, 25 Nov 1949, Dennis 376 (K, type), not studied. This species appears to belong in subsection Insignes, and on account of the rel-

atively small spores, it is different from all other species including C. 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 redescribed as C. trinitatis by Dennis (1970). Since I have not restudied Dennlis 376, the description given above is an adaptation of Dennis's (1951a) description (data derived from the illustration in 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. Crinipellis perniciosa (Stahel) Singer, Lilloa 8: 503. 1942. TYPE. Stahel, from Surinam (collections in Paramaribo ?), not seen. Marasmius perniciosus Stahel, Dept. Landbouw in Suriname Bull 33: 16. 1915. Pileus crimson red, generally faintly so according to Stahel, later bleaching to

whitish and dried material often whitish, with a red-black spot in the center, and with others which are radially arranged and have the same color and are visible on dried

material, radially grooved, campanulate, later expanded and often with a depressed center, 2-25 mm broad, mostly 5-15 mm in diameter. Lamellae white in dried mater-

ial, rather thick (0.2 mm) in mature material, medium broad to rather broad (1-2 mm) distant, (8-20 through-lamellae, mostly + 15), collariate. Spore print pure white. Stipe white except at the thickened subbulbous base 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 according to Stahel) in the

pileus, reviving. Spores 7-9 X 4-4.8 , (from print), but according to Stahel up to 11 X 5 ,, ellip-

soid, hyaline, smooth, inamyloid. Hymenium: basidia 31-32 X 7-(8.8),, 4-spored; cystidia none; cheilocystidia 35-50 X 9-14p, rather regularly bottle-shaped. Hyphae with

clamp connections. Covering layers: hairs of the pileus not numerous and 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-obtuse at the tip, pseudoamyloid, 80-150 X 4-12,; hypotrichium consisting of thick-walled, inamyloid hyphae.

The mycelium infecting the living plants of Theobronia but 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, apparently originally only in northern South America, but the disease has spread to wherever Theobrolia has been planted, from Ecuador to the West Indies. Known hosts: Theobroma cacao, T.

bicolor, T. speciostum (a wild species). MATERIAL STUDIED. SURINAM. Paramaribo: Stahel (FH), authentic, additional mater-

ial (FH). BRITISH GUIANA. Coil. & det. F. L. Stevens (FH). Data on the infection and the effect on the host as well as suggested control

measures should be looked up in Stahel's original paper or in the short digest given by Singer (1942). The fact that the fungus causing the "krulloten disease" is still fre-

quently quoted asMarasmius perniciosus, 30 years after its transfer to Crinipellis, is a

good illustration of the "conservatism" of some phytopathologists and their reluctance to adopt the results of mycological work.



Chaetocalathus 53

ILLUSTRATIONS. Singer (1942), fig 2 a; see also Stahel (1919) pl 1-8; (1915), pl 1-12; Dennis (1970), pi 3, fig 4.

2. Chaetocalathus Singer, Lilloa 8: 518. 1942. TYPE SPECIES. C. craterellus (Durieu & L6veillV) Singer. Habit pleurotoid; pileus with thick-walled pseudoamyloid (to almost amyloid)

hairs which are long and distinctly separated from 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-spored ones; 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 pseudoamyloid; stipe rudimentary or more rarely absent, not directly attached to the substratum; cheilocystidia always present; trama inamyloid; hyphae with numerous clamp connections. On wood, bamboo stalks, bark, leaves and stems.

Key to the Sections, Subsections, and Species of Chaetocalathus

1. Pseudoamyloid seta-like or metuloid sterile bodies in the lamellae absent; only inamyloid cystidia and/or cheilocystidia present. [Not represented in the neotropics]. sect Chaetocalathus.

1. Pseudoamyloid setoid or metuloid bodies in the lamellae present, rarely confined to the edge of the lamellae (in young specimens) and intermixed with inamyloid cheilocystidia. 2. Pseudoamyloid bodies relatively thin, seta-like, acute, branched. sect Meristocystis, p 53.

1. C. columellifer. 2. Pseudoamyloid bodies reaching more than 6ji broad, metuloid, or restricted to the

edge (at least in young specimens) and cystidioid, simple or 2(-4) forked at the apex and reaching over 7p diam. sect Holocystis, p 54. 3. Pigment none, only old specimens may assume a cream colored tinge.

4. Cystidia thickwalled metuloid, pseudoamyloid, often thickly incrusted, on the side of the lamellae, numerous, all simple and entire; cheilocystidia often apically forked. Common neotropical species. 2. C. liliputianus.

4. Cystidia not as above; cheilocystidia, often forked, but also entire and merely appendiculate or muscronate along the edge, later pervading the sides of the lamellae. 3. C. niduliformis.

3. Pigment present; pileus colored from the beginning. 5. Spores relatively not broad: 9.5-10.5 X 5-5.3. Pileus grayish-leather-color. On

living Murraia. 4. C. asperifolius. 5. Spores relatively broader; pileus purple red or orange.

6. Pileus purple red. 5. C. carmelioruber. 6. Pileus dull orange. 6. C. aurantiacus.

Chaetocalathus sect Meristocystis Singer, Lilloa 8: 522. 1942. Characters as indicated in the key above. TYPE SPECIES. C. africanus (Patouillard) Singer Only one species is known from the neotropics.

1. Chaetocalathus columellifer (Berkeley) Singer, Sydowia 9: 398. 1955. Fig 23. Agaricus columellifer Berkeley, Hook. Journ. Bot. Kew Misc. 4: 161. 1852. Marasmius columellifer (Berkeley) Pilat, Trans. Brit. Mycol. Soc. 33: 219. 1950, invalid. TYPE. Borneo (K). Pileus white or more rarely yellowish white, strongly radially pilose-woolly or

woolly-subsmooth with substrigose hairs, 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 sublateral area of the sterile surface and appearing cup-shaped, circular to sub- elliptical, sometimes with a short pseudostipe, 1-8 mm broad. Lamellae white, dry pale buff-tan (to umber when dried), rather narrow to medium broad, intermixed with lamellulae when mature, moderately close to distant, concurrent to the stipe rudiment to which they are adnexed or adnate, spore print white. Stipe only rudimentary and 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 pseudoamyloid (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 appearing on the sides of the lamellae, but may be absent in some sections, 31-90, long, entire or forked, especially near the apical region, narrowly fusoid to needle-shaped or cylindric, but often with a ventricose swelling below, tip and branch-tips acute 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 reaching up to 7.5 p across, often deep-rooting without a septum and then reaching up to more than 100 p in length. Hyphae of the regular hymenophore hyaline, inamyloid, with some inflated cells intermixed, with clamp connections. Covering of 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 intermediate between those of the pileus and the hymenial cystidia.

On palm fragments and frondose twigs fallen to the ground, gregarious, fruiting in the rainy seasons in the lowland forests.

MATERIAL STUDIED. PANAMA. Barro Colorado Island, 30 Jun 23 Aug 1952, Martin & Welden 7279, 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.

Chaetocalathus sect Holocystis Singer, Lilloa 8: 526. 1942. Characters, see key above. TYPE SPECIES. C. carnelioruber Singer,

2. Chaetocalathus liliputianus (Montagne) Singer, Lilloa 8: 527. 1942. Fig 24, 25, 26.

Agaricus liliputianus Montagne, Ann. Sci. Nat. IV. 1: 99. 1854. Pleurotus liliputianus (Montagne) Saccardo, Sylloge Fungorum 5: 385. 1887. Marasmius nidulus Berkeley & Curtis, Jour. Linn. Soc. 10: 299. 1868. Crinipellis nidulus (Berkeley & Curtis) Patouillard, Jour. de Bot. 3: 336.1889 ("nitidulus"). Crinipellis calosporus Patouillard in Duss, Flore Cryptogamique des Antilles frangaises,

Champignons, p 255. 1904. Marasmius calosporus (Patoulliard) Saccardo & D. Saccardo, Sylloge Fungorum 17: 47. 1905. Pleurotopsis calospora Murrill, North American Flora 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 substratum by a central to sublateral area and appearing cupuliform, smooth or with sulcate margin, sometimes with a short pseudostipe, convex to almost resupinate, 1.5-10 mm broad. Lamellae white, dry



Chaetocalathus 55

often yellowish white, narrow to broad (mostly about 1-2 mm broad), subdistant or distant, intermixed when mature, concurrent to the rudiment of the stipe which is either central or sublateral, and then often merged with the pseudostipe, the lamellae attenuated and narrowly adnexed or free. Spore print white. Stipe rudimentary unless lateral and confluent with a pseudostipe, sometimes so poorly developed that it is practically absent but more frequently knob-like, low, white pubescent, snow- white when seen under a lens. Context extremely thin to relatively thick, white; odor none.

Spores 7.5-11.5(13) X 5-8 p, rather broadly ellipsoid, without a distinct suprahilar applanation or depression, with thin, later slightly thickened wall which is inamyloid or in some specimens, especially when over-aged more 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 appendages like the cheilocystidia, with pseudoamyloid 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 similar but more often with 1-4 apical appendages or 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 appendages or burge, but usually the main body simple, rather polymorphous, 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 irregularly shaped but mostly filamentous, in an upper layer more radially arranged and more persistently filamentous, not gelatinized, inamyloid, hyaline, with clamp connections. Covering layer of the pileus consisting of dense hairs 100-400 long which have thick (3-5.5 p), hyaline walls which are pseudoamyloid and incrusted by a thin, finely granular incrustation, or without it, with acute to obtuse tips; hypotrichium consisting of prolongations of the hyphae of the trama where the last member below the hair is slightly swollen and more or less cheilocystidium-like but, instead of the appendage bearing the hair, divided from it by a clamped septum, inamyloid.

On fallen branches and 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; Clibadium erosum; 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, GuznaLn 4524 (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 Flora Neotropica

3. Chaetocalathus niduliformis (Murrill) Singer, Lilloa 8: 521. 1942. Pleurotus niduliformis (Murrill) Saccardo, Sylloge Fungorum 23: 117. 1925. Pleurotopsis niduliformis Murrill, North American Flora 9: 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. Lamellae white, subclose to subdistant or distant, medium broad, concurrent to

stipe rudiment, adnexed to or free from it. Stipe rudimentary and 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, pseudoamyloid. 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 ampullaceous or clavate with

one to three apical appendages which have obtuse apices, most frequently bifurcate at

apex, appendages 2-25 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 gradually becoming more numerous (but always sparse) outside the broad edge region as pleurocystidia, but these not different from cheilocystidia. Hyphae hyaline, with clamp connections, inamyloid. Hymenophoral trama regular, in

places appearing subirregular because of swelling and inflated hyphal ends (31-70 X 14- 19 u). Epicutis: hairs long but ordinarily not reaching 300 u, 2.8 , broad rarely partially inflated and there reaching 9.5 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-sharp acute. On stipe rudiment pseudoamyloid smaller hairs.

On dead twigs of Juniperus barbadensis, according to African collectors (reported by Pegler) also on Morinda citrifolia and Uvaria chazae, solitary or gregarious. Bermudas, also in West Africa (Sierra Leone, Ghana, Congo).

MATERIAL STUDIED. BERMUDA. Brown, Britton & Sealer 1350 (type, NY), 1352. Zaire. Kwango: Kinanga, H. Vandervst, 11 Apr 1910 (BR).

This species is somehow intermediate between 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. Chaetocalathus asperifolius (Patouillard) Singer, Lilloa 8: 530. 1942. Crinipellis asperifolia Patouillard, Jour. de Bot. 3: 336. 1889. Marasmius asperifolius (Patouillard) Saccardo, Sylloge Fungorum 9: 70. 1891. Pleurotopsis asperifolia (Patouillard) Murrill, North American Flora 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. Lamellae concolorous or paler, medium broad, attenuate, medium distant to

subdistant, intermixed. Stipe rudimentary, very small, hemispheric, hirsute, concolorous or paler to almost white, disappearing in 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,, projecting beyond 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-walled incrusted, strongly pseudoamyloid with rounded tips. 4-6.8 ,broad.

On bark of living Murraya exotica, after heavy rains, gregarious. MATERIAL STUDIED. MARTINIQUE. Duss 35 (FH), type. GUADELOUPE. Duss 67

(FH).

5. Chaetocalathus carnelioruber Singer, Lilloa 8: 526. 1942. TYPE. Linder, from Surinam (FH). Pileus "carnelian Red (R.) when fresh becoming a color between "Perilla Purple"

and "Mineral Red" (R) when dried, with paler margin, subtomentose, strongly pilose under a lens, appearing almost cup-shaped, attached to the sterile surface and mostly reflexed, reniform in circumference with a deep incision on the concave side, 2-7 mm broad when dried. Lamellae cream buff when fresh, brownish when dried, very narrow distant or moderately distant, attenuate towards the incision of the pileus. Stipe rudimentary, almost absent in the young carpophores but under a lens visible as a minute, verruciform, lilac-brown, hirsute, button-like, function-less structure. Context very thin.

Spores 11 X 7 p, ellipsoid-ovoid, with granular contents, thin-walled, Hymenium: basidia 34.8 X 8.8 ; cystidia 38-100 X 13-32p fusoid, hyaline or yellowish in NH40H, pseudoamyloid or nearly amyloid, with very thick walls, which are polystratous and 5-8.5 1 thick in the largest cells, incrusted by finely granular material in the upper portion, all entire. Hyphae with clamp connections. Hairs of 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. MATERIAL STUDIED. SURINAM, Bartica, 9 Dec 1923, D. H. Linder (FH), type. ILLUSTRATION. Singer (1942), fig 6 f i.

6. Chaetocalathus aurantiacus Singer, sp nov TYPE. Singer B 2201, from Bolivia.

Pileo obtuse aurantiaco; ceterum C. carnelioruber similis. Typus in F conservatur.

Pileus dull orange, finely pilose, finely long-pilose when 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. Lamellae orange-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 suprahilar depression or applanation, smooth, hyaline, inamyloid. Basidia 25-29 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 pseudoamyloid (exceptionally some inamyloid); cheilocystidia 30-32 X 4-6 u, narrowly ventricose to subfusoid, often constricted in the middle, often appendiculate or with bi- to tri- lobed apex, wall thin to slightly thickened (to 1 ,), inamyloid. Hyphae hyaline, not gelatinized, inamyloid, with clamp connections. Cortical layer: hairs of the surface of the pileus dense, remarkably thin (3-4 p broad) and long, wall 1-1.2 p thick, pseudoamyloid, tips obtuse or subacute.

On dead dicotyledonous branches in hylaea. MATERIAL STUDIED. BOLIVIA. Pando, Manuripi, Conquista, 25 Mar 1956, Singer B

2201 (F), types.



58 Flora Neotropica

3. Lachnella Fries, Corpus Florarum Sueciae 1. Floram scanicam sceipsit, p 343. 1835.

Characters as given in the keys (p ), see also Singer (1973), p TYPE SPECIES. Lachnella alboviolascens Albertini et Schweinitz ex Fries

Key to the Subgenera and Species of Lachnella

1. Cystidia metuloid. Subgenus Metuloidifera Singer. 1. L. cecropiae. 1. Cystidia none. Subgenus Lachnella.

2. Spores elongated and almost sickle-shaped, 15.5-17 X 5-6.5p; carpophores very small, up to 0.25 mm high and 0.4 mm broad. 2. L. subfalcispora.

2. Spores less elongated or fruiting bodies larger. 3. Tropical species from northern South America; spores 12-17 X 5-8,. 3. L. caracassana. 3. Subtropical species; spores 10-13.5 X 5-8.3p. 4. L. villosa.

Lachnella subgen Metuloidifera Singer, Darwiniana 14: 13. 1966. Characters as given in the key above.

Fig 27. 1. Lachnella cecropiae Singer, Darwiniana 14: 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. Lamellae none, hymenial surface smooth, white, not deeply excavated but convave. Stipe or pseudostipe none. Context

very thin, white, waxy, slightly harder when 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 pseudoamyloid wall, 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. Covering layers: 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 separated by a sedondary (clampless) septum and thin-walled, but generally thick-walled all over and aseptate.

On leaves of Cecropia in the Amazonas region. MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, Ivon 3 Apr 1956, Singer B 24901

(LIL, BAFC), type.

Lachnella subgenus Lachnella Characters given in the key above. TYPE SPECIES. Lachnella alboviolascens Albertini & Schweinitz ex Fries.

2. Lachnella subfalcispora Reid, Kew Bull. 15: 267. 1961. TYPE. Dennis 1472, from Venezuela (K). Pileus a small cup, white, becoming subglobose or turbinate when dry, densely

crowded, the sterile surface villous, 0.1-0.4 mm broad. Lamellae none, hymenial surface 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: basidia up 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

carpophore more or less appressed to the sterile surface and mostly overtopping the 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 p broad, with thickened hyaline wall which thins out towards the apex which may be slightly swollen, pseudoamyloid, incrusted by a granular substance which dissolves in KOH (in which medium the hairs become locally swollen and distorted).

On twigs, associated with Cyphellopsis subglobispora. MATERIAL STUDIED. VENEZUELA. Dto Federal, Caracas Botanic Garden, 30 Jun 1958,

Dennis 1472 (K), type (seen, not sectioned; description adapted from Reid). ILLUSTRATION. Reid (1961), fig 2, 2; 3, 6.

3. Lachnella caracassana Reid, Kew Bull. 15: 267. 1961. TYPE. Dennis 1876, from Venezuela (K). Pileus in form of a cup, white, villous on sterile surface, subglobose when dry,

0.3-0.4 mm broad. Lamellae none: hymenial surface smooth, white. Stipe none but carpophores narrowed into a very short, stipe-like base which is usually buried beneath the superficial layer of bark.

Spores 12-17 X 5-8 1, elongated, somewhat asymmetrically obpiriform, hyaline, smooth, inamyloid. Hymenium: consisting of basidia, no cystidia. Hyphae as in the

preceding species but in the basal region of the carpophore more agglutinated. Cor- tical layer: hairs sheathing the fructification much like those of the preceding species.

On twigs. Gregarious but carpophores scattered. MATERIAL STUDIED. Venezuela. Dto Federal, Caracas Botanic Garden 14 Jul 1958,

Deiinis 1876 (K), type (seen, not sectioned; description adapted from Reid). ILLUSTRATION. Reid 1961 fig 3, 4

4. Lachnella villosa (Persoon ex Schweinitz) Gillet, Champignons de France, Dis- comycetes, p 80. 1881.

Peziza villosa Persoon ex Schweinitz, Schriften, Naturforschende Gesellschaft Leipzig 1: 120. 1822 (non Chevelier 1826).

Cyphlella villosa (Persoon ex Schweinitz) Crouan, Flore Finist4re, p 61. 1867. Trichopeziza villosa (Persoon ex Schweinitz) Fuckel, Jahresbericht des Nassauer Vereins 23-24:

296. 1870. Chaetocv 'pa villosa (Persoon ex Schweinitz) O. Kuntze, Revisio Generum Plantarum 2: 848.

1891. C'plhella dochimiospora Berkeley & Broome, Ann. Mag. Nat. Hist. IV, 11: 343. 1873. Cliaetocvpha dochiniospora (Berkeley & Broome) O. Kuntze, Revisio Generum Plantarum 2:

847. 1891. TYPE. Persoonl, from Germany (L). Pileus white, in form of a cup which has inrolled margin when dry, hairy-villous

on the sterile surface, nearly 0.5 mm broad. Lamellae none, 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, pseudoamyloid wall, incrusted by hyaline small crystals.

On dead wood and leaves of Dicotyledones. Known host in South America: Abius jorullensis var spachii. Probably confined to the montane and subtropical zone. In the neotropics said to occur in Ecuador; also in Europe and North America, Australia.



60 Flora Neotropica

MATERIAL STUDIED. ARGENTINA. Tucuman, Tafi, Tafi del Valle 1 May 1949, Singer T 516 (LIL). GERMANY. Konigstein, Krieger, Fungi Saxonici 1457 (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 agreement with the data obtained by Donk (1959) from the type preserved in the Persoon Herbarium.

ILLUSTRATION. Patouillard (1900), fig 38, 8.

4. Amyloflagellula Singer, Darwiniana 14: 14. 1966. Characters, see key p 4. and Singer (1966), p 14.

Type species: Cyphella pulchra Berkeley & Broome Only one species known in the neotropics.

1. Amyloflagellula pseudoarachnoidea (Dennis) Singer, Darwiniana 14: 14. 1966. Fig 28.

Marasnius pseudoarachnoideus Dennis, Trans. Brit. Mycol. Soc. 34: 425. 1951. Crinipellis pseudoarachnoidea (Dennis) Singer, Sydowia 15: 60. 1961. TYPE. Dennis 137, from Trinidad (K). Pileus white or pale buff, deeply sulcate, convex, 3-4 mm broad. Lamellae pale

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 X 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 rhizomorphs consisting of relatively broad, erect or ascendant hyaline and inamyloid terminal cells of hyphae, many of these forked or twice forked, their apices rounded-obtuse and with a disc-like strictly apical pseudoamyloid zone from which 4-10 flagellum-like thin, apical appendages arise, these strongly pseudoamyloid, 1.7-4.8 ulong.

On forest litter, solitary. MATERIAL STUDIED. TRINIDAD. Arena Forest, 5 Oct 1949, Dennis 137 (K),

type. The macroscopical description is given according to Dennis (1951 a) who illustrated

this species (pl 19, fig 20); also Dennis (1970), pl. 2, fig 20.

5. Flagelloscypha Donk in Singer, Lilloa 22: 312. 1951. Characters: See key characters. TYPE SPECIES. Cyphlella minuttissinla Burt. Only one species thus far observed in the neotropics.

1. Flagelloscypha polylepidis Reid, Kew Bull. 15: 269. 1961. TYPE. Dennis 1809, from Venezuela (K). Pileus in form of cups, white, villose, becoming globular when dry, shallowly dis-

coid in profile, up to 0.7 mm broad. Lamellae none; smooth hymenial surface in the concave portion of the cup white. Stipe or pseudostipe none; only a rudimentary stipe-like base present.

Spores 8-10 X 5-6.5 ,, broadly ellipsoid to pin-shaped, smooth, inamyloid. Hymen- ium: basidia 23.3-41.6 X 5.5-6.5 I, 4-spored; cystidia none. Hyphae of the monomitic trama thin-walled, 2.5 pin diameter, with clamp connections. Cortical layer: epicutis



Flagelloscypha 61

consisting of hairs densely covering the sterile surface of the cups, up to 150 X 2.5-3 Iu, densely covered by detersile rod-shaped crystals, divergent, with slightly thickened wall, thinning out towards the strongly tapering whip-like apex and also towards the often somewhat narrowed base, long, narrow, hyaline, continuous (non-septate).

On gall of Poll'lepis sericea. MATERIAL STUDIED. VENEZUELA. Merida: Sierra de Santo Domingo, Laguna Negra,

3440 m alt 27 Jul 1958, Detnnis 1809 (K), type (seen not sectioned; description above according to type description by Reid).

ILLUSTRATION. Reid (1961), fig 3, 1.

Tribe Marasmieae subtribe Marasmiinae Singer, Sydowia 15: 60. 1961. TYPE GENUS. 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. Hymenophore lamellate, the lamellae intervenose or not. 2. Epicutis not hymeniform {see Marasmnius, p 62). 2. Epicutis hymeniform.

3. Epicutis consisting of or containing many broom cells (see Marasmius p 62). 3. Epicutis consisting of smooth, mostly entire cells.

4. Capitate and/or metuloid and/or oleo-cystidia present in the epicutis and/or the hymenium; stipe central, eccentric, lateral or absent; small carpophores (pileus < 15 mm, <5 mm if centrally stipitate), some instititious. Gloiocephala, p 284.

4. Capitate, metuloid and oleo-cystidia absent unlessthe pileus is centrally stipitate and rising from basal mycelium and then pileus reaching larger size.

Marasmius, p 62. 1. Hymenophore with lamellae which are reduced to a few veins or so strongly anas-

tomosing that they appear poroid or merulioid, or else hymenial surface smooth. 5. Epicutis reduced since the pileus is head-like and hollow or else the portion of the

pileus (head) around and near the apex of the pseudostipe sterile; trama always very thin in the pileus (head). [Other types of pileus configuration of the "physalacrioid" type not represented in the neotropics.] Physalacria, p 304.

5. Epicutis on the sterile upper side of the solid pileus well developed and not reduced to a sterile zone; trama thin to rather thick. 6. Epicuticular cells in the form of broom cells (Rotalis-type); carpophores minute,

pileus less than 5 mm broad. 7. Hymenial cystidia none; basal mycelium present; hyphae at least partly

pseudoamyloid. Epicnaphus, p 312. 7. Hymenial cystidia present; basal mycelium absent; hyphae all inamyloid.

8. Cystidia absent; stipe rising from a rhizomorph. Manuripia, p 312. 8. Cystidia present; stipe rising from the substratum (see "3" above).

6. Elements of the epicutis not containing broom cells. 9. Carpophores with a central, eccentric or lateral stipe, more rarely sessile. Epi-

cutis of the pileus hymeniform. 10. Carpophores rather large with long central stipe and smooth hymenial

surface, with basal mycelium; pigments of all parts of carpophore bright (red or yellow). Hymenogloea p 313.

10. Carpophores with pileus to 15 mm broad, more often less broad, and never bright red or yellow if centrally stipitate; stipe often institious (see "4" above).

9. Carpophores with a central or eccentric, rarely lateral pseudostipe (attached to the side of the pileus opposite to the hymenial surface). Epicutis of the pileus hymeniform or not. 11. Capitate dermato- or oleo-cystidia present (see Physalacria, p 304). 11. Capitate dermato- and oleo-cystidia absent. Rimbachia, p 314.



62 Flora Neotropica

6. Marasmius Fries, Genera Hymenomycetum, p 9. 1836. Heliomyces LUveille, Ann. Sci Nat. III. 2:177. 1884 Androsaceus (Persoon) Patouillard, Hymenomycetes Europaei, p 105, 1887. Chanaeceras 0. Kuntze, Revisio Generum Plantarum 3, 2: 454. 1898. Mycenitis Earle, BulL New York Bot. Gard. 5: 414. 1909. Collybiopsis (Schroter in Cohn) Earle, Bull New York Bot. Gard. 5: 415. 1909. Basionym: Marasmius subgenus Collybiopsis Schroter in Cohn, Kryptogamenflora von Schlesien 3a: 559.

1889. Tephrophana Earle, BulL New York Bot. Gard. 5: 27. 1909. Scorteus Earle, Bull. New York Bot. Gard. 5: 415. 1909. Polvinarasmius Murrill, North American Flora 9: 286. 1915. ? Vanromburghia Holtermann, Mycologische Untersuchungen in den Tropen, p 104. 1898. TYPE SPECIES Marasmius rotula (Scopoli ex Fries) Fries Habit collybioid (to almost mycenoid) or pleurotoid, mostly somewhat toughish

and reviving after drying out when remoistened; hymenophore lamellate, rarely smooth or venose, lamellae sometimes intervenose or even anastomosing and appearing poroid: epicutis either hymeniform or not hymeniform: if not hymeniform consisting of broom cells or at any rate strongly diverticulate-nodulose hyphal elements (and then either the whole trama or only that of the stipe more or less distinctly pseudoamyloid or at least the stipe seta-like and shining, blackish, insititious, or absent to small and eccentric to lateral); trama of the pileus monomitic, pseudoamyloid or inamyloid, no gelatinous layers present; clamp connections present, or, more rarely, absent; if absent, epicutis with broom cells; black rhizomorphs often present, sometimes stipe rising from these: telepods often formed; if the basal mycelium present, the epicutis always hymeniform; some species forming endotrophic mycorrhiza, none ectotrophs or lichens. Most frequently on wood or leaves, dead or living, more rarely on other kinds of litter, among mosses or grasses on earth, parasitic or saprophytic. Development of the carpophores hemiangiocarpous at least in some species (paravelangiocarpous according to Reijnders; 1963; pileostipitocarpous in the type species according to Reijnders). Almost cosmopolitan, but much more numerous in carpophore production and number of species in the tropical regions than in the temperate and frigid zones.

Key to the Sections and Subsections of Marasmius

1. Epicutis hymeniform, rarely later subhymeniform. 2. Stipe insititious and central and lamellae with a collarium; epicutis hymeniform,

consisting of broom cells. sect Marasmius. p 91. 3. Epicuticular broom cells of the Rotula type. subsect Pararotulae, p 92. 3. Epicuticular broom cells of another type.

4. Epicuticular broom cells of the Siccus-type. subsect Penicillati. p 121. 4. Epicuticular broom cells, even those of the marginal region of the pileus, with

erect finger-like appendages interspersed with some large setulae (setoid bodies) arising directly from the subjacent hyphae and undivided.

subsect Penicillati, p 121. 2. Stipe insititious and central, or not insititious or not central; lamellae without a

collarium, free or directly attached to the apex of the stipe; epicutis of pileus with or without broom cells. 5. Stipe insititious and central, if not central, epicutis of smooth cells exclusively.

6. Upper part of stipe, lamellae and larger marginal portion of the pileus white or whitish-cream; lamellae sometimes absent, or replaced by veins or a merulioid hymenophore; epicuticular cells and cheilocystidia smooth (if diverticulate, not with pigmented setulae). Section Epiphylli, p 87. 7. Apex of stipe consisting of inamyloid hyphae. subsect Epihyllini, p 87. 7. Apex of stipe pseudoamyloid. subsect Eufoliatini, p 90.

6. Pileus and stipe more strongly pigmented; lamellae well formed; epicuticular cells and/or cheilocystidia, at least some of them, appendiculate or beset with setulae.



Marasmius 63

8. Elements of the epicutis smooth; cheilocystidia corncob-like (not smooth). sect Scotophysini, p 86.

8. Elements of the epicutis at least in part with appendages or else with setulae and then of the Rotalis-type, or many broom cells seriate. 9. Small species; most epicuticular cells are broom cells of the Rotalis-type

or they are aggregated-seriate. sect Hygrometrici, p 79. 9. Medium sized species; many epicuticular elements smooth, the others

with apical or subapical finger-like appendages (not of the Rotalis-type). sect Leveilleani, p 85.

5. Stipe either not insititious or not central (or neither). 10. Stipe central or subcentral; basal mycelium generally strongly developed and

often strigose; epicuticular elements either smooth or in form of broom cells of the Siccus-type.

11. Elements of the epicutis all smooth without setulae. 12. Hyphae of the trama pseudoamyloid; spores cyanophilic. sect Globulares, p 269. 12. Hyphae of the trama inamyloid; spores acyanophilic. sect Alliacei, p 263.

11. Elements of the epicutis partly smooth and partly in form of broom cells or else all cells of the epicutis in form of broom cells of te Siccus-type.

sect Sicci, p 159. 13. Hyphae of the trama inamyloid. subsect Inaequales, p 159. 13. Hyphae of the trama pseudoamyloid. subsect Siccini, p 159.

10. Stipe strongly eccentric to lateral or even rudimentary to absent in mature specimens, only very young carpophores may be centrally stipitate; epicuticular elements in the form of broom cells (which may be of the Rotalis- or Siccus-type, in the latter case setulae often forked or diverticulate), but occasionally many or even the majority smooth; base of stipe or point of attachment of the pileus either insititious or carpophore rising from a tomentose or fibrillose, usually white, small patch of 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; pleurocystidia in form of fusiform entire cystidia with rather thin walls but resinous colored incrustation present; stipe present and not insititious; most hyphae of the pileus-trama pseudoamyloid (see "15" below).

14. Epicutis of pileus hymeniform or subhymeniform, in the latter case cystidia either not incrusted or incrusted cystidia only on and very near the edges, or cystidia absent (excepting for broom cells which may occur at the edge of the lamellae as cheilocystidia); stipe present and either insititious or not, or pileus sessile: hyphae of the pileus trama pseudoamyloid or inamyloid. sect Neosessiles p 253.

1. Epicutis of the pileus definitely not hymeniform or subhymeniform but in form of interlaced hyphae or terminal bodies which bear setulae. 15. Stipe central, insititious or inserted directly in a dark rhizomorph; conspicuous

incrusted pleurocystidia none. sect Androsacei, p 63. 15. Stipe eccentric or sublateral, sometimes (rarely) none, rising from a small white

mycelial patch appressed to the substratum (wood); conspicuous incrusted pleurocystidia always present. sect Fusicystides, p 251.

Schematic drawings showing the principal differences between the sections of the genus Alarasmiius may be compared in Singer (1965), plates I-IV (except for section Leveillea ni).

Marasmius sect Androsacei Kilhner, Le Botaniste 25: 60. 1933 (as "Androsaceae"). TYPE SPECIES. 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 generic position of the specimen under investigation is doubtful, a monograph of the neotropical Marasmielli should also be consulted (Singer 1973).

Key to the Species of Section Androsacei

1. Clamp connections present. 2. Stipe glabrous and smooth (even under a good lens).

3. Combining the habitat on wood sticks of Alnus or other woody material in montane alneta with white, distant (through-lamellae 7-11) lamellae; hypodermium with inconspicuous or no incrustation; stipe less than 15 mm long; spores 5.5-9.5 X 2.5-3.5, most frequently 7-8.8 X 2.5-3.2, 1. M. valae.

3. Not combining the above characters; on leaves (if on woody sticks or roots see "5" below). 4. Dried pileus papillate or strongly umbilicate, 2-3.5 mm broad; stipe relatively

long: 25-75 mm; through-lamellae relatively few (7-10); species growing on Polylepis leaves in the fog forest and sub-paramo. 2. AI. pol)ylepidis.

4. Pileus either larger, or not or weakly depressed when dried; through-lamellae in mature specimens at least 11. 3. M. cyrillidis.

2. Stipe, at least when young and fresh, finely pubescent or pruinate, pilose, or finely setulose, sometimes macroscopically appearing glabrous, but under binocular dissecting microscope distinctly covered by microscopical hairs, setae, or dermatocystidia (if stipe rising from rhizomorph, see "15" below). 5. Spores 10-12 X 2.7-3.7,. 4. M. bactrosporus. 5. Spores less elongated or broader, not strictly rod-shaped.

6. Lamellae gray when fresh, nine; stipe tending to be eccentric; on dead bark of leguminous trees. 5. M. legziminosarulm.

6. Lamellae not gray when fresh, (7-)10 or more through-lamellae; on dead wood or dead leaves. 7. Covering of the stipe consisting of short, acute, subulate-spinose diverticula

which are numerous but not crowded, erect to ascendant, subhyaline, 3.3-12 X 1-4u. West Indies. 6. Al earlei.

7. Covering of the stipe different. 8. Growing on leaves.

9. Growing on (wood and) fallen leaves of Liquidambar in Mexico; pileus vinaceous brown only in youth, soon with very pale margin which is, when mature, radially rugose or sulcate up to 6/7th of the radius sulcate; hypodermium only in the disc-region spadiceous incrusted, otherwise incrustation on the hyphae rather inconspicuous, pigment where present mainly intraparietal; cystidioid elements of the covering of the stipe < 70. 7. .M. liquidambari.

9. Growing on fallen leaves of various dicotyledonous hosts; pileus and hypodermium as above or less sulcate and with deeper colored and/ or more conspicuously incrusting material; cystidioid hairs of the covering of the stipe as above or different. 10. Stipe densely beset with villous hairs especially in the upper

portion, these hairs long with thin (0.3p) to slightly thickened hyaline to brownish hyaline wall, obtuse above, and sometimes grossly diverticulated, in our material 8-60 X 4.5-9.5u; stipe color deep (chestnut) brown; spores 7-9 X 3-4p; on fallen leaves of fagacious trees. 8. M. splachnoides.

10. Stipe finely setulose or pruinate, elements of the stipe covering different from those described above, or at least different elements present; stipe cinnamon, chestnut or black; spores smaller or narrower; on fallen leaves of oak or more often on other substratum.

11. Lamellae medium close to subdistant, more rarely close or distant; stipe when seen under a dissecting microscope in young and fresh condition distinctly covered by long, acute, hyaline bristles which are often incrusted, but are not persistent, often breaking off but sometimes later regenerated, in the basal third of the stipe also long Crinipellis-like hairs present, these thick-walled, hyaline to aple melleous, the inner wall layer often pseudoamyloid, tapering gradually to



Marasmius 65

the apex, up to 150 or 170u long; stipe always with setae or pubescent in the lower portion. 9. M. hakgalensis.

11. Lamellae close to medium close; covering of the stipe often consisting only of small elements, if there are large hairs, these are characteristically effinate; stipe pilose only at the apex when young, or pubescent only when seen under a strong lens.

12. Long, thinly effilate hairs present at the apex of the stipe; on Quercus leaves in central Mexico. 10. M. cryptotrichus.

12. Hairs of stipe short, not effilate. 13. Lamellae fresh white; carpophore accompanied by

thin, dark rhizomorphs; trama of pileus and hyaline hyphae of stipe inamyloid or very slightly and slowly pseudoamyloid; rain forest species of the lower altitudes. 11. M. chiapasensis.

13. Lamellae fresh pale leather brown to buff-whitish; carpophores not accompanied by dark rhizomorphs; some hyphae of the pileus trama weakly pseudoamyloid, many pseudoamyloid in the apex of the stipe; subtropical and tropical-montane species (see "4" above).

8. Growing on wood. 14. On Lituqidambar in Mexico (see "9" above). 14. On roots and woody rootlets of other trees in South America;

lamellae rather broad; stipe not accompanied by dark rhizomorphs. 12. M. radicellicola.

1. Clamp connections absent in the hyphae of the carpophores. 15. Stipe only rarely rising from a black rhizomorph, usually longer than the diameter

of the pileus. 16. Odor of Allitium sativuIm.

17. Spores ellipsoid to piriform, 6.2-9 X 2.5-4.8u; hypodermial elements roughish from incrusting pigment but pigment subhyaline to pale melleous and therefore incrustation not conspicuous. 13. M. dysodes.

17. Spores mostly oblong, (6.5-)7-8.5 X 2-4,; hypodermium strongly sepia- incrusted. 14. M. atroincrustatus.

16. Odor none. 18. Basidia 4-spored; on dicotyledonous leaves.

19. Stipe pigmented. 20. Stipe indistinctly pilose, beset with thick-walled hairs of the Crinipel-

lis-type. Inodorous forms of M. atroincrustatus. 20. Stipe with dense fibrillose-furfuraceous white covering underneath the

spadiceous-sepia and somewhat shining surface of the stipe rind, this covering consisting of a tangle of mostly clavate dermatocystidia with walls 0.2-0.5 thick, few thick-walled and then wall sepia, some hyphous, at times appendiculate. Florida. 15. M. atlanticus.

19. Stipe white. 16. M. eorotula. 18. Basidia 2-spored; on monocotyledonous leaves. 17. M. defibulatus.

15. Stipe always or almost always rising from conspicuous rhizomorphs, its length about equal with the diameter of the pileus. 18. M. tomentellus.

1. Marasmius yalae 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 2.5-3.5 X. 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, regularly inserted 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-fuliginous 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-subfusiform to 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 numerous but discontinuous, 16-24 p long ventricose or hyphous but at times forked and the majority diverticulate, with irregularly distributed diverticula 1-3.5 1 long, or with apical appendages up 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 pseudoamyloid strands in the pileus, parallel and colored in the lower portion of the stipe and there weakly pseudoamyloid or nearly inamyloid, all with clamp connections and not gelatinized. Cortical layer: epicutis of pileus formed by branched-diverticulate hyphae as in M. androsaceus with 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 epicuticular elements cystidiform, elongated and diverticulate on the upper side.

On dicotyledonous sticks and wood particles fallen to the ground in Alnetum,

solitary in the montane forests. MATERIAL STUDIED. ARGENTINA. Jujuy: Lagunas de Yala, 1900-2300 m alt,

13 Feb 1966, Singer T 5131 (F), type, 11 Feb 1966, Singer T 5059 (F).

2. Marasmius polylepidis Dennis in Singer, Sydowia 18: 341. 1965. TYPE. Dennis 173 (K), from Venezuela. Pileus evenly colored between "kermanshah" and "kaffa" M&P (fuscous), dried

gray brown, not shining but glabrous, smooth at first then finely sulcate-striate in

marginal half and there eventually plicate-furrowed, hemispherical to convex, the center

umbilicate depressed or not, not umbonate, 3-4 mm broad. Lamellae paler than pileus to sordid white with white or whitish edge, intermixed with some lamellulae, about 7-10-

(14) through-lamellae, rather close, adnexed with a collarium. Stipe purplish-brown- black, shining when fresh, glabrous, not setose under a lens, insititious on the substra-

tim, not accompanied by 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 , according to Dennis to 20 X 6 u, and interspersed with fusiform cystidia; in Yungas material only fusiform basidioles seen and cystidia none; nature basidia 4-spored; cheilocystidia about 6-7 A broad, hy-

phous and often forked, some swollen to vesiculose and pedicellate, all with mainly

apical diverticula which are somewhat irregularly distributed and short, much as the

epicuticular elements of certain Marasmielli, hyaline, numerous; base of some basidia



Marasmius 67

partly pseudoamyloid at times. Hyphae of hymenophoral trama and margin of 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-hymeniform layer of elements similar to those of the edge of the lamellae, also hyaline. Hypodermium consisting of hyphae in radial arrangement, strongly incrusted by a

deep brown pigment which appears bright brown in the Melzer reagent. 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 appears to be a geographic race restricted to the southern part of the Poly-

lepis area can be collected commonly under Polylepis australis in Argentina. This race differs but slightly in having non-umbilicate pileus; it is mostly accompanied by black rhizomorphs. Characteristic material of this southern form was collected by me in Argentina: Jujuy, Lagunas de Yala, 16 Feb 1966, T 5202 (F).

3. Marasmius cyrillidis Dennis, Kew Bull. 22: 76. 1968. Fig 30. TYPE. Detnnis J 40, from Jamaica (K). Pileus cinnamon to dark reddish brown when fresh, rarely paler and about gil-

vous, tending to become wood-brownish or gray-brown when dried (fresh e g between "Kermanshah" and "kaffa" M&P in the center, the margin usually paler), smooth at first, later sulculate to long sulcate, glabrous, convex, sometimes with applanate apex but neither umbilicate nor papillate, 3-7.5 mm broad, mostly only up to 4 mm broad. Lamellae pale leather brown to buff-whitish (not pure white except at the edge when fresh), medium distant, 12-14 through-lammellae intermixed with some lamellulae, not or scarcely intervenose, narrow, to rather broad, not collariate, broadly adnexted to adnate. Stipe deep chestnut to blackish-brown, shining when fresh, glabrous, smooth, insititious, equal or slightly tapering upwards, 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-ellipsoid or oblong-ellipsoid, to cylindric- oblong, smooth, hyaline, inamyloid. Hymenium: basidia 12.5-22 X 3.5-6.3 p, 4-spored, sterigmata 2-6 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 including its interior, where the hyphae are almost subhyaline and parallel with each other. Cortical layer: epicutis of the pileus of interwoven versiform elements, not in hymeniform arrangement, these often swollen to 7 p diameter and some like the cheilocystidia, most distinctly diverticulate but ir-

regularly so, fewor many here and almost all in the hypodermium distinctly incrusted by a stramineous or fuscous pigment; hyphae of the hypodermium mainly radially arranged: hairs of stipe none or inconspicuous at 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 (Quercus humboldtii, Cyrilla racemosifolia, 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 Flora Neotropica

Andr6s Hidalgo 9 Jul 1969, Singer M 8356 (F). BOLIVIA. La Paz: Nor-Yungas below Mlururata, 1400 m alt, 9 Feb 1956, Singer B 1048 (LIL, BAFC).

I confused this species with M. polylepidis when first encountered, then separated it under a herbarium name 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 broader than indicated by Dennis, but young spores are often exactly as those of 1/. cyrillidis. The incrustation of the hyphae of the hypodermium is indeed less coarse than that of M. androsaceus but I have always found it present. In 1A. androsaceius the hyphae even of the rest of the pileus and hymenophoral trama are pigment-incrusted whereas in M. cyrillidis they are not.

4. Marasmius bactrosporus Singer, Sydowia 18: 334. 1965. TYPE. Singer T 3521, from Argentina. Pileus beige in youth 1 l-B-4, later 11-C-6 M&P with fuscous gray ("soapstone")

disc, glabrous, smooth, eventually sometimes more or less uneven, convex, later convex- applanate, 2.5-3.5 mm broad. Lamellae whitish, later light sordid gray ("almond, biscuit" M&P), medium close, intermixed, not regularly di- or tridymous, 12-17 through- lamellae, medium broad, adnate, sometimes with brown filbrils at the plane of attachment so that they appear collariate (but not!), eventually somewhat decurrent. Stipe fuscous, apex concolorous with the lamellae, macroscopically glabrous but hyaline- pilose under a good lens, smooth, insititious, equal, 11-16 X 0.1-0.7 mm; black rhizomorphs accompanying the stipe, thinner than stipe, concolorous with it, also extremely finely pilose but only partially so, both stipe and rhizomorph eventually glabrescent. Context subhyaline in 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 suprahilar depression, hyaline, 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 versiform and variable in size, about 11-12.5 p broad, hyaline to brownish hyaline, few smooth, most with obtuse short

(rarely reaching 2.5 u length) diverticula all over upper portion, making the edge heteromorphous. Hyphae of the pileus and hymenophoral trama hyaline, thin-walled, filamentous, 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 smaller and hyphous elements which are likewise diverticulate, hyaline or pale brownish hyaline, not hymeniform; pilosity of stipe consisting of scattered thin-walled hyaline hairs which are sub-

cylindric or subampullaceous or subulate, often with widened base, obtuse, rarely subacute, not acute, very rarely forked, not diverticulate, erect, 4-40 X 3-8.5 ,, hypodermium of pileus, a cutis of filamentous hyphae which are subparallel to parallel with 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 appearing which are homologous to hypodermium of pileus and consist of bunches of parallel or subparallel hyphae which are 2.5-8 p thick and incrusted by fuscous pigment.

On fallen dead leaves of Myrtaceae in subtropical-montane forest.



Marasmius 69

MATERIAL STUDIED. ARGENTINA. Tucuman: San Javier, ravine northwest of Ciudad Univ- ersitaria at 1000 m alt, 9 Jan 1960, Singer T 3521 (LIL), Type.

ILLUSTRATION. Singer (1965) pi 4, fig 2.

5. Marasmius leguminosarum Singer, Sydowia 18: 348. 1965. Fig 31. TYPE. Singer B 3394, from Brazil. Pileus brown ("auburn" M&P), darker in the umbilicus, paler on the margin, there

shining, dried more grayish ("suntan" or "tanbark" in the middle, "grain" towards margin, M&P), sulcate, glabrous, convex with an eccentric umbilicus, (2-)5-7 mm broad. Lamellae grayish, dried sordid isabelline-cinnamon (11 E 6 to "bran" M&P), narrow, distant (about 9 through-lamellae) adnate. Stipe black or blackish, macroscopically glabrous but 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), rather thin, inodorous.

Spores 6.5-7.5 X 3.3-3.7 ,, ellipsoid to oblong, smooth, hyaline, inamyloid. Hymen- ium: basidia 31 X 6.5 ,, cystidia none; cheilocystidia filamentous but some here and there

partly widened, sometimes moniliform or diverticulate, often ramose, hyaline. Hyphae of the pileus trama hyaline, pseudoamyloid, with clamp connections, not gelatinized. Cor- tical layers: epicutis of the pileus not hymeniform, consisting of elements rising from the hypodermium and ascendant, often interwoven, similar to the cheilocystidia; hypodermium characteristically dark incrusted by the pigment; superficial hyaline hairs both on pileus and stipe surface, on pileus rare, on stipe rather numerous, there thin, mostly acute, stiff, often incrusted.

On wood of dicotyledonous trees, mostly Leguminosae (fallen twigs, bark, etc). Known host: Bowdichia virgilioides.

MATERIAL STUDIED. 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), Type.

The eccentric stipe is characteristic. The collection from Venezuela has no spores.

6. Marasmius earlei Murrill, North American Flora 9: 258. 1915. TYPE. Earle 269, from Jamaica (NY). Pileus whitish with brownish center, in herbarium almost uniformly colored brown-

ish, glabrous, smooth, then weakly to distinctly-sulculate at least when dry, convex, then

shallowly but distinctly depressed in the center, but neither umbilicate nor papillate, 3-5.5 mm broad. Lamellae white, subclose to subdistant (10-24 through-lamellae) some forked, not intervenose, rather broad, adnate, not collariate or subcollariately separating 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, inamyloid. Cortical layer: epicutis of the pileus a weak to rather strong continuous Rameales- structure, there and in the hypodermium with an ochraceous intraparietal pigment which is not or scarcely incrusting; covering of the stipe consisting of small, thorn-like ascendant or erect subhyaline excrescences which are numerous but 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. Marasmius liquidambari Singer, sp nov Fig 32. TYPE. Singer M 8418/8420, from Mexico (F). Pileo vinaceo-brunneo dein cinnamoneo, 5-11 mm lato; lamellis albidis vel cremeis:

stipite subtiliter subpruninato-subflocculoso, rhizomorphis nigris haud adhaeso. Sporis 5.5-8.5 X 2.2-4 j. Ad folia ramosque Liquidambari styraciflui in Mexico. Typus in F conservatus est.

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 depressed in 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-lamellae and 0-10 lamellulae, narrow to rather broad, subcollariate-separating or attenuate-subfree to adnexed. Stipe chestnut to blackish in lower two thirds with the apex concolorous with the lamellae, subglabrous but under a lens finely white pruinate subflocculose but eventually somewhat glabrescent and then more or less shining, insititious, equal, 10-15 X 0.3-0.7 mm; black rhizomorphs thinner than the stipe, glabrescent or glabrous, accompanying the 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 subdiverticulate apex, hyaline, sometimes merely a continuation of the epicuticular structure; scattered pale melleous incrustations and other extracellular material often found in preparations of the hymenium. Hyphae of the pileus-trama filamentous with firm to thick (0.3-1 j,) walls, with clamp connections, not gelatinized, hyaline, some with fine granular hyaline to pale melleous incrustations, those of the regular hymenophoral trama either thin-walled or 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 strands weakly to moderately strongly pseudoamyloid, others, especially in the pileus, inamyloid. Cortical layers: epicutis of the pileus consisting of not hymeniformly arranged irregular, often cystidiform elements, most of these distinctly diverticulate (diverticula 2-11 X 0.2-1 Iu) hyaline to pale melleous in the marginal region by an intraparietal pigment, in young specimens in the center showing a very thin and intermittent, not conspicuous incrusting resinous pigment, no gelatinization; hypoldermium likewise brownish spadiceous pigmented in a manner similar to that of the epicutis, of thick-walled elements; covering 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 tapering upwards or with swellings here and there, with rounded tip, hyaline straight or bent at an angle of ? 90

On branches as well as on leaves of Liquidambar stvraciflua. 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, Guznman 3666 (ENCB).

8. Marasmius splachnoides(Fries) Fries, Epicrisis 384. 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-brownish center when fresh, sulcate, convex, with eventually shallowly depressed or subumbonate center, 5-7 mm broad. Lamellae white, close to subdistant (10-11 through-lamellae, numerous lamellulae, often didymous), narrow to rather broad, not intervenose but at times anastomosing or forked, adnexed to adnate, with a tendency to separate from the apex of the stipe and becoming subcollariate. 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 continuous (although occasionally interrupted by basidia) fringe, much like the elements of the epicutis of the pileus, strongly and densely diverticulate. Hyphae hyaline and inamyloid in the pileus, pseudoamyloid in the stipe, with clamp connections, mainly filamentous. Cortical layers: hypodermium with thin shee-like, deep brown incrustations 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 numerous but not crowded, versiform hairs 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-hyaline thin (0.3 u) to slightly thickened (0.6 u) wall.

On fallen leaves of Quercus. MATERIAL STUDIED. COLUMBIA. Cundinanarca: Cerro de las Mercedes, Mun. Boyaca,

18 Aug 1964, Gurznwn 4630 (ENCB). FRANCE. Yerres (S&O), Sep 1933, Romagnesi(LIL). In the sense of Kihner and Romagnesi, this species appears to occur mainly on

oak leaves both in the temperate zone of the Northern Hemisphere and in the tropics south to Colombia (southern limit of Quercus).

Fries, who had seen only dried material when describing the species, obviously gave an incomplete but too inconclusive description which was emended by Quelet (1886): the stipe was described as rufous, which I have not observed on the French material nor on the Colombian one, but this color may be present in younger, fresher specimens. The description given above is exclusively on the basis of neotropical material.

9. Marasmius hakgalensis Petch, Trans. Brit. Mycol. Soc. 31: 35. 1947. Marasmius hyalinotrichus Singer, 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 marginal region, glabrous, smooth, soon slightly to moderately strongly sulcate, convex, with weakly umbonate or umbilicate center in the flattened to shallowly depressed central third, 2-6 mm broad. Lamellae pale cream to almost concolorous with the margin of the pileus, not pure white in fresh condition, dried buff to light avellaneous, rather narrow to rather broad, close to distant (-14 through-lamellae and inserted with a few lamellulae) often some forked or even anastomosing but not intervenose, with entire not discolorous or with whitish edge, entire, horizontal adnate or collariate adnate; Stipe young colored like the center of the pileus, gradually becoming 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 rhizomorphs accompanying the carpophores, also pilose or mycelioid, usually thinner at the stipe, long-creeping. Context thin, inodorous.



72 Flora Neotropica

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, versiform but often clavate, with diverticula (which are sometimes fasciculate on a finger-like appendage) mostly several to 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 numerous cheilocystidia. Hyphae of the pileus-trama and the regualr hymenophoral trama hyaline, thin-walled, rather loosely arranged but not gelatinized, inamyloid or at times vaguely pseudoamyloid, hyphae of the stipe parallel, here many thick-walled and melleous in KOH, pseudoamyloid or merely vaguely pseudoamyloid, all with clamp connections (but often numerous secondary septa present). Cortical layers: epicutis of the pileus not hymeniform, its elements cylindric to vesiculose, hyaline or somewhat fuscous in their lower portion laterally and/or apically diverticulate with diverticula 2-6.5 X 0.8-1.8 1, many of them pigmented by an

incrusting pigment but this incrustation thin, resinous-sheet-like, fuscous or fuscidulous; the same slight to rather strong pigment incrustation also on many elements of the hypodermium, elements elongated but often branched; covering of the stipe characteristic 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 + pseudoamyloid wall and acute apex, entire and hyaline, fragile but rigid, incrusted by 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-filamentous or gradually tapering to an obtuse tip, thick-walled, hyaline, the inner wall-layer pseudoamyloid, the outer wall layer and the interior inamyloid, sometimes incrusted by hyaline, amorphous material, generally of the appearance and structure of Crinipellis hairs, wall 1-4 thick but sometimes much thinner, and less pseudoamyloid in the apical portion, more numerous than 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-montane forests, from Mexico to Brazil, also in tropical Africa and Asia.

MATERIAL STUDIED. MEXICO. Morelos: NW of Tepoztlan at 2,200 m alt, 27 Jun 1969, Singer M 8208 (F). BOLIVIA. Beni: Vaca Diez: Guayaramerfn, 9 Mar 1956, Singer B 1 776/III (LIL); 9 Mar 1956, Singer B 1779 (LIL), type of M. hyalotrichus; 17 Mar 1956, Singer B 2094 (LIL). BRAZIL. Pernambuco: Camaragibe, 6 Jul 1960, Singer B 3117 (BAFC); B3111 (BAFC); Paraibo: Joao Pessoa, 11 Mar 1960, Singer B 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 sometimes difficult to distinguish from the preceding species unless the hairs of the stipe are studied. These are

always entire, smooth in M. hakgalensis; also, the spores are slightly narrower in the latter.

ILLUSTRATIONS. Singer (1965) pl 6, fig 8-9; Petch (1947), fig 12.

10. Marasmius cryptotrichus Singer, 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 conservatus est.

Pileus fuscous gray, glabrous, sulcate over about half of the radius, convex, obtuse, not umbilicate, 5-6 mm broad. Lamellae cream white, close (about 14 through-



Marasmius 73

lamellae, some lamellulae), with entire edge and not intervenose, adnexed, not collariate. 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 tapering upwards, insititious, about 40 X 0.3 mm; dark rhizomorphs accompanying 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-trama and the regular hymenophoral trama mostly thin-walled, not gelatinized, hyaline, inamyloid, with clamp connections, in the stipe parallel, some with thin, some thick (1.3 p) or intermediate wall, multiseptate, varying between hyaline and brown, inamyloid, the brown ones very indistinctly pseudoamyloid. Cortical layers: epicutis of the pileus consisting of not hymeniformly ar-

arranged versiform elements, these mostly erect and often cylindric to clavate, inamyloid, 7-10 X 4-7 p (main body), sometimes branching, some hyphous, all or most diverticulate, the hyphous ones often arising from the clavate ones, hyaline, setulae or diverticular equal or more often tapering upwards, to a narrow but obtuse tips, 3-5.2 X 1-(1.2) L; hypodermium contrasting with the epicutis in its pigmented elements which are

rather irregular in shape, mainly radially-elongated with hyaline to brownish wall and thinly incrusted by an inconspicuous sheet-like spadiceous pigment-incrustation; covering 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 broadened base- portion up to 25 , long, the apex thin and piliform, 2-3 A broad, 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, corresponding to the basal portion of the long hairs, rounded obtuse above and not effilate, also inamyloid, all entire.

On dead leaves of Quercus, solitary. Mexico. MATERIAL STUDIED. MEXICO. Morelos: NW of Tepoztlan, 27 Jun 1969. Singer M

8250 (F) type. This differs from other species growing on oak leaves in the characteristics of the

hairs of the stipe which are not thick-walled nor acute (as in M hakgalensis) nor long and diverticulate (as inM. splachnoides) but at least many of them characterized by the thin-walled, effilate apex. The spores are rather small in comparison with those of M splachnoides and relatively broader than those of M. hakgalensis.

Marasmius cryptotrichus differs from the following species (which is closest) in the presence of effilate hairs on the stipe, the more fuscous-gray rather than tan, ochraceous brown color of the pileus and the less diversified cheilocystidia as well as by the host.

11. Marasmius chiapasensis Singer, sp nov Fig 35. TYPE. Singer M 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 plerumque fibulatis, in hypodermio fortiter pigmento incrustatis. Ad folia delapsa dicotyledonea in Mexico. Typus in F conservatus est.

Pileus (ochraceous)-brown, with paler marginal region (fresh about "oak" in center, "papyrus" toward margin, M&P), dried varying from pale tan to almost blackish brown (in center), convex, eventually and on drying with depressed center and dried sometimes with a slight umbo in the depression, somewhat rugulose and obscurely



74 Flora Neotropica

sulcate, 4-5.5 mm broad. Lamellae white, subclose, with 11-16 through-lamellae and some lamellulae, medium broad, adnate or adnexed, not collariate. Stipe dark chestnut to fuliginous and somewhat shining, setiform, appearing glabrous but with a fine

pallid pruina or pubescence when seen dried under a lens, insititious, stuffed, equal, 16-19 X 0.1-0.3 mm; accompanied by thin, glabrous, black rhizomorphs. Context thin, inodorous.

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 subapical setulae, main body about 12 X 7 j, setulae 2-6 X 1-2 ,u, (3) fusoid-mucronate to ampullaceous 20330.5 X 5.5-7 u, rather few; often one or another type missing. Hyphae of the regular hymenophoral trama and 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 parallel, 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 diverticulated versiform elements, none or few of these cystidioid, hyaline, or with very scattered pigment incrustations but in the hypodermium with 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 subclavate or subvesiculose, with broadly rounded-obtuse tip, without effilate

appendages or setulae anywhere, with stramineous-ochraceous, pale melleous. or sub-

hyaline thin to sometimes somewhat thickened (0.2-1 1u, most frequently 0.4 p), inamyloid wall; acute, incrusted and long, thick-walled hairs 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. Marasmius radicellicola Singer, 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 subtiliter pruinato vel subpubescente; rhizomorphis nullis visis; odore nullo. Sporis 4-7.3 X 3-3.7 p; hyphis fibuligeris, in hypodermio spadiceo vel sepiaceo pigmenti incrustatione causa. - Ad radices arborum in Brasilia. Typus in F conservatus est.

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, eventually often slightly depressed in the center, 2.5-3.5 mm broad. Lamellae dirty ocher (not white when fresh or dried), relatively very broad, subclose to mostly subdistant (about 7-12 through-lamellae and some lamellulae), rounded-adnate to emarginate- adnexed. Stipe black, at first with chestnut to white apex, setose, macroscopically glabrous but under a lens finely pruinate to subpubescent, shining, smooth, insititious, 10-19 X 0.1-0.2 mm; black rhizomorphs not accompanying the 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-spored at times; basidioles fusoid; cystidia none; cheilocystidia versiform, many diverticulate like the diverticulate elements of the

epicutis. Hyphae of the pileus-trama thin-walled, with clamp connections, hyaline, filamentous, inanyloid to very slightly pseudoamyloid in places; hyphae of the stipe all parallel with 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 scattered depth and some secondary septa. Cortical layers: epicutis of

the pileus consisting of diverticulate hyphae and broom cells of various shapes, with diverticula 2-10 mostly about 3 ,uhigh, hyaline; hypodermium at least in the middle of the pileus distinctly spadiceous to sepia incrusted by sheets of pigment, its elements

irregularly interwoven; covering of the stipe consisting of numerous but mostly small (4-29 X 5-11 I, rarely longer) obtuse hairs, those of the upper portion of the stipe all thin-walled and 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-walled but otherwise similar, wall here 1-3 p thick.

On dead roots and woody rootlets of trees (Ficus?), gregarious. Brazil. MATERIAL STUDIED. BRAZIL. Pernambuco, Dois IrmSos, 14 Apr 1964, Singer B 4078

(F), type.

13. Marasmius dysodes 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 depressed in the center, 6 mm broad. Lamellae white, becoming deep brown dotted and eventually entirely dirty gray brown, rather broad, subdistant or moderately close (about 10 through-lamellae), intermixed with a few lamellulae, adnate, not collariate. Stipe black, smooth, macroscopically glabrous when fresh but with small, erect, white hairs when seen under a lens, pilose from apex to base, setose but slightly broader in the middle (0.2 mm), 19 mm long, insititious on the substratum; accompanied by blackish-brown rhizomorphs which 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 similar to the

Rotalis-type of epicuticular broom cells, diverticular colorless, some few terminal elements of edge without any diverticular Hyphae without clamp connections, inamyloid or almost so, but the inner-most wall-layer of the thick-walled elements of the context often pseudoamyloid, excepting the few connecting hyphae, the others thick-walled, subhyaline to melleous, and strictly parallel with each other. Cortical

layers: epicutis of pileus consisting of versiform elements, varying from hyphous to swollen in the manner of the cheilocystidia, diverticulate, diverticula melleous hyaline to pale melleous, 1-3.5 X 1-1.5 p; hypodermium more or less thick-filamentous, its hyphae roughish from incrusting pigment which is not conspicuous since it is very pale melleous, often rather voluminous; 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 tapering slightly towards the apex which is almost always always broadly rounded and smooth and entire, the innermost wall layer more or less

pseudoamyloid.



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. ILLUSTRATION. Singer (1965) pl 5, fig 3.

14. Marasmius atroincrustatus Singer sp nov TYPE. Singer M 8320, from Mexico. Pileo cinnamomeo-alutaceo, plerumque 3-3.5 mm lato; lamellis pallide isabelli-

nis; stipite pallide pubescente, glabrescente. rhizomorphis haud adhaeso, 17-19 mm longo; odore alliaceo. Sporis 7-8.5 X 2-4 p; hyphis defibulatis, in hypodermio pigmento sepiaceo fortiter incrustatis. Ad folia dicotyledonea delapsa in Mexico. Typus in F conservatur.

14a. Marasmius atroincrustatus var atroincrustatus TYPE. Singer M 8320, from Mexico. Pileus cinnamon buff, sulculate on margin, glabrous, convex, eventually slightly

applanate or depressed in the center, and neither papillate nor umbilicate, but sometimes with a slight umbo in the depression when 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-lamellae and 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, insititious, setiform, equal, 17-19 X 0.2-3 mm; associated with but not rising from dark rhizomorphs which are also glabrescent. Context thin, white, with a distinct garlic odor.

Spores (6.5-)7-8.5 X 2-4 ,, mostly 2.7-3.5 p broad, ellipsoid, oblong or subcylindrical, hyaline, smooth, inamyloid. Hymenium: basidia 19.5-23 X 5.5-7 ,, 4-spored; cystidia none; cheilocystidia versiform with a main body 15-24 X 3.5-7 1, diverticulate with diverticula erect 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 filamentous, not or scarcely pseudoamyloid, not gelatinized, those of the stipe parallel, subhyaline 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: epicutis of the pileus not hymeniform, its elements towards the upper surface beset with hyaline diverticular 2-7.5 X 1 u, these elements versiform and 3-4 p broad, hyaline, but some, like most of the hypodermium, incrusted by a distinct spadiceous pigment incrustation, some cystidiform and then much like the cheilocystidia; hypodermium hyphous, the broader or all elements with abundant dark pigment incrustation, with- otu clamp connections; covering of the stipe ,consisting of Crinipellis-like hairs, these 29-90 jp long and 4-7 u broad, thick-(1-2 l)walled, cylindric or tapering gradually to an obtusate tip, all entire and smooth, most of them distinctly pseudoamyloid.

On dicotyledonous, dead, fallen leaves in tropical-montane forest. Mexico. MATERIAL STUDIED. MEXICO. Oaxaca: Rancho del Cura, 7 Jul 1969, Singer M 8320

(F) type.

14b. Marasmius var inodorosus Singer, var nov Fig 36. TYPE. Singer M 8816, from Mexico. Odore nullo.



Marasmius 77

This differs from the type variety mainly in inodorous carpophores and perhaps slightly less elongated spores (6.5-7.5 X 3.5-4 j) and somewhat paler margin and stripes along the ridges of the sulci of the pileus.

On oak leaves in Mexico. MATERIAL STUDIED. MEXICO. Chiapas: 15 km W of San Cristobal de las Casas, 2400

m alt 2 Aug 1969, Singer M 8816 (F), type of variety.

15. Marasmius atlanticus Singer, sp nov Fig 37. TYPE. Singer F 347, from Florida, U.S.A. Pileo rubescente-brunneo in 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, incrustationibus inconspicuis. Ad folia quercina in Florida. Typus in F conservatus est.

Pileus reddish brown in center, white on margin, grooved-sulcate, glabrous, size and shape as in Marasmius androsaceus. Lamellae white, rather thick, somewhat irregular, close, adnate. Stipe deep spadiceous-sepia, somewhat shining under a dense white fibrillose-furfuraceous covering, 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 diverticulate or 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 regular hymenophoral trama, deep brown, parallel and pseudoamyloid in the rind of the stipe, there with thick walls. Cortical layers: epicutis of the pileus not hymeniform, a rather thin layer with terminal cells often but not consistently showing wavy outline or irregular diverticula, here and in the hypodermium some sheet-like bt., very thin and inconspicuous pigment incrustations present, but these intermittent, dark brown; covering of the stipe consisting of cystidioid hairs which are dense and often interwoven, somewhat irregular, 1244 X 6.5-13 p, fewer hyphous and 3 1 broad, subhyaline to hyaline, mostly ending in a clavate or ventricose apex which may be appendiculate by one oblique finger-like appendage, few sepia-brown and 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 Quercus laurifolia in subtropical forest. MATERIAL STUDIED. U.S.A. Florida: Highlands County, Highlands Hammock State Park,

21 Aug 1942, Singer F 347 (F), type. This differs from the other oak leaf inhabiting species in the absence of clamp

connections.

16. Marasmius eorotula Singer, sp nov Fig 38. TYPE. Singer B 6380, from Colombia. Pileo albo, 2-7.5 mm lato; lamellis albis, plus minusve collariatis; stipite albo, ad

basin interdum pallidissime stramineo, 4.5-25 mm longo; odore nullo. Sporis 6.5-8 X 25-3 1; lhyphis defibulatis. Ad folia dicotyledonea delapsa in Columbia. Typus in F conservatus est.

Pileus white, glabrous or slightly furfuraceous under a strong lens, smooth or with short-folded-furrowed margin, convex, obtuse, often with flattened or somewhat narrowly depressed center when old or dried, 2-7.5 mm broad. Lamellae white, medium close to subdistant (about 12 through-lamellae), inserted with some irregularly



78 Flora Neotropica

disposed lamellulae, horizontal, moderately broad to rather broad, not intervenose, adnexed but removed from the stipe and connected by a more or less distinct collarium leaving a radially fibrillose, narrow zone between the collarium and the apex of the stipe. Stipe white, sometimes becoming palest stramineous below, never dark at base, glabrous macroscopically but minutely flocculose or pruinate in the lower portion or all over when seen dried under a lens, insititious and equal or with a vaguely disc-like widening at the base, hollow, 4.5-25 X 0.1-0.3 mm. Context white, thin, inodorous.

Spores (5.5-)6.5-8 X 2.5-3 ,, ellipsoid to mostly oblong, hyaline, smooth, inamyloid. 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 4-16 j, clavate to vesiculose, with apical and subapical as well as often also divergent lateral diverticula or setulae, these varying from finger-like to verruculose, very variable in size but mostly 2-4 X 1 E, the main body often forked or tripartite and each fork beset with setulae, the wall thin and hyaline. Hyphae in the pileus-trama hyaline, 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. Cortical layers; epicutis of the pileus in places almost subhymeniform or in palisade but in others irregularly arranged, consisting of terminal bodies which are more or less erect ot ascendant, 14-38 X 6-27 ,, beset with numerous diverticula like those of the cheilocystidia, also often branched or appendiculate witih diverticulated branches or appendages, the appendages of the ascendant cells 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: hypodermium hyaline, of elongated horizontal hyphae inter-mixed with 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, cylindrical or ampullaceous, vesiculose or conic-obtuse, sometimes mucronate with thin or, more rarely, up to 1 thick, hyaline to pale stramineous wall 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. MATERIAL STUDIED: COLOMBIA: Valle, Buenaventura, Campamento Pulpapel 25 Apr

1968, Singer B 6380 (F), type. San Joaquin, 22 Apr 1968, Singer B 6290 (F); 19 Apr 1968, Singer B 6234 (F).

This has the appearance of a representative of the section Marasmius or of Mara- smiellus sect Defibulati, but clearly differs from the former innon-hymeniform epicutis and the irregular and variable shape of the setulae, and from the latter in the distinctly pseudoamyloid hyphae of the stipe. It differs from all other species of section Androsacei in the almost complete absence of pigment, in the carpophores.

17. Marasmius defibulatus 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 rather distant (10-12 through- lamellae, intermixed with shorter ones), narrow to broad, mostly broad, collariately adnexed to a narrow and 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, accompanied by but not attached to black rhizomorphs.

Spores 6-9 X 3.2-4.5 ,, mostly 7.8-8 X 4,, ellipsoid, smooth, hyaline, inamyloid. Hymenium: basidia 18-26 X 4.5-6 M, 2-spored; basidioles fusoid; cystidia none; cheilocystidia ventricose to subvesiculose like the epicuticular broom cells but hyaline. Hyphae hyaline, not gelatinized, regular in the hymenophoral trama, there 2-3.7 ,lbroad, inamyloid. Cortical layers: epicutis of the pileus not hymeniform, consisting of appressed, multiseptate hyphal elements, the terminal ones mostly cystidioid and densely and regularly beset with setulae, main body vesiculose to ventricose, often constricted in the middle but also hyphous, setulae also on the hyphal ends, directed mainly upwards, 2-3 X 0.8-1.2 ,, all hyaline, many terminal cells ascendant or erect, all inamyloid; covering of the stipe consisting of hairs 18-530 X 3.5-10 ,, hyaline, tapering upwards gradually but with rounded tip, entire, thick-walled (wall 0.7-4 ,).

On moist, decaying palm fronds. MATERIAL STUDIED. SURINAM. Coppename River, Calaback Creek, 13 Nov 1923, D.

H. Linder (FH), type. ILLUSTRATION. Singer (1958a), fig 38.

18. Marasmius tomentellus 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, adnexed. Stipe badious to blackish, pubescent, curved, rising from a common black rhizomorphs, solid, subequal with slightly to distinctly narrowed apex, 1-4 X 0.2-0.6 mm; rhizomorphs black, long, as thick or thicker than the diameter of the stipe.

Spores 11-11.5 X 4-5 ,, cylindric to oblong with mostly recurved basal 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 pseudoamyloid rind-zone of the stipe, without clamp connections, hyaline except for the hypodermial zone of the pileus and the rind-zone of the stipe, which are pigment-incrusted, not gelatinized, those of the pileus radially arranged and those of the stipe parallel. Cortical layers: epicutis of the pileus consisting of irregularly arranged, irregularly diverticulated elements which are hyaline to subhyaline, forming a very thin layer; liypodermium very 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 ,u thick) and seta-like hairs 50-125 X 5-7 ,, with hyaline, thick (1 ,) pseudoamyloid wall.

On heaps of dead twigs in accumulated litter. Southern U.S.A. and West Indies. MATERIAL STUDIED. U.S.A. Louisiana: La Salle parish, May 1936, Frank Smith, 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 epicuticular elements in chains, often some of them smooth, (but never all of them); spores usually medium-sized; lamellae free to adnate, not or

incompletely collariate, rarely absent; 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. Pileus rough or spiny from chains of thick-walled, often ornamented cells, accompanied by single broom cells. 2. Spores 8.7-9.3 X 3.5-4.3u1. On Magnolia in Florida. 22. M. magnoliae. 2. Spores 6-8.5 X 3.2-4u. On various monocotyledonous and dicotyledonous branches,

petioles and leaf blades in tropical America and Africa south of the Equator. 23. A. echinosphaerus.

1. Pileus smooth or merely radially sulcate, not with chained cells, only with broom cells of the Rotalis-type, some of these occasionally smooth. 3. Lamellae absent or hymenial surface radially veined. 21. M. sphaerodermus. 3. Lamellae distinct, 7-14 through-lamellae.

4. Spores more than 3.3, broad, at least many of them. 5. Pileus brownish-yellow to yellow-brown, 1-7 mm broad; lamellae not pure

white when fresh; stipe macroscopically not glabrous (see note under M. ilicis: M. corbariensis).

5. Pileus smaller (1-3.5 mm) and differently colored. 6. Pileus brownish white, drying fuscous-brown (see note under M. ilicis: M.

exustus). 6. Pileus with an orange-brown to chestnut tone. 19. M. ilicis.

4. Spores up 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 indicated above or more sepia-ferruginous to

deep ferruginous or grayish-badious; 4-8 through-lamellae present (see note under M. crescentiae: M. capillipes).

19. Marasmius ilicis Singer, Lilloa 26: 142. 1953. Marasmius sphaerodermus var major Spegazzini, Anal. Soc. Ci Argentina 10: 163. 1881. TYPE. Singer B 88/88a, from Brasil. Pileus orange brown to chestnut brown ("spice" 13-B-12 M&P to deeper red-

dish brown) with pallid to light saffron orange (almost "saffrony" i e paler than 10- K-8) margin between 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 campanulate but tending to have truncate or at least somewhat applanate disc, glabrous, 1-3.5 mm broad. Lamellae white with white edge, seven to ten, in mature caps also some few lamellulae occasionally present, neither 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 extremely thin, white, unchanging inodorous.

Spores 6.7-9.5 X 3.5-5 ,, ellipsoid to ellipsoid-fusoid, thin-walled, hyaline, smooth, often granular inside, inamyloid. Hymenium: basidia 22 X 7.5 j,; basidioles fusoid; cystidioles and cystidia fusoid like those of the Epiphylli, hyaline, "empty," with an ampullaceous to subcapitate apex, smooth, projecting 20-27 X 4-7 A; cheilocystidia scattered and similar to epicuticular broom 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. Hymenophoral trama regular, made up by similar hy-



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. Cortical layers: epicutis 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-like outgrowths up to 2.5 pprojecting; dermatocystidia like the cystidia of the hymenium also present in most collections, these hyaline, e g 23 X 8 ,, apex 7.2 X 1.8 ,; superficial hyphae of stipe often distantly diverticulate by spinose diverticula.

On dead dicotyledonous leaves, petioles, stems, most frequently on fallen coriaceous leaves and petioles of Ilex and other trees and shrubs in tropical, subtropical and marginal (gallery) forests. Known hosts: Ilex, Ligustrum, Persea, Eryngium.

MATERIAL STUDIED. VENEZUELA. Sucre: Along Rio Media, 15 Jul 1972 Dumont et al VE 5124 (NY). BOLIVIA. La Paz: Nor-Yungas, Rio Llolosa, 31 Jan 1956, Singer B 758 (LIL). BRAZIL. Rio Grande do Sul: Taimbesinho 5 Nov 1951, Singer B 88, 88a (LIL, MICH), type. ARGENTINA. Buenos Aires: Parque Pereyra 20 Jun 1962, Horak, comm. Singer S 201 (BAFC).

The Venezuelan collection, the northernmost seen, is the only one that grew on herbaceous stems; it differs also by slightly broader spores and less developed cystidia. These small differences do not seem to justify a special taxon.

I have included in the key two related extralimital species: Marasmius corbariensis

Roumeguere (Marasnzius hygrometricus Briganti) and Marasmius exustus Berkeley & Curtis (Fig 40).

Since Olea is planted in some regions of the neotropics, it is quite possible that MIarasmlius corbariensis has 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 discovered on the Bonin Islands; the

type (FH) is much like Mlarasmius ilicis microscopically (spores 5.5-9.2 X 3-4.5 p). ILLUSTRATION. Singer (1965) pl 7, fig 14.

20. Marasmius crescentiae Murrill, North American Flora 9: 259. 1915. Fig 40. TYPE. Earle 184, from Cuba. Pileus ochraceous to ochraceous brown, center deep ochraceous brown, dried

rusty ochraceous brown with dark brown center, long sulcate, glabrous, campanulate- convex, then convex, obtuse, not depressed and not papillate, 2-5 mm broad. Lamellae cream-white then pale dirty-cream, the edge only in the marginal zone narrowly brown- marginate or 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, macroscopically glabrous, setiform, hollow, equal and insititious with the base

slightly lobed and volva-like widened, 10-40 X 0.1-0.2 mm. Context thin, inodorous.

Spores about 6.5 X 2.7,u, oblong, hyaline, smooth, inamyloid. Hymenium: basidia clavate; basidioles fusoid; cystidia on both edges and sides numerous, 22-30 X 7-7 .5 ,, fusoid-ventricose below, ampullaceous with a rather narrow obtuse apex which is sometimes subclaviculate but mostly cylindric or gradually attenuated 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, stramineous to chestnut brown above and there thick (1-1.5 1) walled,



82 Flora Neotropica

the upper three quarters or two thirds covered with rod-like setulae (Rotalis-type). Hyphae of pileus and hymenophoral trama 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 subhyaline hyphae of the core thin-walled to moderately thick- walled, inamyloid. Cortical layers: epicutis of the pileus hymeniform, consisting of broom cells much like cheilocystidia, some hyaline, some chestnut brown to stramineous, thick-walled above if pigmented; there are also some smooth, thick-walled, strongly pigmented bodies, similar to the broom cells in shape but rising from the hypodermium; they do not reach the level of the epicutis and remain somewhat imbedded; covering of the stipe: in the middle of the stipe of an adult specimen consisting merely of some external hyphae which are nodulose-verruculose, some inconspicuous 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, Quercus.

MATERIAL STUDIED. MEXICO: Morelos, NW of Tepoztlan, 2200 m alt, 27 Jun 1969, Singer M 8251 (F). CUBA. 5 miles E of Santiago de las Vegas, 1 Sep 1904, Earle 184 (NY) type.

Another narrow-spored species 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 subtropical belts, introduced into gardens or parks. It was indicated for Chile by Spegazzini but his descriptive data and illustration refer to Marasmnius ilicis.

21. Marasmius sphaerodermus Spegazzini, Anal. Soc. Ci. Argentina 9: 163. 1881. Fig 41.

TYPE. Spegazzini Herbarium, La Plata, from Argentina (partly destroyed). Pileus orange buff to brownish orange ("Saratoga" M&P), scarcely changing when

dried, glabrous, smooth, slightly granular, hemispherical, subumbonate, 0.3-0.7 mm broad. Lamellae none or replaced by few radial veins, hymenial surface cream white. Stipe maroon, chestnut or brown-black at first with cream-white, later with paler apex, eventually unicolorous dark, macroscopically glabrous, 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 claviculate and 2-3 , wide, hyaline, with obtuse tip; there are also, very scattered, some broom cells in the hymenium, much like the epicuticular broom 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 hymeniform arrangement, varying from hyaline to deep brown, many melleous-ochraceous, darkest usually in the upper portion, the pigment being concentrated in the thick portion 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 dermatocystidia occasionally found among



Marasmius 83

the broom cells, the dermatocystidia much like the hymenial cystidia; covering of the

stipe consisting of scattered dermatocystidia, much like those of the hymenium but much smaller, gradually disappearing in old specimens.

On dead leaves and twigs. Known hosts: Salix humrboldtiana, Spathodea campanulata

MATERIAL STUDIED. HAWAII. Manoa Valley, Oahu, Jan 1969, R. D. Goos (F). AR- GENTINA. Buenos Aires: La Plata River (no more carpophores present in the type envelope), (LPS),type. Tierra del Fuego: Ushuaia, authentic material but only some stipes left (LPS).

This species is evidently of wide distribution but has been rarely collected, probably because of its small size. However, Spegazzini's original description (aside from the spore measurements which are obviously wrong) fits the Hawaiian material so well and the lack of lamellae is so characteristic for this one species in section Hygrometrici that I have no doubt but that they are identical with each other.

22. Marasmius magnoliae Singer, Mycologia 37: 435. 1945. TYPE. Singer F 1906, from Florida, U.S.A. Pileus deep brown, then "Amber Brown" Ridgway, with the depressions of the

radiately sulcate-rimose margin pale buff, eventually somewhat pallescent and the whole marginal zone "Clay Color" (R) pilose when mature, eventually somewhat glabrescent, the smooth disc about 0.5 mm broad, the extreme margin fimbriate-ciliate at first, hemispheric when convex, flattened eventually and becoming subumbilicate or with a slight depression in the middle of which there may be a small papilla, up to 5.5 mm broad. Lamellae white, distant, entire, equal, moderately broad (1 mm), subfree. Stipe blackish brown, macroscopically subglabrous but at least partially subfibrillose when seen under a lens, opaque, rarely slightly shining, insititious, more or less flexuous, slightly tapering at the apex, 10-40 X 0.2-0.5 mm. Context white, very thin, inodorous.

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; cheilocystidia fusoid, about 4-7 v broad, acute, smooth, entire, hyaline. Hyphae of the pileus- trama hyaline, filamentous, inamyloid, with clamp connections. Cortical layers: epicutis of the pileus consisting of short, vesiculose elements seriate in chains forming bristle-like hairs (the ornamentation of the pileus surface) these elements beset with conical to rod-shaped diverticula 2.5 Mhigh; among these "hairs" there are also ascendant clavicular elements also seriate to form long chains and arising from each others lower-apical side or forming a chain of normal filamentous hyphae, rarely the hairs consisting of smooth members of free erect hyphal chains, the individual elements in all these cases 1140 C 7-17v, the shortest ones e g 27 X 23 ,, all thick-walled, most with brown walls, the terminal members often with blackish brown apex.

On the petioles of fallen leaves of Magnolia grandiflora, very rarely also on other parts of the tree, very gregarious in "bay heads," fruiting in spring.

MATERIAL STUDIED U.S.A. Florida: Alachua County, Gainesville 11 May 1943, Singer F 1906 (FH), type; Newnan's Lake, 12 May 1943, Singer F 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-Levecq 169, from Zaire.



84 Flora Neotropica

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-spinulose all 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-subfree to free. Stipe concolorous with the

pileus below but at first with whitish apex, later and dried almost uniformly chestnut brown, macroscopically glabrous but finely echinulate-flocculose-rough when 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 , ampullaceous, lanceolate or acicular, subacute, acute, or somewhat obtusate, hyaline, thin- walled, often or rarely covered by a finely granular hyaline mass, (2) broom cells of the Rotalis-type, 10-34 X 4-9 p, vesiculose to subvesiculose, beset with divergent setulae 0.5-3 p projecting, melleous-hyaline to brown above, mostly hyaline below, wall 0.2-0.5 p thick. Hyphae of the pileus-trama hyaline, 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 changing at 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 hymeniform layer composed of four kinds of elements (a) broom cells like the cystidia of

type (2), above these reaching a wall diameter at their apex of up to 5 P, often broadly clavate ventricose of subvesiculose and more pigmented (these dark, thick-walled, even-

ly distributed among the others; (b) similar elements agglutinated in parallel rows con-

sisting of seriate catenulate broom cells and making up the ornamentation of the pileus, in the marginal zone 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 terminal cells, many of them without setulae or merely rough; (c) occasional dermato-

cystidia corresponding to the cystidia of type (1) or slightly smaller: (d) broom cells of the Siccus-type or entire, 12-20 X 5 p, often ramose-forked into branches about 2 P

broad, or simple, with obtuse tips, main body cylindric to slightly ventricose, chestnut colored, scarce and sometimes wanting; hypodermium a 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 similar to the epicutis elements of type (a) and (d), the latter scarce and of some versiform rough to subsmooth brown, thick-walled elements 18-23 X 3-5 M.

On fallen branches, sticks, leaves and petioles of dicotyledonous trees. MATERIAL STUDIED. ARGENTINA. Jujuy: Posta de Lozano, 1500 m alt, 21 Feb 1966,

Singer T 5429 (F). ZAIRE. Haut-Katanga, Kipushi, Kipopo, 1250 m alt, Schmitz-Levecq 169 (BR) type.

ILLUSTRATION. Singer (1964b), fig 14, E-H. The African Marasmius kroumirensis (Patouillard) Saccardo & Sydow has less

ornamented pileus, adnate and fewer lamellae, broadly umbonate pileus and shorter stipe, according to Pegler's (1966) type analysis.

However, I have found a fungus in Ecuador which is somewhat intermediate between M. kroumirensis and M. echinosphaerus. The material is too scanty to determine



Marasmius 85

whether it belongs to one or another of these species or whether it represents a third form. It would differ from M. echinosphaerus in being smaller, echinulate-spinulose only at the apex of the pileus and the spores perhaps being somewhat narrower; it would differ from Al. kroumirensis in being echinulate on the pileus, having a longer stipe and from both perhaps in the ornamentation of the stipe-hyphae and the occasional insertion on black rhizomorphs. Description see below.

23A. ?Marasmius kroumirensis (Patouillard) Saccardo & Sydow, Sylloge Fungorum 14: 105. 1899.

Pileus brown, in apical portion brown echinulate, otherwise glabrous and often glabrescent, sulcate, campanulate, obtuse, about 1 mm broad and high. Lamellae white, distant (about 8-11 through-lamellae), with partially eventually brownish edges, adnate. Stipe black, glabrous, insititious on rhizomorphs or directly on the leaf without a ring or broadening, 5-18 X 0.1-0.2 mm; black rhizomorphs about 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 numerous and much like the epicuticular broom cells. Hyphae inamyloid with clamp connections, stramineous in the stipe but more brown in the rind-zone, all parallel. Cortical layers: epicutis of the pileus in the marginal zone 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, projecting 0.5-1.2 p, subhyaline as in the cheilo-

cystidia or concolorous with main body; covering of the stipe none seen (perhaps fugacious) but the outermost hyphae beset with numerous setulae like those of the epicuticular broom cells; no broom cells proper seen.

On leaves and leaf petioles of a dicotyledonous tree in tropical forest, gregarious. MATERIAL STUDIED. ECUADOR. Tungurahua, Rib Topo, at about 1450 m alt, 30 Apr

1973, Singer B 7208 (F).

Marasmius sect Leveilleani Singer, Bull. Jard. Bot. Bruxelles 34: 362. 1964. TYPE SPECIES. Marasmius leveilleanus (Berkeley) Patouillard. Pileus medium sized to rather large (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. Marasmius leveilleanus (Berkeley) Patouillard, Bull. Soc. Mycol. France 33: 55. 1917. Fig 42.

Heliomlyces leveilleanus Berkeley, London Jour. Bot. 6: 490. 1847. Maras:mius umbraculum Berkeley & Broome, Jour. Linn. Soc. 14: 36. 1873. TYPE. Gardner, from Ceylon (k) (not studied). Pileus a deep and bright chestnut-rust color, 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-campanulate then applanate around a distinct umbo but eventually depressed in the center and subumbonate, 7-39 mm broad. Lamellae white to pale cream with not discolorous edge, sometimes



86 Flora Neotropica

anastomosing, mostly intervenose when quite mature, subclose to (mostly) subdistant, narrow then medium broad, with few lamellulae and sometimes all entire, narrowly adnexed to remote, free, not collariate. Stipe bister to chestnut, shining, hollow, glabrous, 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 epicuticular broom 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-trama parallel or subparallel, inamyloid or pseudoamyloid, with clamp connections, with thin to moderately thick wall (0.2-0.4 p) of variable diameter (some inflated ones to 12 , broad but not dimorphic). Cortical layers: epicutis of the pileus consist-

ing of broom cells in hymeniform arrangement but many cells smooth and entire, ventricose to claviform, fewer cylindrical, often with eccentric bulge or central mucro, rarely forked, with thin to moderately thick wall, with apical or subapical finger-like

appendages which are straight and erect to somewhat oblique, 2-10 X 1.7-3-(5)Pu, entire cell hyaline to stramineous or 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. Pantropical but rare in the neotropics.

MATERIAL STUDIED. MEXICO. Veracruz, 3 km S of Montepio, 22 Jun 1969, Singer M1 8137 (F). ZAIRE. Yangambi, J. Louis 11010 (BR).

The type was studied by Patouillard and is obviously identical with the collections described above.

ILLUSTRATION. Singer (1946b), fig 15, A-C.

Marasmius sect Scotophysini Singer, Sydowia 18: 342. 1965. TYPE SPECIES. Marasmius scotophysinus Singer. Pileus small, well pigmented, with hymeniform epicutis consisting of smooth, glo-

bose elements which are pedicellate; spores oblong to subcylindric, medium-sized; lamellae adnexed to adnate, not collariate, minutely white fringed from corncob-shaped cheil-

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 hairs.

25. Marasmius scotophysinus 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, gradually widening 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-like diverticulate in its upper portion, diverticula short and dense,



Marasmius 87

obtuse, about 1.5 X 1.5 p, both main body and diverticula hyaline, 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 radially arranged, otherwise similar, also inamyloid but having pseudoamyloid contents at times, opaque and long, often irregularly branched; 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 intraparietal pigment, globose part (without pedicel) 6-14 I in diameter, pedicel up to 13 , long, these elements in hymeniform arrangement with the contents partly pseudoamyloid; hypodermium consisting of filamentous hyphae with membrana pigment.

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 Hygrometrici in the smooth epicuticular elements,

from the Epiphylli in the pigment and the absence of pleurocystidia, from Pararotulae in inamyloid hyphae and in non-collariate lamellae, 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 therefore justified to erect a special section for this species even though the section has thus far only a single representative.

ILLUSTRATION. Singer (1965), pl 6, fig 11.

Marasmius sect 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 diverticulate elements and/or de- nnatocystidia (in the latter case dermatocystidia neither pseudoamyloid nor capitate) without long, bristle-like hairs; spores narrowly cylindric 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, epicuticular hyphae always smooth); stipe central or eccentric, attached 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 similar to fusoid basidioles), mostly fusoid to ampullaceous, rarely vesiculose, generally thin- walled and hyaline, but may have pigmented resinous incrustation; stipe with conspicuous hairs or dermatocystidia.

Marasmius sect Epiphylli subsect Epiphyllini Singer, Sydowia 18: 134. 1965. TYPE SPECIES. A1. epiphyllus (Persoon ex Fries) Fries. Tramal hyphae at least in the apex of the stipe inamyloid, epicuticular cells

weakly diverticulate or smooth; hymenophore veined radially, merulioid or lamellate, sometimes absent.

Key to the Species of Subsection Epiphyllini

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 vesiculose.



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. Marasmius euosmus Singer, Sydowia 12: 69. 1958. TYPE. Martin & Welden 8702, from Panama. Pileus white, glabrous, convex, then irregularly flattened, membranous, 3-8.5 mm

broad. Lamellae white, narrow, with obtuse edges, almost venose, with anastomosing cross-veins as high as the through-lamellae forming a poroid-merulioid hymenophore, adnexed. Spore print white. Stipe white, long-filiform, pruinate, equal to slightly tapering upwards, insititious, 8-13 X 0.3-0.5 mnm. Context white, thin, odor of flowers, pleasant.

Spores 6.6-4.3 X 2-2.2 u, hyaline, cylindrical, smooth. Hymenium: cystidia vesiculose or ventricose or clavate, hyaline, not refringent. Hyphae: hymenophoral trama regular, hyaline, all hyphae hyaline with clamp connections, inamyloid. Cortical layers: epicutis of pileus cellular, consisting of smooth, hyaline, subisodiametric or 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. Marasmius martini 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. Lamellae none; 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 cartilaginous 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, hyaline, thin-walled, inamyloid. Hymenium: basidia 30-35 X 7 , 4-spored; cystidia 30- 40 X ? 9 ,, ventricose-cylindrical or apiculate, hyaline, little or not projecting, not conspicuous, smooth. Hyphae filamentous, hyaline, inamyloid, with clamp connections. 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. Marasmius carpenterianus Singer, Sydowia 12: 70. 1958. TYPE. Carpenter 136, from Peru.

5 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. Lamellae white, rather distant (10-12 through lamellae, of those some simple, the others simply or doubly forked), with some lamellulae, rather narrow but not venose except in immature state, broadly adnate. Stipe brown, at first with white apex, setose, insititious under binocular lens pruinose, increasingly pruinose-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 suprahilar applanation or 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, ampullaceous with the widest diameter in the middle, hyaline, smooth, thin-walled, rather numerous. Hyphae: subhymenium appearing subcellular. Trama of pileus of hyphae which are parallel with each other, slightly thick-walled, not metachromatic in cresyl blue mounts, inamyloid, without clamp connections. Cortical layers: epicitis of pileus consisting of subisodiametric or isodiametric cells of various diameters, e g 1 1.5-16 X 6.5-11.5 u, smooth, thin-walled; surface of stipe with dermatocystidia which are hyaline, or more often melleous, with moderately thin walls, very versiform, usually rounded above, ampullaceous or 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: Tingo Maria, Cueva de los Pavos 700 m alt, J. B.

Carpelter 136 (LIL), type. MEXICO. Veracruz: Estaci6n Biologica de las Tuxtlas, 21 Jun 1969, Singer M 8115 (F).

ILLUSTRATION. Singer (1958a), fig 2.

29. Marasmius munyozii Singer, Beih. Nova Hedwigia 29: 100. 1969. TYPE. Singer M 6517, from Chile. Pileus white, with a flush or very pale cinnamon-ochraceous in the center, short

and slightly striate at the margin, glabrous, convex, 5-5.5 mm broad. Lamellae white, 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 heteromorphous like the epicuticular cells. Hyphae filamentous, with clamp connections, 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 rather dense, 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 , diameter) and/or in the lower portion lobed-appendiculate with 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

epicuticular elements, the European Marasamius epiphylloides Rea, by its much smaller



90 Flora Neotropica

spores and habitat on grasses rather than Hedera. It is a somewhat marginal species in a vegetation of predominantly southern origin, but comes from the subtropical zone of North Chile.

Marasmius sect Epiphylli subsect Eufoliatini Singer, subsect nov TYPE SPECIES. M. eufoliatus Klhner. Hyphis partim manifeste pseudoamyloideis, fibulatis aut defibulatis. Hyphae of the carpophore or at least the stipe distinctly pseudoamyloid; hyphae

with or without (in the neotropics without) clamps.

Key to the Species of Subsection Eufoliatini

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 claviform cells 12-23 X 7-14p, no dermatocystidia present; spores 2.5-3.2 u broad, long-fusoid. 31. M. caliensis.

30. Marasmius sanctixaverii Singer, Sydowia 18: 342. 1965. TYPE. Singer T 3156, from Argentina. Pileus white, glabrous, smooth, convex with depressed center, 7-8 mm broad.

Lamellae white, distant, about 12 through-lamellae, not very broad, at maturity becoming distinctly forked-anastomosing intervenose, even young ones forked but then not intervenose, adnate-subdecurrent. Stipe white at apex, chestnut brown below, macroscopically subglabrous but 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 suprahilar depression or 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 hymenophoral trama and pileus rather irregularly arranged although with axial respectively radial trend, filamentous but not parallel with each other, those of subhymenium not to weakly pseudoamyloid, those of pileus, stipe and hymenophoral trama distinctly pseudoamyloid; all hyphae without clamp connections. Cortical layers: epicutis of pileus consisting exclusively of dermatocystidia forming a hymeniform layer, many of the dermatocystidia almost globose below but only seemingly cellular since even these are mucronate and cystidioid, all hyaline, thin-walled, crowded, inamyloid, versiform, varying from ventricose-clavate to ventricose-fusoid ampullaceous or constricted, most frequently pear-shaped, the larger more cystidioid ones 23-32 X 6.8-10.5 ,, the smaller ones e g 20 X 6.8 , but with all sorts of transititons and the larger ones not distinctly or consistently projecting; Hypodermium hyphous, scarcely pseudoamyloid; surface of stipe beset with numerous dermatocystidia which form its pubescence, even more variable 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) ascending into 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 lepidopterous insect, gregarious.

MATERIAL STUDIED. ARGENTINA. Tucuman: Ciudad Universitaria at 1100 malt, Singer T 3156 (LIL) type.

ILLUSTRATION. Singer (1965), pi 6, fig 12.



Marasmius 91

31. Marasmius caliensis 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 castanescente in parte inferiore, albo vel albido ad apicem; odore nullo. Sporis 6.5-10.5 X 2.5-3.5 j,, fusiformibus; hyphis stipitis defibulatis, pseudoamyloideis; epicute hymeniformi ex elementis 12-27 X 7-14 p clavatis, vesiculosis, ventricoso-subelongatis efformata, dermatocystidiis nullis. Ad folia dicotyledonea in Columbia et Venezuela. Typus in F conservatus est.

Pileus white, smooth, later rugose or short-sulcate, glabrous, convex, 2.2-7 mm broad. Lamellae white, distant (up to 12 through-lamellae which are sometimes forked, few lamellulae) in age distinctly intervenose, narrow but not veniform, or only a few so, adnexed, adnate or subdecurrent. Stipe white or whitish above, light cinnamon

tawny to chestnut below, macroscopically subglabrous but pruinose-pubescent under a lens, curving near apex if growing on the underside of 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 numerous on edges than on sides of lamellae, 22-37 X 4-9 p, ampullaceous or fusoid with rather thin and short neck, rarely subcylindric, rarely the neck widened in the middle, hyaline, sometimes slightly incrusted (hyaline), at edge sometimes partly replaced by cells similar to those of the epicutis, especially near the margin of the pileus, all cystidia thin-walled, inamyloid. Hyphae of the pileus-trama filamentous, 2-6 pbroad, thin-walled, weakly pseudoamyloid or inamyloid, without clamp connections; hyphae of the stipe parallel with each other, hyaline at the apex and strongly pseudoamyloid, filamentous, multiseptate, without clamp connections, 2.5-5 p broad. Cortical layers; 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 dermatocystidia differentiated; covering of the

stipe consisting of dermatocystidia and short, hair-like excrescences which are 3-30 X 3-9 p, versiform, cylindric, conical, filiform, hemispherical or ventricose, sometimes bi-lobed but smooth, not appendiculate, thin-walled (wall 0.1-0.3 p) and hyaline, entire, with rounded tip, numerous.

On dicotyledonous leaves, singly but gregarious, in tropical forest. MATERIAL STUDIED. VENEZUELA. Dto Federal: 7 km South of Chichiriviche, 30

Jul 1972, Dumont et al VE 6707 (NY); between Colonia Tovar and El Tigre, 29 Jul 1972, Dumont et al VE 6555 (NY). COLOMBIA. Valle: Cali, Saladito, 15 Apr 1968, Singer B 6149 (F), type.

Marasmius sect Marasmius Rotulae Fries, Epicrisis p 384. 1838. Collariati Bataille, Flore Monographique Marasmius d'Europe p 26. 1919. TYPE SPECIES. 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 specimens is 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 rather long erect or divergent setulae without long conspicuous hairs; spores small to large, ellipsoid to oblong, mostly (or always) acyanophilic; lamellae subcollariate to 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 rhizomorphs which are



92 Flora Neotropica

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

Marasmius sect Marasmius subsect Pararotulae (Singer) Singer, stat nov Marasmius sect Pararotulae Singer, 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 specimens essentially white or whitish (excepting the central knob or papilla which may be white or pigmented) on drying sometimes becoming very pale argillaceous or gilvous. 2. All or the great majority of the stipes attached not to the substratum but to conspicu-

ous black rhizomorphs. 3. Lamellae 9-16, mostly 12-14; pileus 6 mm broad; Gulf area. 32. M. multiceps. 3. Lamellae 7-9; pileus 0.5-2 mm broad: Amazonas region. 33. M. cupressiformis.

2. All or the great majority of the stipes attached to the substratum and accompanied or not by black rhizomorphs. 4. Central dot absent or light colored or brownish and soon disappearing; spores up

to 8.5 , long. 5. Spores small, 5-7 X 2.2-3.5 ; central knob or dot gilvous or cork-color or pale

flesh; lamellae 13-17; pileus 10 mm broad or broader; on leaves. 34. M. rotuloides. 5. Spores as small as above or larger; central dot, 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; central umto white. 35. Al. panamenisis. 6. On leaves.

7. Spores reaching more than 4.8p broad; pileus 1-1.8 mm broad; lamellae 5-8. 36. MI. pararotula.

7. Spores up to 4.8 M broad; lamellae 9-17. 8. Species of the Amazon region, Colombia or Venezuela; dark central

dot absent in fresh pileus. 9. Region of the central dot of the pileus not elevated but flat:

spores 5.5-8.5 X 2.5-4 p lamellae 11-17; tropical-montane species. 37. M. vergeliensis.

9. Region of the central dot somewhat to strongly elevated, showing at least in the dry stage a central knob or a small umbo or papilla in the umbilicus or in the smooth central apex of the pileus; spores as above or different; lamellae as above or more numerous or 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 (under a lens) surface when dried.

11. Lamellae 9-12. 38. M. apateliuls. 11. Lamellae 13-17. 39. M. rotalis.

10. Spores 5-6 X 2.5u; pileus 5 mm broad or rarely broader; umbo surrounded by a conspicuous white zone (dried).

40. M. castellanoi. 8. Species of the Southern Hemisphere. Yungas to Selva Boliviano-

Tucumana (see "12" below). 4. Central dot either fuliginous, deep ferruginous, deep brown or dark gray or, if

paler, spores larger than 10 long. 12. Spores 14.5-16.5 X 3.7-4.3p. 41. M. pandoanus 12. Spores smaller.

13. Spores 10-14 X 3-4 ; lamellae 10-16. 42. M. oaxacanus. 13. Spores smaller or broader or lamellae fewer.

14. Lamellae 6-9; spores 7.5-11 X 4.5 p. 43. M. manuripiensis. 14. Lamellae more numerous or spores either larger or smaller but

always narrower or broader than indicated above. 15. Pileus pale flesh pink; spores 8.5-9 X 3.5-4 p (see "24"

below).



Marasmius 93

15. Pileus white or whitish when fresh or mostly so, but never flesh pink ever; spores different.

16. Pileus with a rather pale or pallid umbo; spores 12.5-14.2 X 7.5-9,u; lamellae about seven. 44. M. psychotriophilus.

16. Pileus with a dark umbo; spores 5.5-7.8 X 2-3(-4)u; lamellae 14-20 (if fewer compare "6" above).

17. Pileus about 6 mm broad; lamellae about 20; on oak leaves. 45. M. cundinamarcae.

17. Pileus up to 5.5 mm broad; lamellae 11-17; on dead leaves other than oak. Southern South America (if in Northern North America. compare "11" above).

46. M. leucorotalis. I. Pileus in mature specimens not white in the larger marginal region (if only old, dried

pale avellaneous, pale ocher or pale argillaceous pilei seen, compare also "2" above). 18. Pileus sepia or mummy brown, 2-3 mm broad; lamellae on edges discolorous (con-

colorous with the pileus); spores 11-14 X 3.5-5 ; on bark. 47. M. arimanus. 18. Not combining these characters.

19. Pileus gray to beige (between blackish gray to pale cinereous when fresh). 20. Spores larger than 9,u long and 4.5g broad, (see also subsection Penicillati).

48. M. griseofuscescens. 20. Spores smaller.

21. Lamellae with cinereous to dark gray edge; spores 7-9 X 3.5-4.7 p; cheilocystidia with mostly sepia colored setulae and hyaline main body. Mexico.

49. M. scototephrodes. 21. Lamellae with white edge which may become brownish after drying;

spores 5.5-8 X 2.5-4ui; cheilocystidia either entirely hyaline or if setulae are brown, main body with hyaline wall but fuscous cell sap. Colombia.

50. M. leucozonitiformis. 19. Pileus not so colored.

22. Pileus when fresh purplish red or red or pale flesh pink with dark colored central dot on papilla; lamellae either with discolorous edge and collarium or evenly flesh-pink all over.

23. Spores 8-11 X 4.5-5u; edge of lamellae and collarium red; stipe 15-35 mm long. 54. M. carminis.

23. Spores narrower; edge of lamellae not discolorous; stipe sometimes longer. 24. Pileus pale flesh pink; lamellae concolorous with pileus. 53. M. carneotinctus. 24. Pileus differently colored; lamellae white 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 Flora Neotropica

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 Aextoxicum. 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 (compare 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

high. 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 collarium 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 different.

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. Marasmius multiceps Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 298. 1869. Pol)ymarasmlius muLlticeps (Berkeley & Curtis) Murrill, North American Flora 9: 286. 1915. Polymarasmius subnulticeps Murrill, North American Flora 9: 286. 1915. Marasmius subimulticeps (Murrill) Saccardo & Trotter, Sylloge Fungorum 23: 155. 1925. TYPE. Wright 160 (K), from Cuba. Pileus white but in age and on drying sometimes tending to become brownish ful-

vous, with chestnut-fuliginous to black umbo both fresh and dried, sulcate, but the umbo and papilla smooth, hemispheric or 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. Lamellae white, distant, with (9-)11-14(-16) through-lamellae, all entire or 1-3 lamellulae present, rather broad, shallowly but distinctly collariate. Stipe black (at first with pallid or brown apex), glabrous, insititious with a ring-joint and rising from black rhizomorphs, equal or tapering upwards, 5-25 X 0.2-0.5 mm . Black rhizomorphs glabrous, creeping or 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, inamyloid. Hymenium: basidia 4-spored, basidioles fusoid, some remaining sterile (cystidioles); cystidia none; cheilocystidia with main body hyaline, 12-13 X 6.8-7 p, beset with divergent setulae 4-5 X 0.8-1 ,u, also hyaline, attenuated to a subacute tip. Hyphae hyaline in the pileus-trama, chestnut-fuliginous in the stipe, there parallel and thin-to thick-walled, many distinctly pseudoamyloid but in the pileus-trama and the hymenophoral trama

(which is regular) mostly thin-walled and only slowly and vaguely pseudoamyloid, those of the rind of stipe and rhizomorphs strongly forking-interlaced, those of the core of the stipe and rhizomorphs often 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 numerous setulae which are either rod-shaped or 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. MATERIAL STUDIED. CUBA. Wright 160 (FH, K), type; Fungi Cubenses Wrightiani 132.

(FH). GUATEMALA. Plateau above Chiatla, 18 Nov 1959, Degener, comm. Petrak, II (BAFC). Dennis (1951c) data on the type of M. submulticeps show that it is conspecific

with A. multiceps. ILLUSTRATIONS. Dennis (1951 b) fig 11; (1951 c) fig 28.

33. Marasmius cupressiformis Berkeley, 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-chestnut colored 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 tapering upwards, insititious on rhizomorphs, 1.2-4 X 0.1-0.4 mm. Rhizomorphs black, long, often ramified and ascendant, 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 similar to the broom cells of the epicutis. Hyphae of the pileus trama hyaline, 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 epicuticular broom cells, (1) of the Rotalis-type, with the main body 11-14 X 7-11 U, clavate to vesiculose, sometimes 2-3-forked, setulae 1-2 X 0.5-0.7 p, divergent or obliquely erect in the subapicular region, hyaline, (2) of the Siccus-type, 20-34 X 6-13 p,, the latter type dense and numerous in the center, widely dispersed to 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-like or obtusely conical, 2-15 X 1.5-4 ,, some of the "broom cells" without appendages or setulae,

Over dead wood and leafmold in tropical rain forest. Pantropical (?), at least in the Amazonas and the Congo regions.

MATERIAL STUDIED. BRAZIL. Amazonas: "Panure" (=Sao Jer6nimo), Spruce 75 (K), type. ZAIRE. J. Ghesquiere 1355 (BR).

ILLUSTRATION. Singer (1964b), fig 15, D, E, F.; Berkeley (1856) pl 5. fig 3.

34. Marasmius rotuloides Dennis, Trans. Brit. Mycol. Soc. 34: 415. 1951. TYPE. Dennis 232, from Trinidad. Pileus white or whitish in tile marginal zone, 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 grayish beige, the central dot or umbo concolorous with the zone around the white zone, not chestnut either fresh or dried (the flesh-ocher zone 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. Lamellae white, distant (13-16 through-lamellae, no lamellulae), broad, distinctly collariate. Spore print white. Stipe dark red-brown to 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 hyaline thin to firm-walled, otherwise like the epicuticular broom-cells in shape and size. Hyphae with thin to often thick walls, filamentous, pseudoamyloid, with clamp connections, hyaline in pileus-trama and the regular hymenophoral trama, not gelatinized. Cortical layers: epicutis of pileus hymeniform, with broom-cells of 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 divergent setulae which are rod-shaped, hyaline to light stramineous, 1-2 X 0.8 8.

On dead, fallen, dicotyledonous leaves in tropical forest. MATERIAL STUDIED. TRINIDAD. Cerro de Aripo, 23 Oct 1949, Deinnis 232 (K), type.

BRAZIL. Para, Estancia Pirelli, 7 Jun 1966, Singer B 4231/I (BAFC), B 4224 (BAFC); Belem, 10 Jun 1966, Singer B 4252 (BAFC); B 4269 (BAFC).

ILLUSTRATION. Dennis (1951a) pl. 19, fig 14.

35. Marasmius panamensis Singer, sp nov TYPE. Martin & Welden 8733, from Panama. Pileo albo; lamellis albis, distantibus, latissimis, collariatis; stipite nigro apice albo,



Marasmius 97

rhizomorphis nigris praesentibus sed stipiti haud adhaesis. Sporis 4.5-7.2 X 2.5-3.6M; elementis epicuticularibus eis Marasmii rotalis analogis. Ad lignum in insula Barro Co- lorado. Typus in F conservatus est.

Pileus white, glabrous, sulcate, barrel-shaped to convex, umbonate in an umbilicus, about 3 mm broad. Lamellae white, 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, accompanied by (but not rising from) black rhizomorphs which 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 epicuticular broom cells. Hyphae of the regular gill 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-fulvous at times, setulae all over the cell, divergent.

On wood, Panama. MATERIAL STUDIED. PANAMA. Barro Colorado Island, 15/16 Aug 1952, Martin &

lelden 8733 (F), type.

36. Marasmius pararotula Singer, Sydowia 18: 340. 1965. TYPE. Singer B 1886, from Bolivia. Pileus white, occasionally with a central or eccentric gray spot, subglabrous or

glabrescent, under a lens aspersed with sugary particles which later disappear, campanulate, some convex, more or less distinctly umbilicate, especially when dried, sulcate when dried, 1-1.8 mm broad. Lamellae white, the edges with sugary fringe, under a lens appearing more or less distinctly collariate, rather broad, 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, glabrous and 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, rather broad, smooth, hyaline, inamyloid. Hymenium: basidia 2-4-spored, 16-17 X 7.3-7.7 u; cystidia none; cheilocystidia causing the sugary appearance of the edge of the lamellae, hyaline similar to the epicuticular elements. Hyphae: subhymenium of very short, small elements; hymenophoral trama regular, not bilateral, hyphae of pileus trama rather irregular in size and shape, subhyaline to hyaline in ammonia, not voluminous as in Mycena but some occasional cells rather broad and short although generally filamentous or subcylindric, interwoven, hyphae of stipe with slightly thickened walls and strictly parallel, hyaline only in upper portion of the stipe, filamentous; all hyphae pseudoamyloid and with clamp connections. 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 irregularly distributed over the upper portion of the surface of the cell, rather few at times, 0.7-5 X 1-2 A; where the trama or hypodermium is exposed, a Rameales-structure of nodose, irregular elements can be observed. No covering layer on the stipe.

On dead fallen leaves of dicotyledonous plants in tropical rain forest. MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, Guayaramerfn, 3 km below the

village in virgin forest, 12 Feb 1956, Singer B 1886 (LIL), type. ILLUSTRATION: Singer (1965) pl 7, fig 13.



98 Flora Neotropica

37. Marasmius vergeliensis Singer, sp nov Fig 44V. TYPE. Singer B 6426, from Colombia. Pileo albo, 3-4 mm lato; lamellis albis, subdistantibus vel distantibus, collariatis;

stipite nigo apice albo; rhizomorphis praesentibus sed stipiti haud adhaesis. Sporis 5.5 -8.5 X 2.5-4p; elementis epicuticularibus eis Marasmii rotalis analogis. Ad folia de-

lapsa dicotyledonea in Columbia. Typus in F conservatus est. 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 champagne color over the marginal zone, the latter strongly sulcate, convex, with a narrow, umbilicate depression in the center, the depression flat-bottomed and even-

tually often slightly tomentose under a lens, 3-4 mm broad and up to 3 mm high. Lamellae white, 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-bister or remaining black, 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 epicuticular broom cells. Hyphae of the hymenophoral trama regularly arranged, almost or quite inamyloid, those of the pileus-trama weakly, of the stipe-trama distinctly pseudoarnlyoid, thin-walled in the pileus, with clamp connections. 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-montane forest. Colombia. MATERIAL STUDIED. COLOMBIA. Valle, Mun. Dagua, Corr. Vergel, 27 Apr 1968, Singer

B 6426 (F), type; 14 Apr 1968, Singer B 6119 (F). This species is closely related to Marasmius capillaris Ellis from North America

which grows on oak and other leaves; common in the Midwest. The latter differs from Mlarasmius vergeliensis in the host, larger spores (7.5-10 X 3.8-4.3 p) and slightly larger carpophores.

38. Marasmius apatelius Singer, Bull. Jard. Bot. Bruxelles 24: 332. 1964. Fig 44A. TYPE. Vanderyst 695, from Zaire. Pileus at first white (?), dried beige to dull pale brownish or beige-ochraceous with

paler margin, long sulcate, glabrous, strongly convex, umbilicate, with a small umbo or

papilla in the umbilicus, 1-3 mm broad. Lamellae white, 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 developing pilei (telepodia), some associated with very thin, black rhizomorphs but 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 epicuticular broom cells. Hyphae of the pileus trama densely packed and inter-

woven, basically running radially, parallel in the stipe, pseudoamyloid, with clamp connections. 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.

MATERIAL STUDIED. VENEZUELA. Sucre: along Rio Media, 15 Jul 1972, Dumont et al VE 5145 (NY). ZAIRE. Bas-Congo, Kisantu, Vanderiyst 695 (BR), type.

ILLUSTRATION. Singer (1964b) fig 16, H- I.

39. Marasmius rotalis 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, grayish beige or concolorous with the margin, sulcate, glabrous or sometimes rivulose-subscabrous in a zone around the disc (and this zone consequently more pallid even in dried material but not

constantly so) the umbo itself not discolorous-dark but pallid to light avellaneous-argillaceous, convex, umbilicate, with a small, low umbo or an obtuse papilla in the umbilicus, 1.5-5 mm broad. Lamellae white to pale cream, with white to (dried) slightly discolorous edge (concolorous with the margin of the pileus), distant (13-17 through-lamellae, no lamellulae or more rarely 1-2 lamellulae present) broad, collariate, the collarium when dried also concolorous with the gill edges. Stipe black, blackish bister, or very dark chestnut, with white apex, glabrous, smooth, insititious, equal, usually accompanied by thin, dark rhizomorphs but 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 epicuticular broom cells. Hyphae of

pileus-trama inamyloid, 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-fuscous setulae 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: Quercus columbiana.

MATERIAL STUDIED. COLOMBIA. Boyaca.: Tunja, Arcabuco, 28 Jul 1960, Singer B 3560 (BAFC). CEYLON. Peradiniya, Thwaites 810 (K), type. ZAIRE. Yangambi, Reserve Isa- lowe, 470 m alt Louis 14934 (BR).

This species is close to Marasmius cundinamrarcae, M. apatelius, and M. leuco- rotalii In the Colombian oak woods it may most easily be confused with M. apatelius which has fewer lamellae and M. cundinanarcae with more numerous lamellae and smaller spores.

ILLUSTRATION. Singer (1964b), fig 16, E-G.

40. Marasmius castellanoi Singer, sp nov TYPE. Singer B 4199, from Brazil. Pileo albo vel albido-pallido, umbone exiguo subericolore in umbilico centrali in-

serto; lamellis albis, distantibus, collariatis; stipite primum nigricante apice albo. Sporis 5-6 X 2.5 :; hyphis pilei plerumque pseudoamyloideis, fibulatis; elementis epicuticularibus typi Marasmii rotalis praesentibus. Ad folia emortua dicotyledonea in Brasilia. Typus in BAFC conservatus est.

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. Lamellae white with white edges, medium 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 epicuticular broom cells. Hyphae of the hymenophoral trama and the pileus-trama hyaline, many of these distinctly pseudoamyloid, with clamp connections. Cortical layer: epicutis of the pileus hymeniform, consisting 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. MATERIAL STUDIED. BRAZIL: Para, Estancia Pirelli, 7 Jun 1966, Singer B 4199 (BAFC),

type.

41. Marasmius pandoanus Singer, 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. Lamellae white, edges concolorous with sides, broad, distant (11), collariate with a very distant collarium. Stipe black with white apex, glabrous and smooth, setose, insititious, 30 X 0.1 mm. Context very thin, inodorous.

Spores 14.5-16.5 X 3.7-4.3 p, ventricose-oblong with suprahilar depression and 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 diverging and short (1-2.8 p) and sometimes (those on summit of cells) branching-forked, entirely subhyaline, clavate or subventricose and tapering to 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 arranged but slightly disarranged in 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 appear almost short-hyphous (as in Marasmzius ntulticeps); surface of stipe with some minute outgrowths similar to 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, Santa Rosa on Rib Mladre de Dios, 22 Mar 1956, Singer B 2145 (LIL), type.

ILLUSTRATION: Singer (1965) pl 7, fig 15.

42. Marasmius oaxacanus Singer, Sydowia 12: 232. 1958. TYPE. Singer M 1519, from Mexico. Pileus white, becoming whitish-argillaceous or argillaceous-pallid when dried, with

a white zone around a pale fuscous to fuscous-subfuliginous dot which becomes light gray, pale fuscous or whitish when dried, smooth in the umbilicus, otherwise long sulcate, glabrous, convex, with deep umbilicus without an umbo or papilla inside but appearing tomentose when seen under a lens (dried), 3-4 mm broad. Lamellae white, subclose at first, subdistant to distant (10-16 through-lamellae no 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, inodorous.

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 epicuticular broom cells but always entirely hyaline. Hyphae of the pileus trama hyaline, thin-walled, with clamp connections, distinctly pseudoamyloid in pileus and stipe. Cortical layers: epicutis of the pileus hymeniform or subhymeniform, consisting of broom cells of the Rotalis-type, these entirely hyaline or some melleous-hyaline except those of the area of the dark, central dot of the pileus, either subhyaline or pale grayish fuscidulous, main body 15-24 X 15-22 m, vesiculose to vesiculose-subcylindric, oftenglobose, setulae thin-rod shaped, 1.3-1.5 M projecting.

On fallen leaves of Inga, Central Mexico. MATERIAL STUDIED. MEXICO: Oaxaca, Huautla de Jimenez, 1660 m alt, 10 Jul 1957,

Singer M 1519 (LIL) type; 8 Jul 1969, Singer M 8330 (F).

Young carpophores of Marasmius tanyspermus should not be mistaken for Maras- mius oaxacanus


43. Marasmius manuripiensis Singer, Sydowia 18: 338. 1965. TYPE. Singer B 2227, from Bolivia.

Pileus white to cream color with a minute ochraceous to pale fuscous central dot, dried pale ochraceous cinnamon or light caf6 au lait with concolorous or mostly cinnamon fuscous central dot, localized in a small umbo inside an umbilicus, sulcate, convex, 2-5 mm broad. Lamellae white, equal, 6-9, rather narrow, collariate to a narrow and moderately distinct collarium, distant. Stipe black with white apex, remaining so in dried condition, glabrous and smooth, setose, insititious, 4-10 X 0.2-0.5 mm.

Spores 7.5-11 X 4-5 A, ellipsoid, smooth, hyaline, with rather thin, inamyloid wall.

Hymenium: basidia 19 X 7.2 ,; basidioles fusoid, often with mucro; cystidia none; cheilo-

cystidia like the epicuticular elements. Hyphae of the pileus and hymenophoral trama partly inamyloid, partly weakly pseudoamlyoid, inamyloid in subhymenium; all hyphae with clamp connections. Cortical layers: epicutis of pileus consisting of elements of the

Rotalis-type, hymeniformly arranged, vesiculose, hyaline with hyaline to yellowish-subhyaline setulae, the main body 20-25 X 15-18 p, setulae divergent over the upper portion 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. MATERIAL STUDIED. BOLIVIA. Pando: Manuripi, Conquista, 25 Mar 1956, Singer B 2227

(LIL), type. ILLUSTRATION. Singer (1965), pl 7, fig 16.

44. Marasmius psychotriophilus Singer, 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 depressed around a central umbo, 1-1.5 mm broad. Lamellae white with white edge, distant (seven through-lamellae only), 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 trama thin-walled, filamentous, inamyloid. Cortical layers: epicutis of the pileus hymeniform, consisting of elements much like the cheilocystidia but setulae mostly pale avellaneous (from dried material in KOH).

On fallen leaves of Psychotriac MATERIAL STUDIED. BRAZIL: Rio de Janeiro, Serra dos Orgaos, 24 Oct 1961, Singer B

4029 (BAFC) type.

45. Marasmius cundinamarcae Singer, 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; rhizomorphis nullis. Sporis 5.5-7.2 X 2-3 A; elementis epicuticularibus typi Mara- smii rotalis. Ad folia delapsa quercina in Columbia. Typus in F conservatur est.

Pileus white to pale buff-white on dehydration and in the herbarium becoming pale beige around a white circular zone surrounding a 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. Lamellae cream white with white edge, moderately distant (+ 20 through-lamellae; no lamellulae) rather broad, collariate. Stipe fuliginous with white apex, glabrous, insititious, not accompanied by black rhizomorphs, equal or slightly tapering upwards, 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: basidia 4-spored; basidioles fusoid; cystidia none; cheilocystidia like the epicuticular broom cells, all hyaline. Hyphae in pileus and stipe pseudoamyloid with clamp connections. Cortical layer: 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 hyaline (fresh or recently dried in KOH).

On fallen oak leaves in oak woods. Colombia. MATERIAL STUDIED. COLOMBIA. Cundinamarca: Bosque de la Mercedes, Bojaca, 8

Apr 1968, Singer B 6009 (F), type.

46. Marasmius leucorotalis Singer, Sydowia 18: 337. 1965.

46a. Marasmius leucorotalis var 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 marginal zone on drying but leaving a 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. Lamellae white with white edge or cream white, subdistant or medium distant (14-15 through-lamellae, no lamellulae) rather broad, distinctly collariate. Stipe black or fuliginous with white apex, glabrous, smooth, equal or slightly tapering upward, setiform, insititious, 15-17 X 0.2-0.3 mm; no black rhizomorphs observed. 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 epicuticular broom cells, hyaline. Hyphae filamentous, hyaline, pseudoamyloid both in the pileus and stipe, with clamp connections. Cortical layer: 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 broader than long, sometimes longer than broad, sometimes pedicellate, setulae dow to the middle (or lower) of the cell, rod-shaped 1-2 X 0.3-1 p,

all hyaline. On fallen dicotyledonous leaves in tropical-montane forest. Bolivia. MATERIAL STUDIED. BOLIVIA. La Paz; Nor-Yungas, Rio Yariza, 23 Feb 1956, Singer

B 1414 (LIL) type. ILLUSTRATION. Singer (1965) pi 7, fig 17.

46b. Marasmius leucorotalis var discopallescens Singer, var nov TYPE. Singer T 3772, from Argentina. Maculo centrali pilei pallescente lamellisque magis distantibus differt. Pileus white, with fuscous-black to light fuscous central dot but this in dried

condition pallescent to whitish or gray and surrounded by a white ring even when the marginal zone has assumed a pale beige discoloration in the herbarium, more often without such a white ring-zone, glabrous, sulcate, hemispherical to convex, umbilicate, with or without a papilla in the depression, 1-3 mm broad. Lamellae white 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 accompanied by 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 epicuticular broom cells, all hyaline. Hyphae with clamp connections. Cortical layer: epicutis hymeniform, broom cells voluminous, with pale brownish to hyaline setulae 2 uhigh and rod-shaped, divergent.

On fallen leaves of Phoebe porphyria in subtropical forest. MATERIAL STUDIED. ARGENTINA: Tucumain, Rio Cochuna at 1350 m alt 27 Feb 1952,

Singer T 3772 (LIL), type.

46c. Marasmius leucorotalis var leucozonites (Singer) Singer, comb nov Marasmius leucozonites Singer, Sydowia 18: 337. 1965. Marasmius pluvialis Singer, Sydowia 18: 341. 1965. TYPE. Singer T 2018, from Argentina. Pileus white to dull ochraceous (fresh), elevations between sulci paler or white,

depression around dark central spot white and remaining so but marginal zone tending to become pale sordid ochraceous to pale argillaceous-beige in 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. Lamellae white, 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 associated 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 epicuticular 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. Cortical layer: epicutis hymeni-




form, broom cells of the Rotalis-type e g 28 X 16 p,, beset with hyaline to (in herbarium material mostly) pale melleous brownish setulae 202.7 p long, divergent.

On leaves of dicotyledonous trees, frequently (but not exclusively) on Phoebe porphyria.

MATERIAL STUDIED. ARGENTINA. Tucuman, Sierra de San Javier, Ciudad Universit- aria, 1000 m alt, 15 Jan 1955, Singer T 2018 (LIL), type.

The preceding variety differs from this in being less pigmented, smaller and with fewer lamellae and the central dark dot as well as the stipe becoming paler on drying, also in slightly smaller and relatively broader spores. Both differ from the type variety by a few rather secondary characters; the type variety appears to be intermediate between the other two varieties.

The binomial Marasmius pluvialis was intended for a species I have now redescribed as Marasmius hvlaeicola Singer (no. 63). Because of an unfortunate printer's error, the description of Marasmius leucozoniites with its type designation (T 2018) has been repeated under the heading Marasmius pluvialis Singer (1965, p 163) so that formally it becomes a synonym of Marasmius leucozonites.

Young specimens of Marasmius platvspermus might be looked for here or under M cundinamarcae. It differs however in habitat and broader spores.

47. Marasmius arimanus Dennis, Trans. Brit. Mycol. Soc. 34: 418. 1951 (as Xi. ari- mana).

TYPE. Dennis 391, from Trinidad. Pileus sepia or mummy brown, striate-sulcate to the center, convex with a

shallow umbilicus, 2-3 mm broad. Lamellae pale buff, distant (7), moderately broad, attenuated to each end, attached to a shallow collarium forming little more than a ridge round the apex of the stipe, with sepia colored edges. Stipe almost black, slender, wiry, smooth and polished throughout. Context thin, white, membranous.

Spores 11-14 X 3.5-4 ,, tear-shaped, inamyloid. Hymenium: cheilocystidia like the epicuticular broom cells, 8 lpbroad. Covering layers: epicutis of pileus formed by pear-shaped elements with reddish brown processes.

On bark of dead logs, Trinidad: Arima, Miss Dora Kong, 28 Nov 1949, comm. Dennis 391 (K), type.

I have studied the material superficially; it consists of a fragment which I did not dare section. It is not fully clear whether the epicuticular broom cells and cheilocystidia are of the Rotalis-type or the Siccus-type. The species seems to be close to Marasmiusfuligineorotula Singer. 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 carpophores much smaller in Dennis' species.

ILLUSTRATION. Dennis (1951a) pl 23, fig 5.

48. Marasmius griseofuscescens Singer, 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 atrocastaneo apice albopallido. Sporis 9-11 X 4-5.5 u; hyphis fibulatis, in tramate pilei inamyloideis; elementis epicuticularibus typi Marasmii rotalis. 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-pallid to cinereous ground of the



Marasmius 105

non-papillate umbilicatus, 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-pallid apex, 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: basidia 21-26.5 X 7-9 ,, 4-spored; basidioles fusoid; cystidia none; cheilocystidia like the

epicuticular broom cells but all hyaline. Hyphae filamentous, some swollen to 15 p,

hyaline in NH40H, with clamp connections, in the trama of the pileus and hymenophore 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 subtropical forest.

MATERIAL STUDIED. ECUADOR. Napo, Lago Agrio, 11 May 1973, Singer B 7361 (F) type.

This species is characteristic because 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 measurements may not cover the entire breadth of variation.

It is probable that the specimens registered by 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, Cataratas del lguazfi, 22/24 Nov 1949 (LIL) and are perhaps specimens which have passed the original gray state.


49. Marasmius scototephrodes Singer, sp nov Fig 46. TYPE. Singer M 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; rhizomorphis tenuissimis nigris praesentibus sed stipiti haud adhaesis. Sporis 7-9 X 3.5- 4.7 u; cheilocystidiis sepiaceo-setulosis; hyphis fibulatis, pseudoamyloideis; elementis epicuticularibus typi Marasmii rotalis. Ad folia, petiolos, frustula Dicotyledonum in Mexico. Typus in F conservatus est.

Pileus ash gray with a white or concolorous zone around a central light gray to dark gray dot, in the intermediate zone and the center smooth but surface tending to become cracked, marginal zone long-sulcate, glabrous, convex, umbilicate, not papillate, 4-6 mm broad. Lamellae white with cinereous to dark gray edge, distant (10-12 through-lamellae, no lamellulae), broad, distinctly collariate, collarium gray. Stipe black, shining, equal, setiform insititious, 20-25 X 0.1-0.2 mm, accompanied by very thin black rhizomorphs but 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 suprahilar depression. Hymenium: basidia 19-21.5 X 5.5-6.8 ,; basidioles fusoid; cystidia none; cheilocystidia like the epicuticular broom 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 numerous connections. Cortical layer: epicutis of the pileus hymeniform, consisting of broom cells of the Rotalis-type, main body 10-20 X




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-shaped and 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. Marasmius leucozonitiformis Singer sp nov TYPE. Singer B 6105 A, from Colombia. Pileo griseo vel pallide griseo, zona albida circum maculam centralem atrum prae-

dito, 1.5-5.5 mm lato; lamellis albidis acie frequenter discoloribus, distantibus, collariatis; stipite nigro acie albo. Sporis 5.5-8 X 2.5-4 /; elementis epicuticularibus typi Marasmii rotalis. Ad folia dicotyledonea in Columbia. Tupus in F conservatur.

Pileus gray, pale gray, often turning beige on drying, retaining however a white or whitish zone around the dark dot in the center of the umbilicus, this zone whitish because minutely rivulose cracked (visible under a lens), showing the trama, the dark spot subfuliginous but 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. Lamellae white 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 numerous black rhizomorphs accom-

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 epicuticular 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, Cordillera Occidental.

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, Singer B 6316 (F).

Among the species with originally white pileus, this pale gray or gray species is closest to Marasmius leucorotalis 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. Marasmius tanyspermus Singer, 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 marginal one surrounding the central dark dot which becomes

pale sordid gray in dried material, sulcate except in the umbilicus and with a small



Marasmius 107

umbo corresponding in extension to the blackish dot inside the umbilicus, convex otherwise, 1.5-3.2 mm broad. Lamellae pale cream, 10-11, rather broad, with white edge, collariate. Stipe umber with white apex, glabrous, insititious, 20 X 0.1 mm. No rhizomorphs.

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 epicuticular elements. Hyphae of pileus, hymenophoral trama 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. Covering layers: 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), divergent.

On dead dicotyledonous leaves in tropical rain forest. MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, Guayaramerin, 16 Mar 1956, Singer

B 2052 (LIL), type. When seen in the white stage, the carpophores may be mistaken for those of

Marasmius panldoalus or Marasmius oaxacanus which do not become brown when mature and have broader spores.

52. Marasmius carneotinctus 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 atrobrunneo apice albido. Sporis 8.5-9 X 3.5-4 A; elementis epicuticularibus typi Marasmii rotalis. In silva tropicali-montana Columbiae. Typus in ENCB conservatur.

Pileus flesh-pink with deep brown papilla, glabrous, sulcate, towards margin finely venose under a lens, convex, deeply umbilicate with a low papilla in the umbilicus, 34 mm broad. Lamellae flesh-pink, subclose (17 through-lamellae, no lamellulae) with an often poorly developed collarium. Stipe blackish-brown with 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: basidia 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 remarkably broad, 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-hyaline to pale melleous, divergent.

In tropical-montane forest, Colombia, west slope of the Cordillera Occidental. MATERIAL STUDIED. COLOMBIA. Valle, along road to Buenaventura. Pacific slope,

1950 m alt, 26 Jul 1964. Guzman 4580 (ENCB) type.

53. Marasmius violeorotalis 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 epicuticularibus typi Marasmii rotalis. Ad folia dicotyledonea emortua in Columbia. Typus in F conservatus est.

Pileus violet-lilac with an almost pallid zone around the fulvous-brown papilla, this zone not distinctly white but appearing pallid because of the rivulose-cracked




visible (under a lens) surface, showing the trama, glabrous, sulcate, excepting the center, strongly convex, umbilicate, with a low papilla in the umbilicus, about 5 mm broad. Lamellae lilac-pallid with 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 epicuticular broom 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-montane forest, Cordillera Occidental, Colombia.

MATERIAL STUDIED. COLOMBIA. Valle, near Dagua road above km 25, at 1900-2000 n alt, 28 Apr 1968, Singer B 6450 (F) type.

54. Marasmius carminis Singer, sp nov Marasmius carminis Singer, Sydowia 18: 156. 1965 (without Latin). TYPE. Singer B 1484, from Bolivia. Pileo purpureo-rubro, circum maculam nigram centralem albo vel rubro, 0.5-2 mm

lato et alto; lamellis albis, rubromarginatis, 12-13 integris tantum; stipite 15-35 mm longo, rhizomorphis nullis. Sporis 7-11 X 4-5.5; ;cheilocystidiis typi Marasmii rotalis. Ad folia emortua dicotyledonea in silva tropicali-montana. Typus in LIL conservatus est.

Pileus deep purple red, in age and particularly on drying fading to red-brown ("chutney"), sulcate, white around a central black dot or concolorous there, umbilicate, in dried material gray to gray-black in umbilicus, bottom of umbilicus somewhat convex very young caps convex and umbonate, later convex-umlbilicate but 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 margin of the pileus and likewise colored on the

edge of the collarium, about 12-13, broad. Stipe buff brown to blackish brown with white apex, glabrous and smooth, shining, setose, insititious, 15-35 X 0.1-0.2 mm. No

rhizomorphs. 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 larger elements 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 epicuticular broom cells but hyaline where edge is concolorous with sides of lamellae. Hyphae in pileus trama moderately strongly but rapidly reacting: pseudoamyloid, with clamp connections. Covering layer: 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-pedicellate or vesiculose and sessile, rarely forked or laterally elongated, hyaline to pale brownish and beset with numerous rod-shaped to conic-spinulose-obtuse setulae 2-3 X 1.5-2 p.

On dead dicotyledonous leaves in tropical montane forest. MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Carmen Pampa, 26 Feb 1956, at

2000 m alt, Singer B 1484 (LIL), type; B 1477 (LIL), paratype. ILLUSTRATION. Singer (1965), p1. 7, fig. 18.



Marasmius 109

55. Marasmius buzae Dennis, Kew Bull. 15: 98. 1961. TYPE. Dennis 1775, from Venezuela. Pileus a rather saturated orange ochraceous to deep orange ferruginous but be-

tween striae and around a central black dot paler to pallid, otherwise "tarragona" or 5-B12 M&P paler regions "sundown" or paler, little changing when dry (between "rust, sorolla" and "Arab" M&P), sulcate over most of the declivous zone, with a flat-bottomed umbilicus, truncate-conic to terete-cylindric or barrel-shaped, about 1.5 mm broad and high. Lamellae whitish with an ochraceous to ocher brown border near the margin of the pileus and on the collarium, subdistant to distant, 7-11, very broad, collariate. Stipe umber to umber-black or black, glabrous and smooth, shining, insititious, 17-24 X 0.1 mm, rising from the substratum but occasionally accompanied by scanty, thick, black rhizomorphs.

Spores voluminous, 8.5-13 X (5-)5.7-9(-10.3) , hyaline, smooth. Hymenium: basidia voluminous, 29-40 X 9.5-11.7 ,, 4-spored, sometimes a few 2-spored ones present; cystidia none observed; cheilocystidia like epicuticular elements but with the hyaline ones predominating at 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. Covering layers: epicutis of pileus hymeniform, consisting of elements of the Rotalis-type, 12-35 X 11-29 u, vesiculose to clavate or cylindrical, setulae diverging, short and rather coarse, 1.3-2.3 X 1.3-1.6 p, often covering the whole upper half of the cell, in some cells hyaline but mostly brown to brownish ochraceous.

On dead fallen dicotyledonous leaves in lower marginal zone of cloud forest. MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Cerro Uchumachi at 2200 m alt

25 Feb 1956, Singer B 1433 (LIL); Cataratas "San Juan" 2400 m alt, 28 Jan 1956, Singer B 656 (LIL). VENEZUELA. Laguna Negra, Sierra de Santo Domingo, Estado Merida, 1 Aug 1958, Dennis 1775 (K), type.

ILLUSTRATION Dennis (1961), fig 51.

56. Marasmius populiformis Berkeley, 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 concolorous 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: epicuticular elements about 8 u in diameter, with short, reddish-brown processes, apparently of the Rotalis-type,

On dead branches. MATERIAL STUDIED. 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.




Pileus Brussels brown (Ridgway), radially sulcate from all inconspicuous central, darker area, convex, 1.5-2 mm broad. Lamellae cream, edge concolorous, equal, 8-10, broad, collariate. Stipe dark brown with paler apex, hair-like, smooth, 4-6 mm long, insititious.

Spores 6-7 X 2-2.5 p, elliptical. Hymenium: cystidia none; cheilocystidia similar to epicuticular elements. Hyphae of trama inamyloid. Covering layer of pileus: epicutis formed by subglobose cells 10-17 y, in diameter, with thin brown walls bearing on their outer surface numerous minute hemispherical processes up to 1 ,projecting.

On fallen, slender twigs. MATERIAL STUDIED. VENEZUELA. Dto. Federal, Caracas Botanical Garden, 950 m

alt, 19 Jun 1958, Dennis 1068 (K), type. ILLUSTRATION. Singer (1965) pl 7, fig 20, Dennis (1961), fig 52.

58. Marasmius dodecaphyllus Singer, Sydowia 18: 335. 1965. TYPE. Singer B 2151, from Bolivia. Pileus ochraceous-avellaneous when fresh, helvolous (much like dried pilei of

M. helvolus) when dried, with a central, deep gray flat dot, the dot remaining deep gray and flat in dried material, around it somewhat rivulose under a good lens, otherwise quite glabrous, sulcate or slightly sulcate in marginal half, not pallid or white around central dot, convex with depressed center, about 9 mm broad. Lamellae white with white edges, buff when dried, moderately broad, 12 through-lamellae and 2 lamellulae, 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 perhaps more variable (few seen in type). smooth, hyaline, ellipsoid, inamyloid. Hymenium: basidia 23-24 X 7.5 ,: cystidia none: chei-

locystidialike epicuticular elements but on an average smaller and hyaline to subhyaline. Hyphae of pileus and hymenophoral trama 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 divergent setulae, main body 19-27 X 22 ;A, setulae 1-1.3 , long.

On a fallen piece of rotting dicotyledonous wood in tropical rain forest. MATERIAL STUDIED. BOLIVIA. Pando: Manuripi, Santa Rosa on Rio Madre de Dios, 22

Mar 1956, Singer B 2151 (LIL), type. This species differs from M. nanuiripiensis in a different (deeper) color, also being

differently colored in the herbarium, furthermore in a larger number of lamellae, flat central dark spot, and large diameter of the pileus.

Among the European species, this comes close to M. wettsteinii (a species often considered conspecific with Ml. bulliardii) on coniferous needles which has 10-16 lamellae but smaller pileus and longer stipe M wettsteinzii is constantly accompanied by blackish, thin rhizomorphs.

ILLUSTRATION. Singer (1965) pl 7, fig 21.

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

59a. Marasmius platyspermus forma platyspermus TYPE. Singer T 203, from Argentina. Pileus at first whitish with gray-brown to dark gray central dot, soon becoming

"Yellow beige" (13-H-7) to "wigwam" (14-G-10) or "bure" (13-H-8), dried about "butterscotch" (14-F-9 M&P) generally remaining whitish 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. Lamellae white, eventually avellaneous pallid, distant or subdistant, 13-20, all equal, broad, sometimes distinctly anastomosing in 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 substratum but sometimes observed to arise from a rhizomorph, 12-75 X 0.2-0.5 mm, accompanied by 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 epicuticular elements. Hyphae of pileus occasionally weakly and slowly pseudoamyloid or inamyloid, those of hymenophoral trama 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 ascending and 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

longer. On trunks, logs, fallen branches, roots, rotting in subtropical forest and less humid

montane woods, also in the montane Alnetum, lower zone, gregarious. Known hosts: Boelznleria, Schilus, Allophylus. Altitudinal range: 800-1600 m.

MATERIAL STUDIED. ARGENTINA. Tucuman: Sierra de San Javier, Parque Aconquija, 12 Mar 1949, Singer T 203 (LIL) type; 5 Mar 1949, Sinzger & Digilio T 195 (LIL) paratype; 24 Mar 1949, Singer 259 (LIL); Anta Muerta, 24 Apr 1949, Singer & Digilio T 501 (LIL); 26 Dec 1951, Singer T 1669 (LIL); paratype; Ciudad Universitaria, 20 Mar 1951, Singer T 1408 (LIL); Mar 1957; Singer T 3049 (LIL); San Agustfn, 1380 m alt, 15 Jan 1957, Singer T 2902 (LIL); Cat- amarca: Andalgala, above Estancia Santa Rosa (Suncho), 18 Jan 1952, Singer T 1 755 (LIL).


59b. Marasmius platyspermus forma scandens Singer, forma nov TYPE. Singer T 3591, from Argentina. Pileo pallidiore et habitatione in arbore viva differt. Typus in LIL conservatur. Pileus white, then pale avellaneous tan in age and when dried, with a flat central

black dot in a depression, otherwise convex, grooved-sulcate, glabrous, 5-9 mm broad. Lamellae white, broad, with the edge concolorous with the sides, without lamellulae, 19-21, widely collariate. Stipe at first white, later becoming black from the base upwards, glabrous, shining, smooth, equal or slightly tapering upwards, 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 suprahilar depression or applanation, thin-walled. Hymenium: cheilocystidia like epicuticular broom 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. Covering layers: epicutis of pileus hymeniform 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 rather thin




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, gregarious but not cespitose. MATERIAL STUDIED. ARGENTINA. Tucuman: Departanento Taff, Ciudad Universit-

aria, Sierra de San Javier at 1200 malt. 24 Mar 1960, Singer T 3591 (LIL), type. This seems to be a habitat form with somewhat paler pileus and very slightly

smaller spores, rare. It should not be confused with AL. leucorotalis which differs in having fewer lamellae and narrower spores.

This common species of the Argentine Northwest is confined to the lower range of the mountains. The preceding species (no. 58) differs in less numerous and therefore more distant lamellae, slightly different colors, generally shorter and thinner stipe and more consistently acanthophysoid (equally and always divergently short setuloid) epicuticular elements. The rhizomorphs are either less or not developed in M. dodecaphyllus.

Among the holarctic species, MI. wettsteinii differs in growing on conifer needles, slightly smaller, especially narrower spores and a slight difference in colors. M bulliardii differs in growing on frondose leaves, fewer lamellae and even narrower spores.

60. Marasmius nebularum Singer, 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. Lamellae white 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, inamyloid. Hymenium: basidia clavate, basidioles fusoid; cystidia none; cheilocystidia hyaline, otherwise like the epicuticular broom cells. Hyphae of the pileus-trama hyaline, with clamp connections, inamyloid. Cortical layer: epicutis of the pileus hymeniform, 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. MaTERIAL STUDIED. COLOMBIA. Boyaca, between Ramiriquiand Rond6n, 30 Jul 1960,

Singer B 3637 (BAFC), type.

61. Marasmius aequatorialis Singer, 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 tramatis pilei inamyloideis, frequenter latissimis, fibulatis; elementis epicuticularibus typi Marasmii rotalis, hyalinis, succineo-setulosis. Ad ramos radicesque Leguminosarum in regione andina Aequatoriae. Typus in F conservatus est.

Pileus bright rusty orange or orange when fresh, sometimes fading to whitish, the low and flat-bottomed umbilicus white with a central darker dot, this however rarely in form of an extremely minute papilla and the umbilicus area brown-scurfy-cracked on white context ground, the orange portion becoming brown and paler striped on drying, 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 diverging hyaline setulae, 1.5-2.5 s long. Hyphae of the pileus trama hyaline, with clamp connections, inamyloid, often very broad (3-19 p diameter in mature specimens); thin- to firm-walled; hy- phac of the stipe dark pigmented, parallel with 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, diverging and

covering a wide area of the cell, smooth cells very scattered or absent.

On dead standing branches or roots of leguminous shrubs and trees in the Andean re,ionl, gregarious.

MATERIAL STUDIED. ECUADOR. Pichincha, Rio Machaingaro, 2600 m alt, 20 May 1973, Singer B 7538 (1), type.

This differs fronm M. icnbularumt in the not uniformly colored pileus, broader yet shorter spores and, as from mlost species in this group, in the fact that the fully mature specimenll shows a large majority of smooth cheilocystidia.

62. Marasmius nothomyrciae Singer, Arkiv Botanik 4: 384. 1959. TYPI:. Sparre F 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 showing the white pileus-trama), dried here deep-gray to buff-gray or whitish to light cin- inamon to 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 grayissh black to black, glabrous, excepting the scurfy zone, where present, sulcate except in the umnbilicus, half-barrel-shaped, then strongly convex to convex, always abruptly umbilicate, 2-7 mm broad. Lamellae white with weakly to decididly discolored ("paloma" M&P) edge and collarium, the discoloration not always complete and ofteii visible only with a lens in dried condition, rather distant (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 accompanied by (but not attached to) numerour black rhizomorphs, insititious, 15-50 X 0.1-0.3 mm, often remiaining sterile ("telepodia"). Context very thin in the pileus, inodorous.

Spores (6-)6.5-10 X 3.5-4.5(-5) p, fusoid, oblong-fusoid or ellipsoid, smooth, hyaline, inamyloid. Hymenium: basidia about 23 X 6.5 p, 4-spored; basidioles fusoid; cystidia lnone: cheilocystidia like epicuticular broom cells but setulae either golden to golden hyaline or hyaline. Hyphae of the pileus-trama hyaline, 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-ferruginous setulae covering the apex or the upper




two thirds of the cell, setulae 1.2-2.7 X 0.9-1.3 ji, cylindric or verruculose, obtuse or subobtuse.

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. Juan Fernandez: Masatierra, path to El Camote, about 400 m alt, 18 Mar 1955, Sparre F 56 (S) type; Coquimbo: Fray Jorge, 16 Apr 1967, Singer M 6515 (SGO). As for Central and South Chilean collections studied see Singer (1969, p. 103).

This species is somewhat marginal as far as the neotropics are concerned since the only locality known in the subtropical belt is a fog forest in North Chile.

63. Marasmius hylaeicola Singer, sp nov Fig 50. TYPE. Singer B 7298, from Ecuador. Pileo brunneo, zona alba vel pallidiore circum maculum centralem nigrum ornanto,

3-5 mm lato; lamellis albis, collariatis; stipite brunneolo apice albo vel pallido; rhizomorphis nullis visis. Sporis 5.5-7.5 X 2-3.5 ,; elementis epicuticularibus typi Marasmii rotalis. Ad folia delapsa arborum dicotyledonearum in hylaea. Typus in F conservatus est.

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 rather wide zone around a black or blackish dot, dried becoming "kis kilim" to "cattail" (M&P) in marginal zone and black or gray in center with the pallid zone remaining + distinct, sulcate, glabrous, convex, umbilicate, with a flat bottom or dark dot slightly elevated, 3-5 mm broad. Lamellae white, with white edge or with an inconsistent and very narrow discoloration on the edge (lens!) which is concolorous with the margin of 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 rhizomorphs seen. 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 epicuticular broom cells but many or all entirely hyaline, often scarce. Hyphae of the pileus-trama hyaline, weakly to distinctly pseudoamyloid, with clamp connections, strongly pseudoamyloid and parallel in the apex of the stipe. Cortical layer: 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. MATERIAL STUDIED. ECUADOR. Napo. Lago Agrio, 8 May 1973, Singer B 7298 (F),

type. BOLIVIA. Beni, Vaca Diez, 3 km below Guayaramerin 12 Mar 1956, Singer B 1873 (BAFC); B 1874 (BAFC).

Marasmius hylaeicola is characterized by 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 Maramius letcorotalis in its color and distribution and from Marasmius minimus Dennis in the presence of a pallid zone and its habitat on dead leaves rather than twigs, more numerous lamellae and larger size. It also differs from all forms of the Marasmius variabiliceps complex in having smaller spores.

ILLUSTRATION. Singer (1965) pl. 7, fig 22.



Marasmius 115

64. Marasmius vigintifolius 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 margin when 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 narrow to broad and conspicuous white zone around the central dark dot which is black to blackish but may become concolorous with margin in dried material, glabrous, sulcate over one half to two thirds of the radius, campanulate to truncate-conical when young, mostly becoming convex when mature, with a rather broad and deep umbilicus, with or without a minute umbo in the umbilicus, 2-10 mm broad. Lamellae white, edges and collarium usually distinctly brown or at least partially so, 17-22, mostly 20, subclose to subdistant, rather broad to broad (1.5 mm), collarium narrow or wide. Stipe umber black to black with initially white apex, dried fuliginous or deep chestnut to nearly black, shining, glabrous and smooth, insititious, setose, or very slightly tapering towards apex, 14-56 X 0.1-0.3 mm; no rhizomorphs or with scanty dark rhizomorphs unconnected 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 diameter which are a special type of basidiole; true pleurocystidia none; cheilocystidia like epicuticular broom cells. Hyphae hyaline, filamentous, slowly and weakly pseudoamyloid in trama of pileus. Cortical layers: epicutis of pileus hymeniform or subhymeniform, its elements vesiculose or vesiculose-pedicellate, sometimes in adult specimens horizontally extended, main body 13.5-20 X 19-23.5 A, very few almost hyphous, of the Rotalis-type, hyaline, beset with diverging short, rod-like setulae which are melleous brownish, deep ochraceous brown or deep sepia, 2-27 X 1-1.3 ,.

On blade and petioles of fallen dicotyledonous leaves in montane tropical forest. MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Coroico, 4 Feb 1956, Singer B 871

(LIL), type; B 873 (LIL), paratype; Carmen Pampa 1 Feb 1956, B 800 (LIL), paratype; 17 Feb 1956, B 1245 (LIL), paratype; 19 Feb 1956, B 1956 (LIL), paratype. BRAZIL. Guanabara, Floresta de Tijuca 12 Oct 1961, Singer B 4006 (BAFC).


65. Marasmius idroboi Singer, sp nov TYPE. Singer B 6925, from Colombia. Pileo brunneo, zona alba vel alutaceo-pallida circum maculam brunneam vel fus-

cum centralem praedito, 4-8 mm lato; lamellis albis, collario pallide coriicolori praeditis, subconfertis; stipite nigro; rhizomorphis nullis. Sporis 6.5-8 X 2.8-3.8 p; hyphis nonnullis tramatis pilei pseudoamyloideis, fibulatis; elementis epicuticularibus typi Marasmii rotalis. Ad folia dicotyledonea delapsa in Columbia. Typus in F conservatus est.

Pileus brown, fresh ("oak M&P) and dried ("Cochin" M&P) with paler radial stripes at least when fresh, with a white to buff-pallid zone around a central brown to fuscous dot which bleaches to buff-pallid or gray when dried, sulcate, convex with steeply declivous margin and abrupt umbilicus which is flattened on the ground, 4-8 mm broad and up to 6 mm high. Lamellae white with pale tan collarium and sometimes also the edges finely and weakly discolored brown to pale tan under a lens, sub-




close (20-23 through-lamellae, no lamellulae), broad, distinctly, collariate. Stipe black (perhaps initially with white apex), glabrous, insititious, not accompanied by rhizomorphs, 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; basidioles fusoid; cystidia none; cheilocystidia like the epicuticular broom cells, those on the very edge more or less pigmented with some hyaline one intermixed, those accompanying them further towards the side of the lamellae, all hyaline. Hyphae of the pileus-trama hyaline, thin-walled, some thick-walled, some distinctly pseudoamyloid, some weakly pseudoamyloid, all with clamp connections; some cleiferous hyphae present. Cortical layers: epicutis of the pileus hymeniform, broom cells of the Rotalis type, main body either entirely hyaline or hyaline 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 ,, verruculose or rod-shaped, mostly brown, fewer subhyaline.

On dead fallen dicotyledonous (ericaceous and rubiaceous) leaves. Colombia. MATERIAL STUDIED. COLOMBIA. Boyaca: Tunja, Arcubuco, Finca de Luis Antonio

Porras, 29 Jul 1960, Singer B 3599 (BAFC); Valle, Cali, Cerro Horqueta, 2000 m alt, 2 May 1968, Singer B 6925 (F), type.

This species is closely related to Marasmius vigintifolius but differs in the narrower spores and the radiately striped pileus.

66. Marasmius eucladopus Singer, Sydowia 18: 336. 1965. TYPE. Singer,B 1478, from Bolivia. Pileus somewhat more ochraceous than "copper br." M&P (rusty), dried turning

deep red-brown ("Mandalay" M&P), dried with a pallid center, margined by a wall, with an insignificant umbo in the pallid zone but umbo not black at least in dried material, sulcate on the large, colored, declivous portion, convex to truncate-conical with a central depression in 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 material collarium wide and distinct, distant (9-10), very broad. Stipe light dirty umber, smooth and glabrous with white, thinner apex, inserted without any ring or rings into a central common rhizomorphic stipe-like strand, individual stipe 5-11 X 0.1 mm; stipe-bearing rhizomorphs black, above more dirty umber, smooth and glabrous, producing several carpophores, insititious.

Spores 8.8-9 X 5.8 p, ellipsoid, smooth, hyaline (few seen). Hymenium: basidia 16-17 X 8 ,; cystidia none; cheilocystidia like epicuticular elements but mostly hyaline. Hyphae of pileus-trama hyaline, practically inamyloid, yellowish even after 30 minutes exposure to Melzer's, reagent, with clamp connections. Cortical layers: epicutis of pileus hymeniform, its elements of the Rotalis-type, the majority covered down to the middle by diverging short setulae, or reminiscent of the Siccus-type (with the setulae although short and diverging all 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. MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Carmen Pampa, 26 Feb 1956, Singer,

B 1478 (LIL), type. The idea that this might be M. populiformis, not rediscovered since 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 different.




Marasmius 117

67. Marasmius tetrachrous Singer 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 ferruginous brown to ferruginous chestnut e g "chutney," fresh showing a rather narrow white ring around a central black dot but this latter usually disappearing on 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, campanulate or conical, then almost cylindrical-drumshaped but with narrower disc 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 rhizmorphs observed.

Spores 7-10.5 X 3.5-5(-6.5), mostly 9.5-10.5 X ? 5p, ellipsoid, smooth, hyaline, inamyloid. Hymenium: basidia broad and short, 16.5-19.5 X 10-1 1u cystidia none; cheilocystidia of the Rotalis-type, vesiculose, 16-17p broad, hyaline. Hyphae of the pileus trama weakly pseudoamyloid. Covering layers: 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.

MATERIAL STUDIED. BOLIVIA. La Paz: Nor Yungas, Carmen Pampa, 26 Feb 1956, Singer B 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 collarium and the medium-sized ellipsoid spores are characteristic.


68. Marasmius tereticeps Singer, Sydowia 18: 343.1965. TYPE. Singer B 2299, from Bolivia. Pileus cinnamon ochraceous, not becoming deep ferruginous chestnut when dried,

the upper flat portion white with a darker dot in the middle of the white field which is surrounded by a wall-like convex zone concolorous with the vertical marginal zone, central 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, extraordinarily broad (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, glabrous shining, 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 epicuticular elements but with hyaline or subhyaline setulae. Hyphae: in pileus and hymenophoral trama regular but very thin, hyphae hyaiine, inamyloid, with clamp connections. Cortical layer: epicutis of pileus hymeniform, of Rotalis-type elements with the main body 18-30 X 12-21 u, 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).




On dicotyledonous leaves fallen to the ground in tropical rain forest. MATERIAL STUDIED. BOLIVIA. Pando: Madre de Dios, Santa Teresa, 28 Mar 1956,

Singer B 2299 (LIL) type. ILLUSTRATION. Singer (1965) pl 8, fig 28.

69. Marasmius variabiliceps Singer, Sydowia 18: 344. 1965.

69a. Marasmius variabiliceps var variabiliceps TYPE. Singer B 1531, from Bolivia. Pileus varying between the color of M. platyspermus and Al. vigintifblius when fresh

but on partial dehydration becoming deep rust brown, fresh e g "russet br," partially dried between "Rust sorolla" and "Arab" and in the herbarium between "chutney" and "Java" M&P (red-brown to brown-red, a deep shade), with a rather broad white zone around a central dark dot which is black or blackish when fresh and gray to palest sordid brown when dried, glabrous, sulcate, convex, with a central depression and not or scarcely umbonate in the depression 5-9 mm broad. Lamellae pallid to white, with brown collarium with 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 tapering upwards or equal, setose, insititious, 58-100 X 0.2-0.4 mm; rhizomorphs scanty, inconspicuous, independent of stipe.

Spores rather variable in shape and size, 8.3-11.5 X 3.5-5.5,, often relatively broad ellipsoid, hyaline, smooth. Hymenium: basidia broad as in i1. vigintijolius; cystidia none; cheilocystidia like epicuticular elements but all hyaline. Cortical layer: epicutis of pileus hymeniform or subhymeniform, of the Rotalis-type, very few epicuticular elements approaching somewhat 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. La Paz: Nor-Yungas, Carmen Pampa, 26 Feb 1956,

Singer B 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 epicuticular elements, also the longer stipe and more variable spores.

ILLUSTRATION. Singer (1965) pl 8, fig 24. There is a number of forms, all characteristic of the Yungas region, which are dif-

ficult to separate from the type form as independent species and which appear to be de- pauperate forms, yet by their spore and lamellae characters varying within certain definite limits they are apparently hereditary races. They are neither geographical races nor mere forms. However, I have listed them as "varieties" (Singer 1941). It will not be possible to determine the correct taxon applicable as long as the infertility between these forms has not been confirmed experimentally. It may be that we deal here with substratum adaptations (mycoecotypes), but it is premature to guess because it was impossible to determine the leaves on which every form occurred inasmuch as 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 varieties here included was found in the continuation of the Yungas towards the south in Argentine territory, in subtropical forest.

69b. Marasmius variabiliceps var mesites Singer, Sydowia 18: 344. 1965. TYPE. Singer B 1210, from Bolivia.



Marasmius 119

Pileus rusty "feuille morte" M&P with margin pale 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. Lamellae pale cream with broad brown- margined collarium, 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. MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Rio Yariza, 16 Feb 1956,

Singer B 1210 (LIL), type. ILLUSTRATION. Singer (1965) p1. 8, fig. 26.

69c. Marasmius variabiliceps var tucumanensis Singer, Sydowia 18: 344. 1965. TYPE. Singer T 1290, from Argentina. Pileus "cognac," "cookie" M&P on marginal portion, dried between "clove" and

"Mandalay" M&P (deep brown with chestnut-ferruginous tinge) with a striking white zone around a central small blackish dot, the dot appearing papillate gray in dried material, glabrous and smooth in the middle, sulcate in the pigmented zone of the margin, convex, depressed in center, 2-8 mm broad. Lamellae white with edge and collarium either brown or almost concolorous with sides, 13-17, rather distant, broad, collariate. Stipe black, glabrous and smooth, insititious, shining, equal, setose, 18-35 X 0.1-0.4 mm; long dark rhizomorphs associated with carpophores but 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 epicuticular elements. 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 subtropical forest, 1000 m altitude. MATERIAL STUDIED. ARGENTINA. Tucuman: Chicligasta: Rio Cochuna, 20 Feb 1951,

Singer T 1290 (LIL) type; Quebrada de los Sosas, 1100 m alt, 12 Apr 1961, Singer T 3705 (BAFC). ILLUSTRATION. Singer (1958a) fig 44.

69d. Marasmius variabiliceps var separatus Singer, 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 maintains itself as a flat gray disc with pallid star- like fibrils around it, sulcate in the colored zone, convex, white zone and central dot depressed, 1.5-4 mm broad. Lamellae white, edges brown, collarium usually at least brown spotted, rarely even edges, not distinctly discolorous, subdistant (11-14, mostly 14 in larger mature 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




pileus with Rotalis-type elements, setulae ochraceous brown, short, obtuse. Otherwise like type variety. On stem of dead vine in montane tropical forest. MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Carmen Pampa, 26 Feb 1956, Sing-

er B 1547 (LIL) type.

69e. Marasmius variabiliceps var heptachroinus Singer, Sydowia 18: 344. 1965. TYPE. Singer B 1529, from Bolivia. Pileus a saturated orange ferruginous ("Ginger, Kaiser br. 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-shaped and vaguely umbonate in truncate top, 2 mm broad and high. Lamellae white, edges usually not discolorous but in dried specimens at least the collarium distinctly 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 epicuticular elements but mostly hyaline. Hyphae somewhat pseudoamyloid in trama of pileus. Cortical layers: epicuticular elements of the Rotalis-type, main body e g 20 X 1 l , 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. Otherwise like type variety. MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Carmen Pampa, 26 Feb 1956,

Singer B 1529 (LIL), type. ILLUSTRATION. Singer (1965), p1 8, fig 25.

69f. Marasmius variabiliceps var nothotereticeps Singer, Sydowia 18: 344. 1965. TYPE. Singer B 1411, from Bolivia. Pileus "Kobe" M&P (deep rufous) with depressed white umbilical zone and a dark

dot in the center, dried between "cocoa" and "Mohawk" (deep red brown) sulcate to white zone, subcylindric or barrel-shaped, 1.2 X 1.2 mm. Lamellae with a brown-rufous margined collarium but 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 compressed below 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 epicuticular broom cells but more often hyaline. Hyphae of trama of pileus very weakly pseudoamyloid at 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 diverging short (to 2 X 1 j) brown setulae.

On tender dicotyledonous leaf fallen to the ground in montane tropical forest. MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Rio Yariza, 23 Feb 1956, Singer

B 1411 (LIL), type.

69g. Marasmius variabiliceps var derubricans Singer, 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 narrow white or paler often indistinct zone in umbilicus, the white zone surrounding a black spot which is flat and becomes dark gray on drying, barrel-shaped and practically as high as broad, 2-4 mm broad. Lam- ellae white with white to slightly browned (dried) edges and brown-margined collarium, 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 epicuticular elements. Hyphae of pileus pseudoamyloid. Cortical layers: 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. Marasmius baeocephalus Singer, sp nov Fig 52B. TYPE. Singer B 7311, from Ecuador. Pileo brunneo, albo-zonato, atro-maculato, minuto; lamellis brunneo-marginatis,

collariatis; stipite atrocastaneo apice pallido; rhizomorphis nullis. Sporis 6-8 X 3.5-4,; elementis epicuticularibus typi Marasmii rotalis. Ad folia dicotyledonea delapsa in Aequa- toria. Typus in F conservatus est.

Pileus brown when dry, (dried near "Sudan br." M&P) with a white or whitish zone in the umbilicus surrounding a 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. Lamellae white with brownish ochraceous edge 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 accompanied by 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 epicuticular broom cells but all with hyaline main body and somewhat lighter colored (than in the epicutis) setulae. Hyphae of the trama of the pileus 2-8s, broad, subhyaline, pseudoamyloid, with clamp connections; hymenophoral trama 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-

shaped. 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 epicuticularibus typi Marasmii sicci. TYPE SPECIES. Marasmius graminum (Libert) Berkeley.




Key to the Species of Subsection Penicillati

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.1 chr?sochaetes

4. Epicuticular broom cells with narrower, less obtuse and generally more numerous 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

6-12. 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 s. 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 drying (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, between 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 species.

15. Pileus neither fresh nor dried umbilicate, umbonate or papillate, 5-6 mm broad; on fallen leaves (see key II below, "12").

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; spores 6.3-7.7 X 2-2.5(-3),u (see key II "3"). 12. All stipes arising directly from the substratum.

17. Rhizomorphs in part pilose (pilosity caused by thick-walled, obtuse, brownish hairs); pileus light cinnamon when fresh becoming rather deep rusty-chestnut when dried; spores 9.3-9.7 X 4-4.5 u; on dead leaves and petioles in tropical rain forest of the Amazon region. 81. M. conquistensis.

17. Rhizomorphs, if present, glabrous; fungi not combining the characters indicated above.

18. Stipe short and black or almost black; pileus avellaneous-brownish but white around a central black dot (papilla), becoming brownish-orange to light sordid orange-brown on drying; spores 9.5-13.2 X 4.5-6.2,i; rhizomorphs abundant; lamellae 9-10, edges mostly concolorous with the pileus; on leaves. 82. M. boliviae.

18. Not combining these characters. Stipe over 20 mm long, not deeper colored than the margin of the pileus or at least not darker than fulvous or medium brown; pileus cinnamon buff, ochraceous buff, light brown, clay color, fulvous or white; without a white zone around the central dot or papilla (unless entirely white excepting the papilla); spores as above or smaller; rhizomorphs less abundant or absent (but "telepodes" often present); lamellae 9-14; edges and collarium not concolorous with the pileus unless the latter is white on woody material of both Monocotyledones and Dicotyledones.

19. Spores (4-)4.5-6.2j broad. 83. AM pallipes. 19. Spores 7.5-9.5 X 4-4.2 . 84. M. aciculaeformis.

11. Pileus (fresh) partly or entirely brighter colored, orange, rufous, rusty- orange, orange-yellow, salmon-orange, red, pink, purple, red-brown. Key II.

2. Spores three times longer than broad or still narrower. Key II.

Key II

1. Pileus red to brown-red or red-brown or purple when fresh, either so colored when wet or becoming so when drying out in situ. 2. Spores three times as long as broad or narrower (Q73).

3. Spores smaller than 7.7 X 3,; rising from rhizomorphs. 85. M. polycladus. 3. Spores larger or stipe not rising from rhizomorphs but directly from the sub-

stratum. 4. Spores in their great majority over 14p long.

5. On leaves and litter of all kinds, tropical; spores (9-)14-18.5 X (1.5-)2.5-3.3/; lamellae distant (11-15); edge of the lamellae pink or concolorous with the pileus, the latter tending to be a rich deep brown when dried. 86. M. rubromarginatus.

5. On wood in subtropical vegetation of South America; spores still more voluminous or lamellae subdistant (16-17).

6. Lamellae >14. 87. M. marthae. 6. Lamellae <14. 88. M. sanguirotalis.

4. Spores not reaching 14/u in length (see "10" below). 2. Spores less than three times longer than broad (Q < 3).

7. Stipes rising from rhizomorphs; both stipe and rhizomorph with seta-like hairs (if stipe glabrous see no. 80, p 131). 8. Pileus pink without a white zone around the dark purple low umbo; spores

7.5-8 X 2.8-3 U. 89. M. microdendron. 8. Pileus red with a white zone around a white to blackish central papilla; spores

7-9 X 3.5-4.2,u. 90. M. hippiochaetes. 7. Stipe either rising directly from the substratum or from rhizomorpns; in either

case both stipes and rhizomorphs (if present) glabrous. 9. On wood; stipe rising directly from the substratum; pileus purple red, 3-10 mm

broad; spores 4-7,u broad; context of pileus and hymenophoral trama pseudoamyloid, collarium often discolorous (brown). 91. M. xerampelinus.

9. On fallen leaves or small twigs or stems; stipe rising directly from substratum or from dark rhizomorphs; pileus as above or smaller; hymenophoral trama



124 Flora Ncotropica

and pilcus-tra ma 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 sonme stage 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 partially (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 discolorous (concolorous with the margin of the pileus) edge or with intermittently to very weakly discolorous edge; stipe rising from the substratum or from rhizomorphs.

13. Edge of lamellae red; pileus 1-3.5 nimm broad, disc umbonate, blood-red to bright pinkish red. 95. M. edvalliaius.

13. Edge of lamellae eitlier not discolorous or miore often very narrowly and intermittently concolorous with the pileus; pileus 2-6 nmm broad, umbonate-papillate or not; collariumi often indistinct.

94. A1. lputtemansii. 1. Pileus orange, orange-rufescent, salmon-orange, golden yellow, yellow ochlr, ora nge-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 present.

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 leaves and monocotyledonous culms. 5. Spores 9.5-13.2 X 4.5-6.2u; edge of lamellae discolorous (see Key I above, "18"). 5. Spores less broad.

6. Pileus orange-ferruginous, drying brown ("cocoa" M&P); edge of lamellae white and cheilocystidia with hyaline setulae; spores 8.5-11 X 3.5-4.1 M. On dicotyledonous leaves in the Amazonas region. 103. M. robertsonii.

6. Pileus orange with deeper colored papilla, sometimes somewhat bleaching in the marginal region; edge of the lamellae intermittently or indistingly discolorous, some cheilocystidia with pigmented setulae; spores 7.5-11(-13) X 3.5-5 p. Mostly on monocotyledonous leaves and culms. 102. M. crinisequi.

1. All stipes rising from the substratum. 7. On wood, generally of Dicotyledones.

8. Edge of lamellae not discolorous. 9. Carpophores accompanied by black rhizomorphs. pileus often rather pale dirty

orange to pale cinnamon buff when wet, without a white zone around the deep brown central papilla. 10. Spores 9.5-12.3u long. 104. M. dicotyledoneus. 10. Spores usually shorter (see "11" and "12" below).

9. Carpophores not accompanied by black rhizomorphs. pileus orange with or without a white zone around a darker papilla or umbilicus; spores 7.5-10l long, about twice as long as broad or slightly broader (if spores more elongated compare no. 105 (M. rhizomorphogeton).

106. M. graminum var schini and var subalpinus. 8. Edge of lamellae discolorous.

11. Spores 4.7-5.3, broad. Southern South America. 107. M. praecox. 11. Spores 3.5-4.3,u broad. Florida to Amazonas. 108. M. ruforotula.

7. On dicotyledonous and monocotyledonous leaves, on culms of Bambuseae and twigs of other Monocotyledones, not on woody material of dicotyledonous trees and shrubs, rarely on stems and fronds of Pteridophyta. 12. Edge of lamellae not discolorous; on monocotyledonous hosts.

13. Spores relatively elongated, 10-13.2 X 3.8-4.2p; pileus rather pale and dull orange ocher, dried deeper colored (rufescent). Amazonas region (see key II above "18").

13. Spores either smaller or broader; pileus well pigmented orange to ferruginous orange or orange red, sometimes pallescent in age. Subtropical, tropical and subtropical-montane zone of both hemispheres.

106. M. graminumn var graminum and var culmisedus. (Note: If on Bromeliaceae, compare A bromeliacearum, Key II, "11" no. 96)

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 shrubs.

15. Spores 9-11.8 X 4.8-6.2 or 9-11.5 X 3.5-4.3 ; pileus bright "ferruginous" (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; rhizomorphs either none or scarce and inconspicuous; lamellae 12-13; collarium white. 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. Spores smaller (cf also M. praecox no. 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; lamellae 7-11. On dicotyledonous leaves.

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.




71. Marasmius peckii Murrill, North American Flora 9: 254. 1915. Fig 52 P. TYPE. M. E. Peck, from Belize. Pileus whitish, sometimes discoloring to gray but mostly remaining whitish when

dried, with blackish or dark chestnut-fuliginous papilla, glabrous, sulcate, hemispherical, unbilicate with a dark, rather low papilla in 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 accompanied by dark rhizomorphs, 20-30 mm long; some sterile stipes (telepodia) present.

Spores 5.8-6.8 X 1.8-2.8 p, oblong to fusoid, smooth, hyaline, inamyloid. Hymen- ium: basidia 15.3-16.7 X 5.5-6.5 ., 4-spored; cystidia none; cheilocystidia like the epicuticular broom cells. Hyphae with clamp connections, parallel and strongly pseudoamyloid in the trama of the stipe. Cortical layer: 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, subacute to obtusate, hyaline.

On dead dicotyledonous leaves in British Honduras. MATERIAL STUDIED. BELIZE (British Honduras), 1906, M. E. Peck (NY), type.

72. Marasmius chrysochaetes (Berkeley & Curtis), Jour. Linn. Soc. Bot. 10: 297. 1868. Crinipellis chrysochaetes (Berkeley & Curtis) Singer, Lilloa 8: 461. 1942. TYPE. Wright 162, from Cuba. Pileus white, dried tending to "pinkish buff' Ridgway, and developing sometimes

a brown umbo, sulcate, glabrous, convex, umbilicate, with a small umbo in the umbilious, 1.2-2 mm broad. Lamellae white, distant (about ten through lamellae, no

lamellulae), rather broad, 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:

basidia 4-spored, basidioles fusoid; cystidia none; cheilocystidia like the epicuticular broom cells but generally hyaline and inamyloid. Hyphae of pileus-trama and hymenophoral trama hyaline, thin, rather thin-walled, inamyloid with clamp connections; those of the stipe parallel, pseudoamyloid. Cortical layers: epicutis hymeniform or subhymeniform, broom cells of the Siccus type, 14-28 X 7-9u, with apical obtuse appendages which are often finger-like and 2-3/u broad, entire wall firm to thick (to 3.5 ,)- walled, pseudoamyloid.

Subcespitose on dead leaves of the evergreen kind. Cuba. MATERIAL STUDIED. CUBA. Wright 162 (FH) type. I have not studied part of the type deposited at Kew but Dennis indicates

similar characteristics as given above although he finds the spores slightly narrower

(9-10 X 3-3.5g). ILLUSTRATIONS: Singer (1942) fig 1, g. Dennis (195 lb) fig 3.

73. Marasmius aspilocephalus Singer, Sydowia 18: 334. 1965. TYPE. Singer B 2214, from Bolivia. Pileus white, in umbilicus pale buff or pale cinnamon, weakly to distinctly

sulcate where white, glabrous without a dark dot in the center, not papillate, convex, umbilicate, 4-7 mm. broad. Lamellae white, with white edges, distant (7-9), equal or with 1-2 lamellulae intermixed, 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; cheilocystidia like epicuticular broom cells. Hyphae of pileus trama hyaline, with clamp connections, inamyloid or almost so. Cortical layers: epicutis of pileus hymeniform, its elements of the Siccus-type, hyaline with hyaline apical setulae, few with golden melleous setulae, main body e g 9-11 X 8-1 1 , setulae up to 5 A. long, obtuse.

On leaves and small culms of Bambusa sp. MATERIAL STUDIED. BOLIVIA. Pando: Conquista, 25 Mar 1956. Singer B 2214 (LIL)

type. ILLUSTRATION. Singer (1965) pl 9, fig 32

74. Marasmius pallenticeps Singer sp nov Marasmius trichorhizus var pallenticeps Singer. Sydowia 18: 178. 1965 (nomen nudum).

TYPE. Singer M 1123, from Argentina. Pileo albo vel pallido, juventute papilla atrobrunnea prominente ornato; lamellis

albis, collariatis; stipite aut e substrato aut e rhizomophis nato. Sporis 7-8.5 X 2.7-4.5 ,; elementis epicuticularibus typi Marasmii chrysochaetis praesentibus. Ad ramos emortuos in silva subtropicali argentinensi. Typus in LIL conservatus est.

Pileus white or pallid, in dried condition stramineous whitish, only a few caps now brown, glabrous, weakly sulcate, with a dark brown prominent papilla in young specimens, later more repand and maintaining the dark papilla or papilla becoming paler, 1-2 mm broad, 0.5-1.2 mm high. Lamellae white with apparently pallid edges, distant (7), collariate. Stipe copper brown, reddish brown, glabrous, smooth, shining, insititious, or pallid and rising from a copper brown shining rhizomorph, the rhizomor-

phogenous stipes 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, insititious where inserted in the substratum. Context white, inodorous.

Spores 7-8.5 X 2.7-4.5 ,, ellipsoid, smooth, hyaline, inamyloid. Hymenium: not studied. Hyphae of pileus trama thin-walled with clamp connections, inamyloid or almost so; of stipe strictly parallel, thick-walled, strongly pseudoamyloid, also the

hyphae of the rhizomorphs. Cortical layers: epicutis of pileus hymeniform or subhymeniform, consisting of broom cells with erect or somewhat spreading-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 intermixed with coarse elements of the Chrysochaetes-type, i e thick-walled and not setulose but with coarse broad verrucose to finger-like processes, or nodose, versiform, hyaline and strongly pseudoam- loid, but strongly colored brown in center (papilla), often forked.

On dead branches in subtropical forest, gregarious. MATERIAL STUDIED. ARGENTINA. Misiones: Frontera, General Manuel Belgrano near

San Antonio, 19 Apr 1957, Singer M 1123 (LIL), type. ILLUSTRATION. Singer (1965) pl 9, fig 33.

75. Marasmius conicopapillatus Hennings, Engler Bot. Jahrb. 22: 100. 1895. TYPE. Dusen, from Cameroons (type lost). Pileus white, eventually and mostly in the herbarium becoming light beige to

brown (e g "Arab" M&P), with the center from the beginning brown to black, glabrous, sulcate, center smooth, papillate, convex, later papilla in an umbilicus, 1-6 mm broad. Lamellae white or whitish with white or whitish edge, distant, (6-12

through-lamellae, 0-2 lamellulae), simple, rather broad to broad, collariate. Stipe black, at first with white apex, glabrous, smooth, setiform, simple and insititious




on the substratum, mostly accompanied by 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 epicuticular broom cells. Hypae of the pileus trama and the hymenophoral trama hyaline, with clamp connections, partly and weakly pseudoamyloid, many remaining inamyloid in Melzer's reagent. 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 forest.

MATERIAL STUDIED. BOLIVIA. Pando: Madre de Dios, Las Piedras, 5 Apr 1956, Singer B 2528 (LIL). ZAIRE. Eala, Goossens-Fon2tana 109 (BR); Yangambi, Louis 14935 (BR).

If indeed the American collection is identical with the African collection, the spores must be very variable. It is possible that the American species with spores 8.2-12 X 4-4.5 u (slightly larger) is a constantly larger-spored race (subspecies) but too few collections have been made to separate them under a subspecific name.

IILUSTRATION. Singer (1964b) fig 18, D-E; (1965), pl 8, fig 31.

76. Marasmius nigrobrunneus (Patouillard) Saccardo, Sylloge Fungorum 11: 37. 1895. Androsaceus nigrobrunneus Patouillard, Jour. de Bot. 5: 308. 1891. Marasmius griseoviolaceus Petch, Trans. Brit. Mycol. Soc. 31: 42. 1947.

TYPE. Bon 4397, from Vietnam. Pileus gray (between "Deauville sand" and "turtledove," "smoke brown")

becoming "limestone" M&P, sometimes whitish cinereous, glabrous, sulcate to deep sulcate-grooved to the umbilicus, with a low black papilla or central dot in the umbilicus or a shallow depression, 2-9 mm. broad. Lamellae white, 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, glabrous macroscopically, but sometimes distinctly scabrous-spinulose under 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 smaller on an average, ellipsoid to oblong-ellipsoid, smooth, hyaline, inamyloid. Hymenium: basidia 24-28 X 6-7.8M, 4-spored; cystidia none; cheilocystidia like the epicuticular broom 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. Cortical layers: epicutis of pileus hymeniform (but sometimes detersile) and consisting of broom cells of the Siccus-type, although often rather coarsely diverticulate at apex, main body hyaline and thin-walled or at least in upper portion sordid gray to deep sepia-fuliginous or 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-fuligineous or umber, 2-5.3 X 0.7-2.81, generally rather broad and almost always rounded-obtuse at tip; covering layer of stipe inconspicuous, consisting of small hairs, sometimes ascending at 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 carpophores often on grasses.

MATERIAL STUDIED. TRINIDAD. St. Joseph, 30 Nov 1949, Deinnis 409 (K). BOLIVIA. La Paz: Nor-Yungas Carmen Pampa, 2000 m alt, 26 Feb 1956, Singer B 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: Lagunas de Yala, 14 Feb 1966, Singer T 5127 (F). VIETNAM. Hanoi, Kes6, 31 May 1890, Bon 4397 (FH), type; Demange 6 7 (H). INDIA. Gurdespur, Jaggat Pur, 15 Aug 1941, Sultan Ahrmad 509 (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. Marasmius fuligineorotula Singer, Sydowia 18: 336. 1956. TYPE. Sinlger B 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, primordia only papillate, sulcate when mature, 4-11 mm broad. Lamellae white, 7-10 equal, collariate. Stipe black with white apex, glabrous and smooth, insititious. 5-10 X 0.1-0.2 mm; black rhizomorphs present, but scattered and

inconspicuous. Spores (7.5-)10.2-12.3 X 4.5-6u, somewhat broader than half their length or about

half their length in width, more rarely narrower than half their length, ellipsoid-subfusoid, 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 epicuticular broom cells, but very few with other then hyaline setulae and setulae more clearly of the Rotalis-type. Hyphae of the pileus trama partly weakly pseudoamyloid, with clamp connections. Cortical layers: epicutis of puleus hymeniform, consisting of elements varying in shape and appearance between 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-pedicellate or subglobose-pedicellate, 10-24 X 4-10.5 u, some cells smooth without setulae, the majority with hyaline or spadiceous to blackish-spadiceous setulae of extremely variable shape and size, some forked, some acute, some obtuse.

On dicotyledonous stickes in montane tropical forest. MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Rio Yariza, 16 Feb 1956,

Singer B 1193 (LIL), type. This species is intermediate between the species with Rotalis-type epicuticular

elements and those with Siccus-type. ILLUSTRATION. Singer (1965) p 8, fig 30.

78. Marasmius magnisetulosus Singer, sp nov Fig 53. TYPE. Dumonit et al VE 6206, from Venezuela. Pileo griseo vel brunneolo-griseo in siccis, sub lente velutino, 0.3-1 mm lato, 0.2-

0.5 mm alto; lamellis albidis vel grisello-albis, acie haud discoloribus, distantibus, collariatis; stipite atrobrunneo apice albido-pallido; rhizomorphis absentibus. Sporis 5.5-6.5 X 3-3.5,; hyphis pilei inamyloideis, fibulatis; elementis epicuticularibus cystidiiformi-




bus dimorphis, nonnullis integris. Ad folia dicotyledonea emortua in Venezuela. Ty- pus in NY conservatus est.

Pileus gray or brownish gray (dried), long-sulcate, macroscopically glabrous, under a lens velvety, semiorbicular to campanulate or 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 interlamellar spaces appear like pores, collariate. 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, 2-5 p 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. Hymenophoral trama regular, of subparallel, hyaline hyphae 2-34i broad. 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 appendages which are il their majority apical (two or four) but some lateral, basically of the Siccus-type but more clavarioid or reminding one of the cheilocystidia of Marasmiellus, definitely not of the Rotalis-type. hyaline below, brown above, some entirely brown, some merely mellous, with walls 0.2-0.5, thick, appendages 3-15 X 2-61 , with obtuse tip, (2) 13-35 X 5-7p, rather numerous, entire, not branched or a few with one very short branch near the base, cylindrical ampullaceous or clavate, some finely granular incrusted, 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 structure of the epicutis and the shape of the pileus, small number of

lamellae and small spores are characteristic.

79. Marasmius misionensis Sydowia 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, concolorous umbo in a central depression, 1.7-2 mm broad. Lamellae white or whitish, with white or whitish edges, broad, distant (7), equal, collariate. Stipe brown, then black, dried umber, shining, glabrous and smooth, rising from a slightly thicker rhizomorph, 14 X 0.1-0.2 mm.

Spores 7.5-9 X 44.5,4, ellipsoid, hyaline, smooth, inamyloid. Hymenium: basidia 18 X 5.5,L; basidioles fusiform; cystidia none; cheilocystidia like epicuticular broom cells but not or not strongly pigmented. Hyphae of pileus trama and hymenophoral trama hyaline, 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, characteristically 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 5.5.

On forest debris. MATERIAL STUDIED. ARGENTINA. Misiones: Iguazfu, Refugio Yaguarete, 16 Apr 1957,

Singer M 1047 (LIL) type. ILLUSTRATION. Singer (1965) p1 10, fig 4.

80. Marasmius schultesii Singer, sp nov TYPE. Schultes & Cabrera 12869, 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 rhizomorphis hirsuto-pilosis nato. Sporis 7.5-9 X 4-5.5,u; cheilocystidiis hyalinis vel subhyalinis, ceterum elementis epicuticularibus simillimis; hyphis tramatis pilei inamyloideis; elementis epicuticularibus pilei aurantiacis vel aurantiaco-luteis, rarius subhyalinis, crasse tunicatis. Ad radicem in silva tropicali. Typus in F conservatus est.

Pileus brown, dried cinnamon, glabrous, sulcate, with a central papilla which is concolorous and not surrounded by a pallid zone, later small-papillate in a central de-

pression, dried up to 0.5 mm broad. Lamellae buff-pallid when dried, not discolorous at the edge, about six through-lamellae and no lamellulae, broad, collariate. Stipe reddish brown, glabrous, attached to the much thicker rhizomorph by a node and sometimes also with a mediane node, rarely branched, equal, about 3 mm long and up to 0.1 mm broad; rhizomorphs concolorous, hirsute-pilose all over, about 65-120 X + 0.3 mm, insititious. Context pallid, very thin. Spores 7.5-9 X 4-5.5 j, ellipsoid to more rarely oblong, smooth, hyaline, inamyloid, tending to germinate from the distal point in situ. Hymenium: basidia about 12 X 5p; cystidia none seen; cheilocystidia like the epicuticular broom cells but mostly hyaline or subhyaline. Hyphae of the trama of the pileus and the hymenophoral trama hyaline 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 1.2-1.5M thick wall, more rarely with thin wall, with few to numerous apical setulae, these obtuse, concolorous with the Imain body, 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 ("highland forest" of the Amazon basin).

MATERIAL STUDIED. COLOMBIA. Amazonas, Soratama, Rio Apaporis, between Rio Pacoa and Rio Kananarf about 250 m altitude 3 Jul 1951, R. E. Schultes & I. Cabrera 12869 (BPI, F), type.

The color of the pileus as indicated by the collectors as well as the spore data and thick-walled broom cells suggest affinity with Ml. misionentsis (no. 79); the hairy rhizomorph suggests affinity with M. microdendron and 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. microdendron because of the much broader spores, 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 margin of the pileus.




81. Marasmius conquistensis 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 disappearing and a small black papilla remaining in a central umbilicus or depression when mature, otherwise convex when mature, sulcate over the convex part of the pileus, 2-11 mm broad, 0.7-1.2 mm high. Lamellae white 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, glabrous and smooth, shining, insititious with a simple very minute ring directly on the substratum, about 7-13 X 0.1-0.2 mm; black rhizomorphs present, accompanying the carpophores but 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, cheilocystidia like epicuticular broom 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 brownish, 11-13.8 X 5.3-7 ,, rather thin-walled, cylindric to clavate or subventricose, setulae 3-5.5 X 1.2-1.6p, either hyaline or deep reddish rust brown, obtuse; covering of the rhizomorphs consisting 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 epicuticular elements of Crinipellis, not incrusted.

On dead fallen dicotyledonous leaves and their petioles in rain forest. MATERIAL STUDIED. BOLIVIA. Pando: Manuripi, Conquista, 25 Mar 1956. Sinlger B

2190 (LIL), type. ILLUSTRATION. Singer (1965) pl 10, fig 43.

82. Marasmius boliviae 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 orange to light sordid orange brown (1 l-C-9, further inwards "caramel" M&P). sulcate to disc, convex with depressed center, 2.5-3.5 mm broad. Lamellae white, with edges concolorous 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 rhizomorphs associated with the carpophores, also insititious where arising from substratum, 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 epicuticular elements. Hyphae of pileus and hymenophoral trama hyaline, filamentous, some with somewhat thickened walls, many pseudoamyloid, others inamyloid, all with clamp connectioIls. Cortical layers: epicutis of pileus consisting of a hymeniform layer of broom cells of the Siccus- type, mottled by regularly distributed 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, rather variable in volume, setulae coarse



Marasmius 133

and at times few, numbering from one forked setular up to 12 setulae, 4-12 X 1.3-1.7 ,, rarely reaching up to 2.8s diameter at base, either tapering upwards to a subacute or obtuse tip, or more frequently cylindric with broadly rounded tip.

On the thick central nerve of dicotyledonous leaves (Inga ? edulis). MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Coroico, 12 Feb 1956, Singer B

1098 (LIL), type. This species, in dried condition, has the appearance of Marasmius dicotyledoneus,

I. gramfiumt var schinii, etc. Only few spores were found in the type collection and it may be expected that the maximum-minimum range is still wider than indicated lie re.

Marasmiius boliviae 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.


83. Marasmius pallipes Spegazzini, Anal. Soc. Ci. Argentina 16: 272. 1883. TYPE. Balansa 3379, from Paraguay. Pileus cinnamon buff, ochraceous buff, light brown or clay color, at times slightly

zonate in the declivous portion, or gradually paler 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 differentiated wit] an extraordinarily small 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. Lamellae white, edges and sides white, eventually or dried becoming somewhat paler than the pileus, broad, distant, equal or with 1-2 lamellulae or 1-2 forked ones when mature, 9-13 through-lamellae, collariate. Stipe paler than the pileus, often reaching light chestnut below and pale stramineous above, dried usually almost uniformly pale cinnamon or pale reddish fuscous, or sordid stramineous, smooth and glabrous, shining, insititious to the substratum with a very minute basal disc, at apex often white in young specimens, setose-wiry, equal 18-40 X 0.1-0.7 mm; rhizomorphs concolorous with stipe or black, not regularly observed, accompanied by "telepodes," all glabrous, often branched, mostly scanty. Context thin, white, inodorous.

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 u ellipsoid to ellipsoid-oblong or even slightly subfusoid, smooth, hyaline, inamyloid, tending to germinate on the lamellae. Hymenium: basidia (19-)23-32 X 6.8-8.10.5, , 4-spored; basidioles fusoid; cystidia none: cheilocystidia like epicuticular broom cells but predominantly hyaline to pale melleous with melleous to melleous golden setulae. Hyphae: trama of pileus and hymenophoral trama weakly 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-shaped or 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




(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 material in forest humus, dead twigs, mossy logs in tropical and subtropical forest, densely gregarious to

cespitose. MATERIAL STUDIED. BRAZIL. Santa Catharina: Sao Canisio de Porto Novo, 1928, J. Rick

642 (FH). PARAGUAY. Guarapf, Mar 1880, Balansa 3379 (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. Guerrero 118 (LIL). BOLIVIA. Beni: Vaca Diez, Ivon, 3 Apr 1956, Singer B 2480 (LIL).

This occurs also in Trinidad on Ficus benjamina since material described and illustrated by Dennis (195 la) under the name of M. aciculaefbrimis B & C. appears to be this.

Spegazzini apparently described the material exclusively from dried material, and it is obvious that he misdescribed the colors.

The Salta and Bolivian material differs from the other materials in 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 gradually paler colored (from chestnut to ochraceous buff and light brown and eventually pallescent to pallid), maintaining a

deep chestnut central dot (umbo), and becoming between "gold br." and "Bombay" M&P when dried. The Brazilian material became more gray-brown when dried and showed less of a darker central umbo. It was also less pseudoamyloid. The material from Misiones has very short and broad basidia (19-20 X 8-10.5,).

Whether these 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. Marasmius aciculaeformis Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 297. 1868.

84a. Marasmius aciculaeformis var aciculaeformis TYPE. Wright, from Cuba. Pileus fulvous, scarcely sulcate, convex, 4 mm broad. Lamellae whitish, 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 pileus 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. MATERIAL STUDIED. CUBA. Wright 121 (K, FH), type.

84b. Marasmius aciculaeformis var albus Dennis, Kew Bull. 15: 98. 1961. Pileus white, sulcate, with a chestnut-brown small papilla in the center, 5-6.5 mm

broad. Lamellae white, 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 apparently like Marasm7ius pallipes Spegazzini (no. 82 above). On dead wood, cespitose.



Marasmius 135

MATERIAL STUDIED. Color sketch and notes by Dennis on the type of the variety. VEN- EZUELA. Dto Federal: Caracas Botanic Garden, 6 Sep 1959, Dennis 1023 (K).

85. Marasmius polycladus 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. Lamellae fuscou (dried), with not discolorous edge, distant, broad, collariate. Stipe spadiceous-black, short, about 0.25 mm broad, inserted in a sheet- to volva-like node of the broad black, shining, glabrous rhizomorphs.

Spores 6.3-7.7 X 2-2.5(-3) u, ellipsoid-oblong, smooth, hyaline, inamyloid. Hymen- ium: basidioles fusoid; cystidia none; cheilocystidia like epicuticular broom 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 melleous thin to thick (1.5 ) wall, with melleous to reddish brown setulae, these acicular to rod-like, acute to obtuse, 4.5-6.5 X 1-2 .

On and around dead branches of trees, French Guyana. MATERIAL STUDIED. FRENCH GUYANA. Cayenne, Leprieur (K), part of type.

86. Marasmius rubromarginatus Dennis, Trans. Brit. Myco. Soc. 34: 415. 1951. TYPE. Deznnis 339, from Venezuela. Pileus carmine-purple to brown-red ("caldera," M/P or more red), dried bay-mahog-

any ("falcon, muskrat," "leafmold" "Rembrandt" M&P or even deeper on disc), without a white zone or with a very narrow one or with a purple-black zone 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. Lamellae whitish, eventually pale buff, or cream color from the beginning, with red, eventually often brown edge, distant (11-15 through-lamellae, no lamellulae), 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 rhizomorphs mostly accompanying the 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-fusoid to 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 epicuticular broom cells but sometimes smaller and narrower on the average and more hyaline ones intermixed. Hyphae of the pileus trama hyaline, 2-20, broad, partly weakly to strongly pseudoamyloid, 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 hyaline, 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- ment.

On fallen leaves of dicotyledonous trees in forest, known host: Ficus doliaria. MATERIAL STUDIED. VENEZUELA. Maracay: Rancho Grande, Mt. Periquito, 22 Nov




1949, Dennis 339 (K) type. ECUADOR. Napo, Lago Agrio, 16 May 1973, Singer B 7512 (F). BOLIVIA. Beni: Vaca Diez, 22 km east of Riberalta, near Arroyo Florida, 7 Apr 1956, Singer B 2565 (LIL). BRAZIL. Pernambuco: Camaragibe, 7 Jul 1960, Sinlger B 3282 (BAFC); B 3250 (BAFC); 6 Jul 1960, Singer B 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 underneath the epicutis. Marasmiuns martlae has slightly broader spores and more lamellae; it is lignicolous.

ILLUSTRATION. Singer (1965) pl 9, fig 35; Dennis (1951a) pl 19, fig 4.

87. Marasmius marthae Singer, 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 portion when fresh, disc blackening in dried condition, convex, 4-8 mm broad. Lamellae

pure white, with deep purple edge, subdistant (16-17), equal, broad, ventricose, collariate. 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 broader at 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: cheilocystidia like epicuticular broom 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 manner of 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 subtropical zone. MATERIAL STUDIED. ARGENTINA. Tucuman: Capital, Garden of Institute Migucl Lillo,

14 Jan 1952, Martha Sinlger & Singer T 1730 (LIL), type.

88. Marasmius sanguirotalis Singer in Singer & Digilio, Lilloa 25: 19. 1952.

88a. Marasmius sanguirotalis var sanguirotalis TYPE. Sitlger, T 1240, from Argentina Pileus dark purple with purplish black or black center, glabrous, sulcate, convex,

2-5 mm broad. Lamellae whitish or cream white, edges concolorous with sides or slightly purplish in the immediate neightborhood of the margin of 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 epicuticular broom cells, otherwise like these. Hyphae of trama of pileus with clamp connections, 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

MATERIAL STUDIED. ARGENTINA. Tucuman: Quebrada de Lules, 18 Feb 1951 ,Singer T 1240 (LIL, MICH), type (LIL-portion fragmentary and deteriorated); Near Tapia, T 1233 (LIL, paratype.

In some specimens, under a good dissecting microscope, there are traces of discolored (red) gill edge in parts of several lamellae 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-trama of the former and in the habitat.

88b. Marasmius sanguirotalis var paucilamellatus (Singer) Singer, comb nov M1arasmius nmarthae var 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," "Egyptian r.," 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. Lamellae whitish 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 gardens in the subtropical zone. MATERIAL STUDIED. ARGENTINA. Tucuman: Capital, 20 Feb 1955, Singer T 2190

(LIL), type; Sierra San Javier, Parque Aconquija, 17 Feb 1955, T 2179 (LIL), paratypes; Ciudad Universitaria, 15 Dec 1956, T 2853 (LIL); 1 Jan 1960, T 3471 (LIL).


89. Marasmius microdendron Singer, 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. Lamellae whitish, with whitish edge, distant (8-9 through- laiellae, no lamellulae), rather broad, collariate. Stipe stramineous-umber, rather light colored, insititious and without nodes rising from rhizomorphs, with sparse hyaline hairs, 1-2.5 X 0.05-0.1 mm; rhizomorphs dendroid-ascendant and somewhat thicker than stipes, 30-35 mm long, otherwise like stipes but eventually glabrescent. Context very thin, inodorous.

Spores 7.5-8 X 2.8-3 p, elongated, smooth, hyaline, inamyloid. Hymenium: cystidia none; cheilocystidia 11-21 X 6-10, hyaline, otherwise like the epicuticular broom

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-ventricose to

broadly clavate, usually entire, with thicker (up to 1.7 p) melleous wall, with apical mucro or with several coarse apical appendages, but more frequently without appendages and broadly rounded above.

On dead dicotyledonous leaves in rain forest. MATERIAL STUDIED. BRAZIL. Amazonas, 30 km N of Manaus, 19 Jul 1960, Singer B 3448

(BAFC).




90. Marasmius hippiochaetes 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, hemispheric to 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. Lamellae white with red edges and red collarium border, 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-umber when fresh, dried sometimes paler (almost stramineous), shining, but not glabrous in well preserved material, except sometimes at the originally white apex, with minute setae pilose, glabrescent and smooth, arising from a minute disc-like node (from a rhizomorph) or more rarely from the substratum, usually several carpophores branching off a rhizomorph in botryoid arrangement or unilaterally (the latter if rhizomorph is creeping or ascending) 3-120 X 0.1-0.2 mm; rhizomorphs like the stipes, also pilose, glabrescent ramose, often

very long, often marked at intervals with 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 epicuticular broom cells. Hyphae in pileus trama filamentous, thin- walled, slowly distinctly pseudoamyloid, in stipe apex strongly pseudoamyloid and parallel. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the

Siccus-type, main body cylindric to clavate or vesiculose-oval, lobate-setulose at 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's reagent, 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 American rain forests.

MATERIAL STUDIED. SURINAM. Bartica, 20 Aug 1937, Sandwith 1580 (K). VENEZUELA. Bolivar, Chimauta massif, 1100-1700 m alt, 30/31 May 1953, Stev'ermark 15605 (NY). BRAZIL. Amazonas: Rio Uaupes, 30/31 May 1853, Spnice (K), type. BOLIVIA. Beni: Vaca Diez, Guayar- amerin, 9 Mar 1956, Singer B 1762 (LIL); 14 Mar 1956, Singer B 1996 (LIL); 17 Mar 1956, Singer B 2069 (LIL).

ILLUSTRATION. Dennis (1951b), fig 9 (above).

91. Marasmius xerampelinus Singer, sp nov Fig 56. TYPE. Singer M 8961, from Mexico. Pileo purpureo vel purpureo-rubro, umbilicato, in umbilico saepe papillato, 3-10 mm

lato; lamellis albis, ad aciem albis vel prope marginem pilei hoc subconcoloribus, distantibus, latissimis, collariatis, stipite nigro vel nigro-brunneo, acie albo demum ambrino, glabro; rhizomorphis praesentibus sed stipiti haud affixis. Sporis 8.5-11 X 5.5-7.5 ,; hyphis tramatis

pilei et hymenophori pseudoamyloideis, fibulatis; elementis epicuticularibus eis Marasmii sicci analogis, nonnullis crassitunicatis, nonnulis pigmentatis. Ad lignum emortuum in



Marasmius 139

Mexico. Typus in F conservatus est. 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 umbilicus, 3-10 mm broad. Lamellae white, with white edges or the edges somewhat concolorous with the pileus near the margin of 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 substratum but accompanied by somewhat thinner (than stipe) black rhizomorphs creeping 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, inamyloid. Hymenium: basidia 26-33 X 9-11 p, 2-4-spored, rarely one or another 5-spored; basidioles fusoid; cystidia none but there are occasionally some cheilocystidia (broom cells) away from the edge proper; cheilocystidia not making the edge strictly heteromorphous but intermixed with basidia and basidioles, near margin of pileus mostly like the

epicuticular broom cells, otherwise similar but hyaline. Hyphae of the pileus-trama and the regular hymenophoral trama hyaline, with clamp connections, not gelatinized, strongly pseudoamyloid, those of the stipe parallel, also strongly pseudoamyloid. Cortical layer: 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 pseudoamyloid with thin to thick wall (0.2-0.9 ,) hyaline to reddish yellow or golden cinnamon, the pigmented ones distributed in such a pattern that the scalp preparation appears mottled, with hyaline to concolorous setulae, these 2.8-6 X 1-2.7 1, rod-shaped or conical and subacute.

On dead wood in tropical forest. Mexico. MATERIAL STUDIED. MEXICO. Chiapas, between Finca Sospiro and El Pozo, 4 Mar 1969,

Singer 11 8961 (F), type; M 8962 (F).

92. Marasmius hiorami Murrill, North Amer. Flora 9: 256. 1915.

92a. Marasmius hiorami var hiorami TYPE. Hioram, comm. Johnston, from Puerto Rico. Pileus fresh red with darker umbilicus zone, without a white or cream colored or

at all paler zone around the central papilla, glabrous, sulcate over half the radius of 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 apparently white or whitish when fresh but darkening somewhat in the herbarium, with not discolorous edge, distant (about 12 through-lamellae), rather broad, adnate to an indistinct or narrow collarium. Stipe fuliginous to blackish, at first with white apex, setiform, smooth, glabrous, insititous on the substratum accompanied by but not attached to black rhizomorphs and sterile stipes ("telepodes") 20-30 X 0.1-0.2 mm. Context thin.

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 epicuticular broom cells but less pigmented. Hyphae of the pileus trama inamyloid or vaguely pseudoamyloid in parts; those of the apex of the stipe deep purple, pseudoamyloid. Cortical layer: 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 assuming the same golden melleous to succineous




pigment as the setulae; setulae 1-3.2 X 0.5-1.2 , golden melleous to succineous, brown in the Melzer's reagent, not initially red; obtuse, sometimes forking; on the umbo of the pileus there are some irregularly clavate broom cells with thick walls turning vinaceous in Melzer's, with few setulae or these replaced by rounded, lobulate protuberances or

simply uneven above. On dead tough leaves of dicotyledonous trees and shrubs. Puerto Rico. MATERIAL STUDIED. PUERTO RICO. San Juan, Brother Hioram7, 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 Polypodiales and sugar cane, dicotyledonous branches, etc. The following variety agrees with the type in almost all characters, certainly in all important characters separating this species from the following related

species excepting the spore length.

92b. Marasmius hiorami var austroamericanus Singer, var nov TYPE. Singer B 2499, from Bolivia. Sporis minoribus a Marasmio hiorami var hiorami differt. Typus in LIL conserva-

tus. Pileus purple red ("Algerian r.") 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 herbarium tend-

ing to become deep ferruginous or red-brown ("Mandalay" M&P or rustier), the center at first paler or concolorous, later much darker in most specimens, glabrous, sulcate over half the radius or somewhat wider, convex, soon umbilicate. with a prominent papilla which is soon the center of the umbilicus, eventually sometimes relatively small

(papilla appearing as a small umbo), 2-3 mm broad. Lamellae white, later discolored

buffy or brownish, with the edge concolorous or near margin red to brown or vaguely

intermittently red (dried) mostly not at all discolorous (edge at first white), subclose to distant (7-13 through-lamellae), broad, narrowly to widely and distinctly collariate.

Stipe succineous to chestnut-fuliginous or black, at first with white apex, shining, setiform, glabrous, smooth, insititious on the substratum, accompanied by but not attached to "telepodes" and/or black rhizomorphs which 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, inodorous.

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 pseudoamyloid contents: basidioles fusoid; cystidia none but some cystidioles like sterile basidioles and occasionally forked sometimes present; cheilocystidia like the epicuticular broom cells but most of them hyaline (pigmented only in the neighborhood of the margin of the pileus or on scarce pigmented dots along the edge where this discoloration is present). Hyphae of the pileus- trama hyaline, with clamp connections, inamyloid, here and in the regular hymenophoral trama filamentous and thin-walled, hyphae of the stipe parallel more or less pseudoamyloid. Cortical layer: 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 Ramaria carpophores, hyaline to brown (melleous, ochraceous brown), near margin of the pileus thin-walled, with setulae apical, erect (but in some cells with somewhat divergent setulae 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, particularly in the center of the pileus some thick-walled and more strongly pigmented (but not red) broom cells with variable shape 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.

MATERIAL STUDIED. BRAZIL. Guapore, Guajara mirim, 10 Mar 1956, Singer B 1820 (LIL). BOLIVIA. Pando: Madre de Dios. Las Piedras, 5 Apr 1956, Singer B 2499 (LIL), type; B 24995 (LTL); Beni: Vaca Diez, Guayaramer'n, 8 Mar 1956, Singer B 1704 (LIL); La Paz: Nor- Yungas, Carmen Pampa, 26 Feb 1956, Singer B 1483 (LIL).

ILLUSTRATION. Singer (1965) pl 10, fig 40 (as AI. aripoensis).

93. Marasmius aripoensis (Dennis) Singer, Sydowia 18: 188. 1965. Marasmlius trichorhizus var aripoensis Dennis, Trans. Brit. Mycol. Soc. 34: 417. 1951. TYPE. Deinnis 231, from Trinidad. Pileus mahogany red (illustration corresponds to "mohawk" M&P) with prominent

black umbo which is later in a shallow depression, otherwise convex, about 2 mm broad. Lamellae pale 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, white.

Spores 6.8 X 3 #, elliptical (appearing to be oblong) smooth, hyaline. Cortical layer: epicutis of pileus consisting of pear-shaped broom cells about 7 u across, with short dark red processes.

On fallen leaves of Inga and other trees in forest. Trinidad. MATERIAL STUDIED. (seen but not sectioned). TRINIDAD. Cerro de Aripo about

2000 feet alt, 23 Oct 1949, Dennis 231 (K), type. No black rhizomorphs are mentioned and the dark red setulae of the epicuticular

broom cells as well as the much smaller spores would differentiate this from the preceding species with which Singer (1965) identified it earlier. It differs from Marasmius edwallianus forma simplex in the colors of both pileus and lamella edges. It appears to be closest to Marasmius puttemansii (the type of which was seen by Dennis) but differs in the absence of a cream or white zone around the papilla which is also more

prominent here than in that species. Furthermore, the spores appear to be different from those of all forms of M. puttemansii known to me and the absence of black

rhizomorphs would also distinguish this species. It is difficult to judge whether the color of the margin of the pileus is similar to

that of Ml. putteiiansii since a purplish red stage in fresh condition, characteristic for both Ml. hioraiii and M. puttenmansii, is not mentioned by Dennis. On the other hand, Dennis (1951a) draws a 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 anld until further fresh material has been collected and more spores been measured, 1 maintain M. ari- poeilsis as a species independent of M. puttemanssii but cannot exclude the possibility that it might be a variety of the latter.

ILLUSTRATION. Dennis (1951a) pl 19, fig 16.

94. Marasmius puttemansii Hennings, Hedwigia 43: 203. 1904. Marasimius oligocladus Singer, Sydowia 18: 190. 1965 ad interim (not validly published).




94a. Marasmius puttemansii var puttemansii Marasmiuls puttemlansii Hennings var subtricliorlhizus Singer, Sydowia 18: 352. 1965. TYPE. Puttemans 860, from Brazil. Pileus "briarwood" M&P (purple-red to red-brown) when drying in situ often be-

coming brown with a whitish to cream zone originating in mature specimens because of cracking of the surface layer around a gray to blackish dot (papilla) but this pallid zone often only on one side of the papilla or with minute "briar wood" spots or obsolete in the herbarium becoming reddish brown or bay at least in part, sulcate, convex, soon umbilicate with or without a small umbo or rather low to medium sized papilla which is eventually in the middle of the umbilicus, 2-5 mm broad. Lamellae white 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, collariate but the collarium sometimes indistinct, narrow and (dried) appressed to the stipe. Stipe dark brown to brown-black with white apex, glabrous, smooth, insititious either on the black rhizomorphs which often accompany the carpophores or 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 epicuticular broom cells but often most of them hyaline. Hyphae of the pileus-trama hyaline, with clamp connections, from slowly and weakly to distinctly pseudoamyloid; 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 Clusiaceae and Myrtaceae, in Brazil) west to the upper Amazonas.

MATERIAL STUDIED. BRAZIL. SSo Paulo, Serra da Cantareira, 21 Mar 1904, Puttemnans 860 (RBR) part of the type collection; Pernambuco: Camaragibe, 7 Jul 1960, Guanabara, Rio de Janeiro, Jaraim Botanico, 28 Jan 1961. Singer C 3165 (F). ECUADOR. Napo, Shushufindi, 13 May 1973, Singer B 7394 (F).

There is a photograph of the type specimen at USDA herbarium in Beltsville (BPI).

94b. Marasmius puttemansii var oligocladus Singer, var nov Mlarasmnius oligocladus Singer, Sydowia 18: 190. 1965 ad interim. TYPE. Singer B 2330, from Bolivia.

Sporis latioribus a typo specieis differt. Typus in LIL conservatus est. Pileus reddish brown 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 umbilicate in age or umbilicate already in the young state, with or without a very low papilla or small umbo, 2-6 mm broad. Lamellae white or whitish, drying cream, white or with a very thin pigmented (concolorous with margin of pileus) line, often intermittently so along the edges, distant (8-1 1 through-lamellae, no lamellulae), broad, collariate 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 epicuticular broom cells but mostly hyaline. Hyphae of the pileus-trama hyaline, with clamp connections, with thin or rather thin wall, dis-

tinctly pseudoamyloid; hyphae of the stipe parallel, pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform, broom cells of the Siccus-type, main body clavate and sometimes above broadened laterally, more rarely ventricose of ampullaceous 9-16 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. MATERIAL STUDIED. BOLIVIA. Beni, Vaca Diez, Riberalta, 29 Mar 1956, Singer B

2330 (LIL), type. BRAZIL. Pernambuco: Camaragibe, 7 Jun 1960, Singer B 3251 (BAFC). In both varieties (which differ mainly in the breadth of the spores), the number

of stipes attached to a rhizomorphs rather than the substratum varies 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 character in this group, I now combine var subtrichorhizus with var puttenansii.


95. Marasmius edwallianus Hennings, Hedwigia 39: (135). 1900. TYPE. Edwall, from Brazil (holotypus lost, syntypus see below). Pileus blood-red, remaining so or fading to brown-red in the herbarium, glabrous,

sulcate, cubic to hemispherical or campanulate, papillate, often flattened around the

papilla, 0.8-2.5 mm broad. Lamellae white with red ege, distant, (about 11 through- lamellae, no lamellulae), broad, collariate. Stipe pale brownish to bister, shining, insititious either on rhizomorphs or directly on the substratum ("forma simplex"), and then relatively long, otherwise 3-6 X 0.1-0.2 mm; rhizomorphs glabrous. 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 epicuticular broom cells. Hyphae pseudoamyloid. Cortical layer: epicutis of pileus hymeniform, 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-shaped to 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. MATERIAL STUDIED. BRAZIL. Bahia, Torrend 56 (BPI); SSo Paulo, Alto de Serra,

Edwall (FH) syntype; Rio Grande do Sul, Theissen (FH), the latter often with white lamella-edge (W), thus doubtfuL

96. Marasmius bromeliacearum Singer 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 leaving a pallid zone around the dark spot in the umbilicus), but dried pileus virtually unicolorous ("hazel," "harvest" or 13 K 10 M&P) excepting the more ochraceous zone toward papilla (in some



144 Flora Neotropica'`

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. Lamellae white or whitish, distant (9-11 through lamellae), rather broad with white or whitish edge and collarium. Stipe sordid ochraceous brown (13 K 10 to "tortoise shell") to sordid umber, with pallid 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 substratum insititious; 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 similar to epicuticular broom cells, but often few and scattered. Hyphae of pileus filamentous, hyaline, varying from inamyloid to distinctly pseudoamyloid, with clamp connections. Cortical layers: epicutis of pileus consisting of elements of the Siccus-type, main body hyaline, subhyaline or golden melleous, sometimes forked at tip and often hand-shaped, clavate to subcylindric if 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 Bromeliaceae in subtropical forest. 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. Marasmius megalospermus Singer, 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, campanulate to convex with steeply declivous sulcate margin and rather high, 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: consisting of basidia; cystidia none seen. Hyphae of pileus often slightly thick-walled, moderately strongly but distinctly pseudoamyloid, with clamp connections. Cortical layers: epicutis of pileus hymeniform, broom cells of the Siccus-type, but sometimes slightly transient towards the Rotalis-type, main body cylindric, clavate or vesiculose, sometimes with geniculate pedicel, hayline or brown, generally thin-walled, 10-14 X 7-9 u, setulae apical but sometimes obliquely-erect immediately below apex, brown, 2-4 X 1-1.2 2.

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. Marasmius chrysocephalus Singer, sp nov TYPE. Singer B 1936, from Bolivia. Pileo aureo-flavo, siccando ochrascente, primum conico, papillato, dein in umbilico

minute umbonato, papilla umboneque nigro, 2-4 mm lato; lemellis albis, distantibus, co-



Marasmius 145

llariatis; stipite nigricante vel fuligineo, primum ad apicem albo; rhizomorphis praesentibus sed stipiti haud affixis. Sporis 11.7-15.2 X 3.2-4.2u, elementis epicuticularibus eis Marasmii sicci analogis. Ad folia delapsa nec non ad ramulos minores in hylaea boliviana. Typus in LIL conservatus est.

Pileus "Florida gold" M&P (golden yellow), dried ochraceous or ochraceous buff, glabrous, sulcate, at first conical with a prominent black papilla, later with a small black umbo in a small umbilicus, hemispheric to convex, 2-4 mm broad. Lamellae white, edge not visibly discolorous, distant, 7-12, collariate, rather broad. Stipe blackish or fuliginous, at first with white apex, smooth and glabrous, insititious on the substratum, rarely on a black rhizomorph, 20-30 X 0.1-0.2 mm; rhizomorphs similar to stipe, accompanying the fruiting bodies.

Spores 11.7-15.2 X 3.2-4.2 u, fusoid, inaequilateral in profile, smooth, hyaline, inamyloid. Hymenium: basidia 14-18 X 4.5-5 p,; cystidia not seen; cheilocystidia like epicuticular broom cells but a larger number of hyaline ones and often scattered over certain distances. Hyphae hyaline to pale melleous, in hymenophoral trama almost inamyloid but slowly and weakly pseudoamyloid in 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-walled main body, main body 8-17 X 6-9 M, clavate, setulae 2.5-4.5 X 1-1.3 ,, obtuse or sometimes acute, generally quite distinctly of the Siccus-type but at times slightly divergent on and very near the apex as in M. gtuanensis.

On fallen leaves and small twigs fallen to the ground in tropical forest. MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez 3 km below Guayaramerin, in virgin

rain forest, 12 Mar 1956, Singer B 1936 (LIL), type. This was formerly (Singer 1965) interpreted as identical with Marasmius hiorami

but after our own type studies on the latter it has been recognized as an independent species.


99. Marasmius xerophyticus 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 macroscopically appearing as a minute black central dot in fresh material, glabrous, sulcate, without a white or pallid zone around the black dot, 2 mm broad. Lamellae white, 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, glabrous and 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. Cortical layers: epicutis of pileus consisting of three types of broom cells of the Siccus-type and of irregular elements similar to those of the Rameales-type (Maras- miellus); the latter elongated and erect or curved, with rounded-obtuse irregular nodulose excrescences, all brownish, without typical setulae, sometimes somewhat thick-walled, of variable size; 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 subxerophytic to xerophytic woods.




MATERIAL STUDIED. ARGENTINA. Salta: Gicmes, lower part of Quebrada de Yaquiasme, 7 Feb 1957, Singer T 2948 (LIL) type.


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

00a. Marasmius guyanensis var guyanensis 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," "cadmium y." M&P) when dry (in situ) turning orange-gold brown to ferruginous ("Punljab," "Arabian br.," "ferruginous," "feuille morte" M&P), often collected in this shade when not quite freshly grown, dried a color between "Punjab" and "feuille morte" or near "Arabian br." M&P or yellowish brown and sometimes if poorly prepared reaching a sordid brown color in the herbarium, fresh with a central black dot which in dried material appears as a deep brown to blackish papilla or small umbo prominent in young material but 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. Lamellae white, in age whitish to cream, with the edges concolorous with sides, in dried material eventually sometimes entirely subconcolorous with 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, rarely minutely pruinate (?) at base, insititious directly on the substratum, 10-40 X 0.1 mm, black rhizomorphs not accompanying 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, clavate, hyaline: basidioles fusoid; cystidia none; cheilocystidia, broom cells of the Siccus- type, hyaline to yellowish hyaline, otherwise like the epicuticular broom cells. Hyphae of subhymenium, hymenophoral trama and trama of pileus hyaline in ammonia, with clamp connections, inamyloid in subhymenium, inamyloid to weakly pseudoamyloid in hymenophoral trama and more distinctly (though weakly) pseudoamyloid in pileus trama accompanied by some oleiferous hyphae; trama of pileus a very thin layer, pseudoamyloidity noticeable only in accumulations of hyphae; hyphae of stipe in strictly parallel arrangement, with numerous clamped septa, weakly pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells generally of the Siccus-type but some with slightly divergent setulae which descend somewhat from the extreme tip of the cell and thus vaguely remind one of some epicuticular elements of the Rotalis- type, clavate, or vesiculose-ventricose and always broadly rounded, even applanate, often forked or cauliflower-like divided in upper portion, 10-25 X 4.8-17 p, either entirely hyaline or hyaline in lower portion or entirely brownish yellow or melleous to deep orange, generally thin-walled but often some thick-walled, setulae 1.5-4 X 0.7-1.5 s, i e rathei thin at base, rather coarse, acute, subacute or obtuse, simple, varying from hyaline to light yellow to deep golden yellow or at times even brownish yellow. Cells of the black-dot-region of the center similar but either without setulae or these replaced by broadened rounded conidia-like appendages, chestnut brown and thick-walled, forming a palisade rather than a hymeniform layer. Stipe surface without a covering layer.



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 determined as Carapa guianensis (sec. Dennis).

MATERIAL STUDIED. TRINIDAD. Arena Forest, 30 Oct 1949, Dennis 255 (K). FRENCH GUYANA (K), authentic type ? BRAZIL. Acre: Sena Madureira, 7 Oct 1968, Prance et al 7904 (NY). VENEZUELA. Sucre: Rio Media, 15 Jul 1972, Dumontt et al VE 5174 (NY). BOLIVIA. Pando: Madre de Dios, Santa Teresa, 28 Mar 1956, Singer B 2305 (LIL); Las Piedras, 5 Apr 1956, B 2527 (LIL); Beni: Vaca Diez, Guayaramer'n, 16-17 Mar 1956, Singer B 2040, B 2101 (LIL); 3 km below Guayaramerfn in virgin rain forest, 12 Mar 1956, Singer B 1935 (LIL).

It was at first thought that Dennis's interpretation of the species is slightly different from the original concept. But after collecting a large number of specimens, it was found that there is enough variation in the shape of the epicuticular broom cells and the Melzer reaction of the trama of the pileus to consider the Trinidad and the French Guy- ana material extremes of the same species and even variety. The color (umber) of the

pileus as indicated by Montagne and copied by others is probably the color of old herbarium specimens, not characteristic for living or well prepared specimens.

ILLUSTRATIONS. Singer (1965), pl 9, fig 39; Dennis (1951a) pl 19, fig 17.

10Ob. Marasmius guyanensis var erythrocephalus Singer, 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 "Brazil r" M&P) when quite fresh and bleaching little (in age "Nasturitium" M&P) pallid around a central black dot but pallid zone very narrow and difficult to observe in dried material, dried pileus fuscous brown. Lamellae white. 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-walled or almost so, hyaline to melleous, setulae 2-5.5 X 1-1.3 u, rarely hyaline, mostly light golden melleous attenuate upwards but tip itself obtuse in most cases, rarely forked.

Otherwise like the type variety. On dead dicotyledonous leaf in virgin rain forest. MATERIAL STUDIED. BOLIVIA. Beni, Vaca Diez, 3 km below Guayaramerrn, 12 Mar 1956,

Singer B 1876 (LIL), type. This is a rare color variant of the species, apparently due to the slight microscopical

differences evident in the structure of the epicutis. ILLUSTRATION. Singer (1965) pl 9, fig 39E.

lOOc. Marasmius guyanensis var guayarensis Singer, Sydowia 18: 336. 1965. TYPE. Singer B 2014, from Bolivia. Macroscopically differing in the fresh color of the pileus which is a deep but pure

orange ("bittersweet o." M&P). Microscopically differing in somewhat smaller spores: 8.5-11.5 X 2.7-4 A. On monocotyledonous branch in tropical rain forest. Otherwise generally like type variety.

MATERIAL STUDIED. BOLIVIA. Beni, Vaca Diez, Guayaramerfn, 16 Mar 1956, Singer B 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 smaller spores.

101. Marasmius trichorhizus Spegazzini, Anal. Soc. Ci. Argentina 16: 173. 1883. TYPE. Caa-Guazfi, Paraguay (no type material found). Pileus testaceous or rufescent, glabrous, hemispherical, 0.5-1 mm broad. Lamellae




concolorous, pliciform, few, ?collariate. Stipe dark, somewhat shining, erect, glabrous, 10-20 X 0.15-0.18 mm rising from thicker rhizomorphs which are creeping and form a very extended (200-500 mm), very loose "rhizomorphoid mycelium," which forms annular joints where the stipes are inserted (description adapted from Spagazzini, 1883).

Microscopical characters see below. On bark of rotting logs in virgin forest Caa-Guazfi, Paraguay. Spegazzini himself says that his material consisted 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 Marasmius equicrinis (M. crinis-equi Mueller ex Kalchbrenner); this species is strongly papillate. Paraguayan material (Piragu, July 1879) is conserved at LPS and does indeeed show a papilla but no spores could be recovered by me or by Dennis (1951a); but Spegazzini found them to be 7-8 X 4-5 p. It is indeed probable that the Paraguayan collections refer to Marasmius crinisequi. There is however a further collection determined Marasmius trichorhizus by Spegazzini. AR- GENTINA. Formosa: Dec 1900, Kermes 723 (LPS). This material has spores 8.2-9 X 2.2-3.5 ,, thus narrower than indicated by Spegazzini. It has epicuticular broom cells of the Siccus-type, 12.3-19.2 X 8.2-11 p, setulae 2.5-3.7 p long, main body pale yellowish, setulae brown. Hyphae of the pileus inamyloid, with clamp connections. The color of the pileus is now brown but its original color is not indicated.

There are two possibilities. Either the original description is taken at face value and the later Paraguayan collection only is taken as identical with the type; in this case Marasmius trichorhizus would become a (lignicolous?) form of Marasmius crinisequi Or else the Formosan material is taken as authentic and Spegazzini's failure to indicate the presence of a prominent papilla and normal (not fold-like) lamellae is considered erroneous or exceptional. In this case Marasmius trichorhizus would be a good species, not fully and partially wrongly described. A decision as to which approach to take, should, I believe, be postponed until an attempt has been made to recollect the species at the type locality, inasmuch as Spegazzini's spore measurements suggest that the type might have been close to our no. 80 (M. schultesii) if the hairiness of the rhizomorphs has disappeared or was overlooked.

102. Marasmius crinis-equi Mueller ex Kalchbrenner, Grevillea 8: 153. 1880. Marasimius equicrinis Mueller in Berkeley, Jour. Linn. Soc. Bot. 18: 383. 1881. Marasmiius gramzintum (Libert) Berkeley & Broome var equicrinis (Berkeley) Dennis, Trans. Brit.

Mycol. Soc. 34: 416. 1951. ? Marasmius repens Hennings, Engler Bot. Jahrb. 23: 548. 1896.

102a. Marasmius crinis-equi var 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 bleaching occasionally, in the

marginal region, glabrous, sulcate, convex, papillate, 1.5-2 mm broad. Lamellae pale orange-yellow with 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; rhizomorphs abundant, long, smooth, glabrous. Context white, very thin, inodorous.

Spores 7.5-11 X 3.7-5 p, ellipsoid to ellipsoid-oblong, few tear-shaped or fusoid,



Marasmius 149

smooth, hyaline, inamyloid. Hymenium: basidia 24.6 X 9,u; cystidia none; cheilocystidia like the epicuticular broom cells. Hyphae of the trama of the pileus hyaline, filamentous, with clamp connections, inamyloid; those of the stipe + pseudoamyloid. Cortical layer: 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 cinnamon, 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 Eastern hemispheres. MATERIAL STUDIED. ZAIRE. Bamania, Eala, P. Stanzer C 325 (BR); Angodia, J. Lebrun

3009 (BR), type. Material similar to the African material cited above has been collected by Petch

in Ceylon (K) and by Dennis on Trinidad (K) but these collections have been checked only macroscopically by 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 described above 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. Marasmius robertsonii 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 marginal zone, at least when mature, at first campanulate to convex and distinctly papillate, later with a small umbo in the umbilicus, otherwise convex, 3-6 mm. broad. Lamellae white, 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 substratum or in the majority of a population, inserted on a black

rhizomorph, 7-20 X 0.1-0.2 mm; rhizomorphs black, glabrous, smooth or with occasional (1-2) nodes and node-bearing ramifications, sometimes insititious in the substratum, sometimes long. Context inodorous.

Spores 10 X 3.5-4.1 ,, ellipsoid-oblong or cylindric, with a slight suprahilar applanation or depression, hyaline, smooth, inamyloid, occasionally with small wart- like processes which are the beginnings of germination on the lamellae. Hymenium: cystidia none; cheilocystidia hyaline, otherwise like the epicuticular broom cells.

Hyphae of the trama of the pileus inamyloid, with clamp connections. Cortical layers: 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. trichorhizus from 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 material of Al. trichorhizus, nor with Spegazzini's.




Marasmius robertsonii differs from the type variety of Marasmius crinis-equi in smaller spores and colors.


104. Marasmius dicotyledoneus (Singer) Singer, comb nov Marasmius equicrinis var dicotyledoneus Singer, Sydowia 18: 336. 1965. TYPE. Singer T 1854, from Argentina.

Pileus fresh pale cinnamon buff in middle portion, white at margin, dried "caramel" except for a minute deep brown central papilla, 1.3-3 mm broad. Lamellae white 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-walled hyphae with clamp connections, weakly partly pseudoamyloid, hyphae of hymenophoral trama inamyloid, of apex of stipe pseudoamyloid. Cortical layers: 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

long). On dead twigs of Compositae and Solanaceae (Baccharis?, Solanum?) in sub-

tropical-montane, rather dry zone. MATERIAL STUDIED. ARGENTINA. Catamarca, Dpto. Andalgala, Rio Potrero, 1400 m

alt, 10 Feb 1952, Singer T 1854 (LIL), type.

105. Marasmius rhizomorphogeton (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 stramineous towards the margin of the pileus, sulcate except in the umbilicate center, but in some weakly, irregularly or shallowly sulcate, in others

strongly regularly sulcate-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. Lamellae cream, with cream to pallid edge, distant (8-11), broad or medium broad. Stipe umber to succineous, reaching blackish in fresh condition, but dried

mostly umber-succineous, glabrous and smooth, insititious on the substratum, simple, equal, setose, 19-32 X 0.1-0.5 mm; accompanied by dark rhizomorphs of the same characters as the stipe, however often with one or two nodes and often branching, the

repent ones at times with fibrils, often very long, where arising from substratum, insititious. Context thin, white, inodorous.

Spores 10-13.2 X 3.8-4.2 p, clavate-ellipsoid to oblong, sometimes subcylindric, thin-walled, smooth, hyaline, inamyloid. Hymenium: basidioles fusoid; cystidia none.

Hyphae of pileus varying from weakly pseudoamyloid to mostly inamyloid with some strands of hyphae appearing very weakly pseudoamyloid in accumulations. Cortical

layers: epicutis of pileus consisting of broom cells of the Siccus-type with some few

slightly transient to the Rotalis-type in occasional collections, i e with setulae slightly diverging although 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 grass culms, sometimes also on tree fern rachis, gregarious, particularly along rain forest brooks but not seen in inundated forest.

MATERIAL STUDIED. BOLIVIA. Pando: Madre de Dios, Las Piedras, 5 Apr 1956, Singer B 2531 (LIL), type.

105b.Marasmius rhizomorphogeton forma arrhizomorphicus Singer, f nov Rhizomorphis nullis. Differs from the type form in the absence of rhizomorphic

filaments. MATERIAL STUDIED. BOLIVIA: Pando, Las Piedras, 5 Apr 1956, Singer B 2518 (LIL), type. The rhizomorphless form is paler than the type form of the variety and it is this

form that was found both on grass culms and ferns. The type form is known only from grass culms.


106. Marasmius graminum (Libert) Berkeley & Broome in Berkeley, Outlines of Bri- tish Fungology p 222. 1860.

Agaricius graminum Libert, Plantae Cryptogamicae Arduennae II no. 119. 1832. Marasmius pruinatus Berkeley & Curtis, Ann. Mag. Nat. Hist. III. 4: 295. 1859. Marasmlius curreyi Berkeley & Broome, Ann. Mag. Nat. Hist. V. 3: 209. 1879. MIarasmius tritici Young, Phytopathology 25: 116. 1925. Marasmiuts culmisedus Singer in Singer & Digilio, Lilloa 25: 193. 1952.

106a. Marasmius graminum var graminum MIarasmius graminurun var brevisporus Dennis, Trans. Brit. Mycol. Soc. 34: 416. 1951. TYPE. Libert 119, from Belgium. Pileus light orange to rufescent "terracotta" to "carrot r." or 1 l-C-l1 M&P and

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, campanulate or convex, papillate or obtuse, soon convex and flattened around the central zone and becoming shallowly umbilicate, 1.5-15 mm broad, in the neotropics mostly about 2-7 mm. Lamellae white to cream, with white edges, distant (8-15 through-lamellae) broad, but sometimes strongly narrowed or evanescent at margin, collariate but the collarium occasionally very narrow or adnate to the stipe, in other carpophores of the same population broad and leaving a somewhat fibrillose, white space between the collarium and the apex of the stipe. Stipe reddish brown or brown, often becoming black from base upwards, mostly at first white at the apex, shining, glabrous, insititious on the substratum and generally not accompanied by 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-oblong but 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 epicuticular broom cells but mostly all or almost all entirely hyaline. Hyphae of the pileus-trama mostly filamentous, hyaline, not gelatinized, slowly and not strongly but undoubtedly at least in their majority pseudoamyloid, with clamp connections; those of the apex of the stipe parallel, distinctly and strongly pseudoamyloid. Cortical layer: epicutis of pileus hymeniform, broom cells of the Siccus type, main body 11.5-16 X 5.2-11 1, mostly hyaline to pale golden and thin to firm-walled, regularly intermixed with more distinctly pigmented and thicker-




walled cells, all with apical setulae 2-5.5 X 1-1.5 i, these spinulose-obtuse, often somewhat curved, hyaline to more frequently golden melleous.

On grass culms, 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, Madamie fi FPraniois (LIL). BELGIUM. Ardennes, Libert 119 (BPI, LE), type.

The spore size alone cannot serve to- distinguish varieties of 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.Marasmius graminum var 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-1 2 M&P) to ferruginous rufous

("henna" to "Kobe"), sometimes bleaching in age or at least in the furrows or in spots, with tan colored to blackish central umbilicus with a knob-like papilla or elevation inside the umbilicus, glabrous, sulcate outside the umbilicus, convex, 3-15 mm broad. Lamellae white in age and on drying reaching pale buff, distant, equal, narrow to rather broad, sometimes not reaching the margin, moderately distant to distant (9-18), collariate but collarium often not well developed or not fully free. Stipe black with white to pallid apex, glabrous and 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, hyaline, inamyloid. Hymenium: basidia 22-35 X 7-9 p, 4-spored; cystidia none; cheilocystidia like epicuticular broom cells but more hyaline ones present, often irregular and with few or no setulae. Hyphae in trama of pileus varying from weakly and only partially pseudoamyloid ( i e some hyphae inamyloid) to inamyloid, hyaline, with clamp connections. 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 5.8-11 1, obpiriform, piriform or cylindric to clavate, sometimes divided like a Clavariella carpophore, hyaline or orange-melleous and then thicker-walled than 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 irregular or forked, some entire (without setulae).

On grass leaves and culms, usually gregarious. Subtropical northwestern Argentina 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 pseudoamyloid pileus- trama, slightly larger spores and the more rusty color.

ILLUSTRATION. Singer (1958a), fig 51.

106c. Marasmius graminum var 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. Lamellae white 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: basidia 4- spored; cystidia none; cheilocystidia like the epicuticular broom cells but all hyaline or a few pale stramineous to golden stramineous. Hyphae of the trama of the pileus hyaline, mostly filamentous, varying from extremely slowly and weakly pseudoamyloid to inamyloid, 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, branches and stems of dicotyledonous plants in Northwestern Argen- tina.

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.Marasmius graminum var subalpinus Singer, Sydowia 18: 336. 1965. TYPE. Silger S 159, from Argentina. Pileus "burnt sienna," umbo in an umbilicus and surrounding highest 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 relatively lower, convex with central depression or umbilicus, sulcate, 3-5 mm broad. Lamellae white with white edge, 8-9, distinctly collariate. Stipe fuliginous to black, 9-10 X 0.1-0.2 mm; no rhizomorphs. Context thin, inodorous.

Spores 7.5-9 X 4.5-4.7 ,, ellipsoid, smooth, hyaline, inamyloid. Hymenium: cheilocystidia like epicuticular broom cells but hyaline. Hyphae of pileus partly slowly weakly 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 stramineous.

On dead wood of Baccharis sp. 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 brevisporus Dennis but differs in the pallid zone of

young specimens, the shorter stipe, fewer lamellae and the habitat. ILLUSTRATION. Singer (1965), pl 10, fig 50.




107. Marasmius praecox 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," "terra cotta" 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 dehydrated to air-dry, so in herbarium) reaching "copper brown" to "Mohawk" with the extreme margin sometimes nearer "chipmunk" or "Arabian br.," 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 appears as 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. Lamellae white, 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 margin of pileus sometimes scarcely visible!), broad, but not ventricose, distant at maturity, equal or more rarely with 1(-2) lamellulae, 8-13, distinctly collariate. Stipe light umber to black with pallid to white apex in

youth, glabrous and smooth, 12-40 X 0.1-0.2 mm, insititious on the substratum but

accompanied by black branched rhizomorphs or rhizomorphs of the color of the stipe which are glabrous and often provided with ring-nodes, insititious on the substratum, rarely stipe insititious on the rhizomorph but not several stipes branching off 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 germinate on lamellae. Hy- menium: basidia 21-24 X 5-6.2 u, clavate, 4-spored; basidioles fusoid; cystidia none; cheilocystidia like epicuticular broom cells. Hyphae: in hymenophoral trama inamyloid or almost so, in trama of pileus varying from inamyloid to very weakly pseudoamyloid, in center of pileus and apex of stipe distinctly pseudoamyloid, in pileus at

maturity varying from hyaline to pale yellowish, of variable diameter, clamp connections present. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, main body rarely golden orange to golden melleous and then often thick-walled, more numerous being the thin-walled cells 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 leaves attached to the main substratum. Identified host: Eugenia. Generally gregarious. Fruiting early in the rainy season: December until mid-February. Basal subtropical forest, here common.

107a. Marasmius praecox 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.Marasmius praecox forma aurantiicolor Singer, 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), dehydrated partially 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 (F).

107c Marasmius praecox forma rubroumbonatus Singer, Sydowia 18: 340. 1965. Differing from the type form in a more orange cinnamon pileus which becomes

reddish orange when half dry, with a red (drying brown) small umbo, dried (in herbarium) "Mohawk" (brown red) with "Arabian brown" margin amd umbo "Mohawk" (M&P).

MATERIAL STUDIED. ARGENTINA. Tucuman: Dpto. Taff, Parque Aconquija, 12 Mar 1949, Singer T 208 (LIL). type.

The spores are generally not larger than 9.7 X 5.3 ,. but in collection T 2852, otherwise typical (f praecox), I observed spores slightly larger: 9-11 X 5-6.2 p. I do not think it possible to separate this form from the others in view of the fact that some collections have intermediate spores e g T 2178: (8-)8.7-10.7 X 4.5-5.3 ,.

The few cases of branching stipe observed may lead to the misconception that this is Al. trichorhizus. This latter species should therefore be compared. Marasmius praecox differs from 1M. robertsonii in being more orange in dried, especially half-dehydrated 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 habitat.

Vlarasmius praecox 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 considerably narrower in the latter than in the former.


108. Marasmius ruforotula Singer, Sydowia 2: 34. 1948. TYPE. Singer F 1456, from Florida (USA). Pileus rufous-red ("rufous," "vinaceous rufous," "ocher red," more rarely between

"Mars Yellow" and "Amber Brown" [Ridgway], "rust, sorollar br" or "chrysanthemum" with 4 G 12 margin, M&P), with deep brown to dark bister center, with sometimes crenate margin, sulcate-plicate, convex, somewhat umbilicate, the zone around the papilla sometimes minutely cracking rivulose, otherwise all glabrous, 4-9 mm broad. Lamellae whitish to cream, the edge concolorous with the sides or narrowly discolorous (concolorous with the margin of the pileus), distant (e g 8-14 through lamellae and 0-2 often venose lamellulae), broad, sometimes somewhat intervenose, collariate but collarium sometimes low and narrow. Stipe cinnamon-fulvous to (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, inodorous.

Spores 7.2-10.2 X 3.5-4.3 E, 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 epicuticular broom cells. Hyphae of the pileus-trama hyaline, with clamp connections, most filamentous but many broad (reaching about 1 2 in diameter), some inamyloid, some pseudoamyloid. Hyphae of




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 rather thin 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 dicotyledonous wood) but also on living roots etc, sometimes among mosses, gregarious in subtropical 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. Marasmius brunneocinctus Singer, sp nov Fig 57. TYPE. Singer M 8982, from Mexico. Pileo atrorufo, rufescente-aurantiaco, maculo atrobrunneo zona brunneolo-brunnea

cincto, 7-8 mm lato; lamellis cremeis acie haud discolori, collario cremeo, distantibus;

stipite nigricante apice albo; rhizomorphis praesentibus sed stipiti haud adhaerentibus

Sporis 6.5-8.5 X 3.5-4 u; hyphis pilei pseudoamyloideis; elementis epicuticularibus crassitunicatis, typi Marasmii sicci. Ad folia dicotyledonea emortua in Mexico. Typus in F conservatus est.

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-bottomed umbilicus and a convex mediane zone,

eventually often recurved at extreme margin 7-8 mm broad. Lamellae cream with not discolorous edge, distant (10 through-lamellae), all equal and simple, broad but sometimes strongly narrowed at margin of pileus and not reaching the 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 substratum but accompanied by thin, black

rhizomorphs, equal, 23 X 0.2 mm; rhizomorphs thin, not very abundant. Context white, thin, inodorous.

Spores 6.5-8.5 X 3.5-4 p, ellipsoid, smooth, hyaline, inamyloid. Hymeniurn: basidia 21-26 X 6.2-7.2 p,, 4-spored; cystidia none; cheilocystidia none seen (possibly eroded). Hyphae in the hymenophoral and pileus-trama as well as in the apex of the stipe hyaline, firm- but not thick-walled, with clamp connections (but many secondary septa observed) distinctly pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform, broom cells of the Siccus-type, more or less strongly pigmented cells evenly distributed and surface in scalp therefore appearing mottled, main body 8.25 X 6-13.5 u, thickwalled and yellow with deep yellow to golden yellow setulae, or thin-walled and hyaline to yellowish hyaline with subhyaline to yellow setulae, the latter apica!, tapering upwards, 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. Marasmius pallidocinctus Singer sp nov



Marasmius 157

11 Oa. Marasmius pallidocinctus var 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 atrocastaneo; rhizomorphis nullis. Sporis 9-10.5 X 3.5-4.3 1; hyphis pilei debiliter lentoque pseudoamyloideis, fibulatis; elementis epicuticularibus typi Marasmii sicci. Ad folia dicotyledonea delapsa in Venezuela. Typus in NY conservatus est.

Pileus (dried) rusty orange with a pallid zone around a central fumosous-fuliginous spot, glabrous, sulcate-plicate, convex, deeply depressed in the center with a low papilla becoming visible in dried material, 3-6 mm broad. Lamellae white with white edge and collarium, subdistant (about 12-23 through-lamellae), broad, collariate. Stipe deep chestnut, glabrous, equal, insititious on the substratum, not rising from nor accompanied by 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: basidia 4-spored; basidioles fusoid; cystidia none; cheilocystidia like the epicuticular broom cells but all hyaline. Hyphae of the pileus-trama yellowish-hyaline to hyaline, weakly and slowly but undoubtedly pseudoamyloid, with clamp connections, filamentous, 4-5 u broad, thin-walled, not gelatinized; hyphae of the stipe parallel pigmented, pseudoamyloid. Cor- tical layer: epicutis of pileus hymeniform, somewhat mottled because of alternatingly paler and darker broom 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, yellowish 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. Marasmius pallidocinctus var latisporus Singer, 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, marginal zone 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. Lamellae white or whitish with white or whitish edges and collarium, distant (12-13 through-lamellae, no lamellulae), collariate. Stipe sordid ochraceous brown (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 accompanied by rhizomorphs, equal, 60 X 0.2 mm. Context thin, inodorous.

Spores 9-11.8 X4.8-6.2 p, ellipsoid, smooth, hyaline, inamyloid. Hymenium: basidia 19-19.5 X 6-6.5 u; cystidia none; cheilocystidia rather scattered, like epicuticular broom cells but less pigmented. Hyphae of the pileus-trama hyaline, 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.




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

111. Marasmius rufomarginatus Singer, 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 "Mars Orange" (Ridgway) becoming "Salmon Orange" or "Mars Orange" 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 concolorous around the brown center, not showing a white zone or, if such a zone is present, it is due to cracking whereby the white trama of the pileus is partly exposed (use lens!), glabrous, sulcate to sulcate-plicate with smooth center, at first campanulate-papillate, then hemispherical to convex, soon rather deeply and often narrowly umbilicate 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. Lamellae white, eventually often becoming cream color, with the edge at least in the marginal half discolorous (concolorous with the margin of the pileus) distant (7-11, more often 10-11 than 7-9 through-lamellae and 0-2 lamellulae), simple or one to several forked near stipe, broad (1-2 mm), collariate but the collarium at first low, later often incomplete and sometimes partly adhering to the apex of the stipe. Stipe chestnut to blackish-brown or 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 strongly elongated and acute in overmature spores, smooth, hyaline, inamyloid. Hymenium: basidia 30 X 8-10.2 ,, 4-spored; cystidia none; cheilocystidia like the epicuticular broom cells or more subhyaline to golden orange. Hyphae of the pileus-trama hyaline, weakly and slowly but distinctly (purple) pseudoamyloid, in the hymenophoral trama 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-shaped to conical, subacute to obtuse, golden 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 subtropical and tropical to tropical-montane forest. Florida to Mexico, south to Colombia. Known host: Nec- tandra coriacea.

MATERIAL STUDIED. U.S.A. Florida: Dade County, Miami, Simpson Park, 23 Oct 1942. Singer F 1219 (FH), type; Matheson Hammock, 30 Oct 1942, Singer F 1219a (FH); 22 Oct 1942, F 1196 and 24 Oct 1942, F 1196a (FH). MEXICO. Chiapas: Tuxtla de Gutierrez, Botanical Garden. 3 Aug 1969, Singer M 8921 (F). COLOMBIA. Valle: Corr. Vergel, Finca La Araucana, 14 Apr 1968, Singer B 6126 (F). ECUADOR. Tungurahua, Rio Verde, 28 Apr 1973, Singer B 7172 (F).

112. Marasmius foliicola Singer, sp nov Marasmius ruforotula var foliicola Singer, Sydowia 12: 136-137. 1958, nomen nudum. Marasmius foliicola (Singer) Singer, Sydowia 18: 205. 1965 "ad interim." TYPE. Singer T 2147, from Argentina. Pileo aurantiaco, in umbilico nigro-maculato, 4-7 mm lato; lamellis albis, acie aur-

antiacis, distantibus, collariatis; stipite atrocastaneo vel nigro, ad apicem primum saepe albo, 15-75 X 0.1-0.5 mm; rhizomorphis nullis. Sporis 7.3-10 X 4-5 p; elementis epicut-



Marasmius 159

icularibus typi Marasmii sicci. Ad folia monocotyledonea et dicotyledonea delapsa in Argentina. Typus in LIL conservatus est.

Pileus orange (shading from "bittersweet o." to "Tokyo" M&P), and margin eventually bleaching in 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 disappearing altogether when drying out, convex around the umbilicus, 4-7 mm. Lamellae white with orange edge (this discoloration at times discontinuous and not very conspicuous or in some caps limited to the region near the margin of the pileus or to the lamellae edges reaching a higher level, i e broader lamellae, than others, but always more or less distinct), flat to arcuate, distant, 10-13, collariate, rarely incompletely collariate. Stipe in very young and fresh material deep brownish orange with white upper portion, soon dark chestnut or black with or without white apex, glabrous and smooth, insititious directly on the substratum, 15-75 X 0.1-0.5 mm, not accompanied by 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- trama weakly but undoubtedly pseudoamyloid, with clamp connections. Covering layers: epicutis of pileus consisting of Siccus-type broom cells 13-23 X 5.5-9(-11.7)A, sometimes spreading to 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 subtropical and 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 illustrations of a Mara- smtius found by Dennis on dead Bauhinia and on dead Mlusa leaves respectively, in Caracas and in Cortada de 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, Ziekten Suikerriet Java 1 p 195, Leiden 1898, is possibly this. I have not seen the type.

MIarasmlius fbliicola might possibly be a synonym of Marasmius gordipes 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 described differently and the number of through- lamellae is smaller. I believe therefore that they are different and accept Mlarasmius foliicola as an autonomous species.

Marasmius exustus Berkeley & Curtis from the Bonin Islands has been mentioned as a possible synonym of Marasmius graminurm or M. foliicola. Indeed, Western Hemi- sphere material determined as suchprobably belongs to Marasinius foliicola 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.

Marasmius sect Sicci Singer, Mycologia 50: 106. 1958. TYPE SPECIES. Maraslius siccus (Schweinitz) Fries. Pileus rather small 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-




smius inaequalis) strongly pseudoamyloid in pileus and stipe; stipe centrally attached, growing from leaves or wood etc, always with basal mycelium; black rhizomorphs absent.

Key to the Subsections of Section Sicci

1. Trama of the pileus inamyloid. subsection Inaequales. 1. Trama of the pileus pseudoamyloid. subsection Siccimi.

Marasmius sect Sicci subsect Inaequales Singer, Sydowia 12: 97. 1959. TYPE SPECIES. Marasmius inaequalis Berkeley & Curtis.

Hyphae of the carpophore inamyloid.

113. Marasmius inaequalis Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 297. 1869. TYPE. WVright, from Cuba. Pileus white, center brown ("cookie" M&P), dried entirely like illarasmiuts helvolus

in color, glabrous, sulcate, campanulate to convex, 2-5 mm broad. Lamellae white with white edge, distant or subdistant (10-12 through-lamellae), rather narrow, subconcolorous with the pileus when dried, narrowly adnexed to free, definitely not collariate. Stipe chestnut brown with pallid or white apex, dried chestnut-amber, smooth, glabrous, shining, 4-32 X 0.1-0.2 mm; basal mycelium tomentose or strigose, white, accompanied by whitish mycelial patches and rhizomorphs in many populations. Context very thin, inodorous.

Spores 11-15 X(2-)3-3.3 ,, guttiform-subclavate to cylindric-subfusoid, often miore or less curved or at least the hilar appendage strongly curved inwards, smooth, hyaline, inamyloid. Hymenium: basidia 20-21 X 5.5-6.5 p, 4-spored; cystidia none; cheilocystidia like the epicuticular broom 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. MATERIAL STUDIED. CUBA. Santiago de Cuba, 1857. Wright 114.4 (FH), authentic.

BOLIVIA. La Paz, Nor-Yungas, Coroico, 30 Jan 1956, Singer B 736 (LIL). ILLUSTRATION. Singer (1958a) fig 18.

Marasmius sect Sicci subsect Siccini Singer, Sydowia 18: 343. 1965. TYPE SPECIES. Marasmius siccus (Schweinitz) Fries.

Hyphae of the carpophore pseudoamyloid.

Key to the Series of Subsection Siccini

1. Setae in pileus-epicutis and hymenophore none; stipe glabrous except for the basal mycelium (tomentose or strigose), more rarely very finely pruinate. 2. Cystidia on the sides of the lamellae absent or not clearly differentiated. series Leonini, p 160. 2. Cystidia on the sides of the lamellae present. series Haematocephali, p 201.

1. Setae in pileus-epicutis and/or hymenophore present, or stipe distinctly pruinate, pubescent or hirsute from dermatocystidioid hairs or setae. series Actinopodes, p 236.

Marasmius sect Sicci subsect Siccini ser Leonini Singer, ser nov Cystidiis ad latera lamellarum vix a cheilocystidiis distinctis nec typo cystidiorum

Marasmii sicci - Typus: Marasmius leoninus Berkeley. Cystidia and setoid hairs absent.



Marasmius 161

Key to the Species of Series Leonili

1. Pileus white over most of its surface when young (in part pale cream or cork color at times). 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 mmn broad, 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 fulvous; 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 Marasmiuzs haematocephalus 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 cheilocystidia present, more rarely there is a thin line of discoloration at the very edge which becomes distinct only when dried material is observed 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.




19. Pileus orange to orange rufescent, 5-6 mm broad, hemispheric and obtuse. In xerophytic and subxerophytic vegetation. 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 1 3.8, 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 black.

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 inconspicuous. 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 1 2.5 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 subsmooth, 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.

1 3. Pileus when dried dull brown, ochraceous buff when fresh; edges of lamellae 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 epicuticular 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-scrobiculate 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 umbilicus; 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 central umbilicus; lamellae free to subfree; surface of stipe




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. Marasmius subrotula Murrill, North American Flora 9: 259. 1915. Marasmtius imelinocephalus 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 applanate and 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 sparse to 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: basidia up to 31 X 8.2-13.7 ,, 4-spored; cystidia often present but few and poorly or not differentiated, basidiomorphous (like basidioles, fusoid, or basidia clavate), neither projecting nor deeper rooting, not appendiculate nor distinctly opaque, but sometimes with a very thin, weak incrustation on apex; cheilocystidia like the majority of the epicuticular broom 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-walled main body and hyaline to pale brownish melleous setulae 1.5-4(-10) X 0.8 p;

regularly scattered among these and giving a scalp preparation a mottled appearance, are broom cells of type II with a main body whose wall gradually thickens towards the apex and is concolorous with the setulae, and setulae deep melleous to spadiceous-fuliginous, some broom cells type II with few or no setulae.

On earth or, more often, dung-covered fallen dead branches of dicotyledonous trees and shrubs, particularly Achatocarpus nigricans in marginal and subxerophytic forest in the subtropical zone, densely gregarious to cespitose, sometimes directly and exclusively 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. Marasmius amazonicus Hennings, Hedwigia 43: 183. 1904. TYPE. Lost (see below). Pileus deep purple to deep lilac violet, the disc blackish purple ( e g 56-J/L - 5

M&P) near center with numerous orange brown ("talavera"), near margin pale buffy orange ("capucin buff') or everywhere buff dots which are oval to round and 2-4 mm in diameter, glabrous, deeply sulcate almost to the disc, campanulate-convex to convex applanate, with umbilicate disc, 42-72 mm broad. Lamellae white or buff-cream, broad (6-16 mm broad), ventricose, intermixed with 0-1 lamellulae, not intervenose, with entire, not discolorous edge, free to removed from the apex of the stipe and then leaving un- covered a white, sulcate disc around the extreme apex of the stipe, distant (19-22 lamellae). Stipe deep chestnut to black, paler brown at apex when young, slightly longitudinally striate when dried, otherwise smooth, glabrous, shining, hollow, equal or gradually and very slightly tapering upwards, terete and sometimes somewhat compressed, sometimes arising from 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: basidia hyaline, 4-spored (?); cystidia none; cheilocystidia like epicuticular elements, but all hyaline. Hyphae hyaline, distinctly pseudoamyloid. Cortical layers: epicutis of pileus consisting of broom cells which are hymeniformly arranged; these hyaline to subhyaline seen individually (NH40H) but in thicker layer deep porphyry brown, individual setulae mostly hyaline or subhyaline but here and there some purple black setulae present, total measurements 15-18 X 9-10,, main body often indistinct and reduced, setulae spinulose and subacute, tapering from a 2-2.7,u broad base, 5-9A

long, entirely inamyloid. On trunks and on very rotten wood among leafmold, in small groups, not ces-

pitose. MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, Guayaramerin, 8 Mar 1956, Singer

B 1706 (LIL); 14 Mar 1956, B 1946 (LIL). The species was originally described from Jurua miry, Amazonas, Brazil, collec-

ted by Ule. The type was conserved in Berlin where it was destroyed during the last war. The description however, leaves not the slightest doubt as to the identity of the material. This is the most spectacular species of South America as far as external appearance is concerned. A similar or analogous species from Cameroon, M. superbus Henn., differs in umbonate pileus with rugolose-venose and not dotted surface and, according to Hennings, collariate lamellae, and broader stipe. It belongs in section Globulares.


116. Marasmius tucumanus Singer 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. Lamellae pink, soon becoming white in the edge half, 1.5 mm broad, not collariate, distant, 10-17 through-lamellae and 0-6 lamellulae, free, later adnexed to adnate. Stipe at first entirely purple-red then black towards the base, glabrous, smooth and shining, with a minute disc or socle at base which is bearing the basal




mycelium 33 X 1.5 mm, base reaching 0.5 mm, basal mycelium tomentose, brown, with alternating hyaline 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 attenuated hilar portion which is either straight or curved towards the inside or outside and then assuming a 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 metachromatic nor opaque. Hyphae pseudoamyloid with clamp connections. Cortical layers: epicutis of the pileus hymeniform, consisting of broom cells which are hyaline to reddish brown or melleous, 17.5-20.5 X 7-7.5., mottled in scalp view, setulae 5.5-7.5M long obtuse.

On dicotyledonous, dead, fallen twigs and branchlets of the humid forest in the subtropical zone, solitary or in small groups in the shade.

MATERIAL STUDIED. ARGENTINA. Tucuman, Taff: Parque Aconquija, 13 Mar 1951, Singer T 1460 (LIL) type; T 1477 (LIL) paratype (cf Singer 1965, p 258, footnote!).

117. Marasmius tageticolor Berkeley, Hook. Jour. Bot. 8: 136. 1856. TYPE. Spruce 37, from Brazil. Pileus red (deep blood red, purple red or carmine red, dried "Macracaibo" M&P)

with white or ochraceous-whitish or (dried) ocher radial stripes, glabrous, smooth on the disc, sulcate on the margin, campanulate-convex, later convex-repand and often umbilicate, 4.5-18 mm broad. Lamellae white with white edges, distant, not or scarcely intervenose, narrowly adnexed 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; cheilocystidia like the epicuticular broom cells but all hyaline to subhyaline. Hyphae hyaline in the pileus, with thin to barely firm wall, with clamp connections, pseudoamyloid. Cortical layer: 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 sometimes slightly spreading, acute or subacute, pale melleous to subhyaline, mostly hyaline in the pale-striped zones, deep melleous in the red-striped zones, 5.5-9 X 1.2-2.5 u.

On twigs as well as on coriaceous leaves, fallen to the ground, in tropical forest. MATERIAL STUDIED. MEXICO: Chiapas, between Ixtacomitan and Solusuchiapas, 1 Aug

1969, Singer B 8804 (F). VENEZUELA. Sucre: NW of Irapa, 8 Aug 1972, Dumont et al VE 4300 (NY). BRAZIL. Amazonas: "Panure" (Ipanur6 = Sao Jeronimo), Spruce 37 (K) type.

ILLUSTRATIONS: Singer (1965), pi 14, fig 77; Berkeley, (1856) pl 5, fig 1.

118. Marasmius poecilus Berkeley, Hook. Jour. Bot. 8: 137. 1856. TYPE. Spruce 139 (part), from Brazil. Pileus purplish brown to fulvous brown (dried almost spadiceous) with radial white

to yellow stripes, deeply sulcate to the umbilicate center, glabrous, campanulate-convex and eventually with uplifted margin, 8-25 mm broad. Lamellae deep cream with not discolorous 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 abundant 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 metachromatic in cresyl blue mounts; cheilocystidia like the epicuticular broom cells and mostly hyaline with hyaline to pale fulvous setulae. Hyphae and cortical layer as in the preceding species.

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 preceding one which I did not separate spec- ifically in the past. However, the colors of the stipe and pileus are different in these species, the stipe of I. poecilus perhaps somewhat 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 appearing through drying, and the collector (Spruce) collected them as different species under different numbers, I prefer to describe them separately.

I believe the type of Marasmius poecilus to be a mixed collection. The leaf with five good and three imperfect carpophores which I consider the holyotypes, undoubtedly bears the species described above as M. poecilus. Berkeley indicated the stipe as insititious which it is not. In some carpophores of the type the stipe has a poorly developed basal mycelium but this is always visible under a lens.

119. Marasmius phaeus Berkeley & Curtis, Jour. Linn. Soc. Bot. TYPE. Wright 18, from Cuba. Pileus dark reddish brown (slightly deeper and rustier than "mohawk" when fresh)

with white radial stripes, glabrous, sulcate to the narrow (sometimes subumbonate) disc, campanulate to conic-applanate, 6-9 mm broad. Lamellae on edges concolorous with pileus, sides fresh white, distant, about ten, ascendant, eventually more horizontal, narrow to broad, adnexed. Stipe umber or light date brown when fresh, often darker in the herbarium, glabrous and smooth, shining, setose, subequal, 12-30 X 0.2-0.8 mm; basal mycelium scanty to well developed, strigose, white or whitish, dried sordid grayish 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; cheilocystidia exactly like the epicuticular broom cells. Hyphae pseudoamyloid with 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 diverge somewhat or main body forked spreading to 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 standing live dicotyledonous trees in rain forest, gregarious.

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. Marasmius bambusiniformis Singer, sp nov Fig 59. TYPE. Singer B 7480, from Ecuador.




Pileo ferrugineo, 4.5-7 mm lato; lamellis candidis; stipite 18-30 mm longo. Sporis 15-18.5 X 3.7-5 ,; cystidiolis frequenter praesentibus. Ad folia delapsa mono- et dicotyledonea. Typus in hylaea aequatoriana lectus 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. Lamellae pure white, often becoming brownish white when dried, with edges concolorous with the pileus, narrow to broad, few. Stipe chestnut or ferruginous, at first with white apex, smooth, glabrous, with slightly thickened base which is pubescent 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, arising from the same level as the basidioles; cheilocystidia like the

epicuticular broom cells. Hyphae in the trama of the pileus hyaline and with clamp connections, thin-walled, pseudoamyloid. Cortical layers: epicutis of the pileus consisting 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 grass culms. 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. Marasmius mazatecus Singer, sp nov Fig 60. TYPE. Singer M 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 conservatus est.

Pileus orange ferruginous, deeper rusty-reddish in 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), rather narrow (2 mm), leaving an orange-ferruginous band at the

apex of the stipe from which they are free when dried. Stipe chestnut, dried brown, fresh with white apex underneath the orange-ferruginous band appearing in dried and

dry material, quite glabrous except for the moderately developed strigose-woolly fulvous 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: consisting entirely of basidia and basidioles; cheilocystidia making the edge heteromorphous, in shape like the epicuticular broom cells. Hyphae of the pileus trama hyaline, with

clamp connections pseudoamyloid. Cortical layer: epicuticular broom cells in hymenial arrangement, main body hyaline or orange, with thin to moderately thick wall, 10-28 X 6-8 p, cylindric to clavate, more rarely ventricose, setulae 2-13 X 1-1.8 1, or-

ange-brown, orange-yellow, yellow, in very rare cases absent. On dicotyledonous wood. MATERIAL STUDIED. MEXICO. Oaxaca: San Andres Hidalgo, Sierra Mazateca, 8 Jul 1969,

Singer M 8368 (F), type.



Marasmius 169

122. Marasmius longisporus (Patouillard & Gaillard) Saccardo, Sylloge Fungorum 9: 68. 1891.

Androsaceus longisponts Patouillard & Gaillard, Bull. Soc. Mycol. France 4: 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 margin between "henna" and "chutney" or between "feuille morte" or "rust sorolla" and "Peruvian br" or pl. 13 A 12, M&P), the margin often paler, not or dried indistinctly narrowly radially striped (ochraceous) but distinctly long-sulcate, glabrous, conic to campanulate, often ? umbonate, 4-10 mm broad, 5-6 mm high. Lamellae white, subdistant to 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, macroscopically glabrous, 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,E broad, clavate to fusoid, smooth, hyaline, inamyloid. Hymenium: basidia 20-28 X 8.5-11 -,

4-spored: cystidia none, but some of the basidioles often remaining sterile, fusoid, ampullaceous or 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 hymeniform, 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 rather thin (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 revised description since my earlier description was based only on the type and fresh material from Bolivia. The latter, however, later turned out to be 1. berteroi var berteroi, whereas newly collected Mexican and Ecuadorian material is obviously identical with the Venezuelan type.


123. Marasmius onoticus Singer, sp nov Marasmliu s onoticus Singer, Sydowia 18: 252. 1965, nom nud

Type. Singer T 2995, from Argentina Pileo aurantiaco vel 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; cheilocystidiis et cellulis epicuticularibus typi Marasmii sicci. Ad lignum dicotyledoneum in silvis subxerophyticis. Typus in LIL conservatus est.

Pileus orange to orange rufescent ("talavera", margin "doubloon"), deeply sulcate, glabrous, unicolorous, hemispheric, 5-6 mm broad. Lamellae white, distant, ten, rather




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 epicuticular broom cells; hyphae hyaline, psuedoamyloid. Cortical

layer: epicutis of pileus hymeniform, consisting of broom cells, 15-16.5 X 7-8.5 u, sometimes spreading because of divergence of setulae to a diameter of 9-11 p, setulae 5.5-8.5 X 1.2-1.8u (at base), golden succineous, few hyaline, obtuse orsubacute, spinulose 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. Marasmius rubricosus Montagne, 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 ferruginous in center and deep chestnut brown on margin, glabrous, densely striate, sulcate on margin, applanate with a very obtuse umbo, dried more convex since margin tends to incurve broadly on drying, about 37 mm broad. Lamellae sordid whitish, drying to light brown, narrow, fresh free and moderately broad, drying narrow and subfree, close with numerous lamellae of various lengths. Stipe chestnut, glabrous or subglabrous, about 50 X 1.5 mm; basal mycelium very abundant, tomentose, forming a socle-like broadening of 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 numerous and not distinctly differentiated but occasional cystidioles occurring (ampullaceous to 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 referred here on the strength of Dennis's remarks (1951a, p 420) which would fit our collection perfectly. These remarks refer to authentic material sent to Berkeley by Montagne.


125. Marasmius rhabarberinus Berkeley, Hook. Jour. Bot. 8: 135. 1856. TYPE. Spruce 140, from Brazil. Pileus rhubarb yellow, orange (e.gr. 1 l-D-11 M&P) dried bright orange fulvous (e

g "spice"), smooth or reticulate-rugose, distinctly sulcate-furrowed in marginal two-thirds when mature or dried, glabrous, convex, with or more rarely without a small umbo, 7-31 mm broad. Lamellae pallid, equal or with some very short lamellulae intermixed, not



Marasmius 171

intervenose, rather narrow to rather broad, (up to 4.5 mm broad), free or even remote- free, distant. more rarely sub-distant (15 through-lamellae or 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-clavate to cylindric-fusoid, straight or 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 narrowly clavate, 35-36 X 6-6.8 p, 4-spored; basidioles fusoid; cystidia none; cheilocystidia hyaline to

pale melleous in upper portion, otherwise like epicuticular elements, making the edge of the lamellae heteromorphous or nearly so. Hyphae of pileus and hymenophoral trama hyaline, thin-walled and 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-dotted in scalp preparations. Stipe without any broom cells or hairs.

On dead fallen leaves in tropical and subtropical forest. MATERIAL STUDIED. BRAZIL. Amazonas: Rio Purua (Purus), Trail 161 (K), authentic;

"Panure" (Ipanure = Sao Jeronimo), Spnuce 140 (K), holotype (FH), isotype (K). ARGENTINA. Misiones: Arroyo Piray Guazi, 24 Feb 1960, R. T. Guerrero 73 (LIL).

It is the medium-sized carpophore of the holotype collection which yielded the

spores which are identical with those found in the Argentine material. A Colombian collection which differs from the other collections in more chestnut and rusty orange pileus, yellowish, adnexed lamellae, and slightly larger size of the carpophores, was referred to AMarasmius berteroi var major in spite of the fact that the spores were some- whlat larger than usual so that it may have keyed out here (with Al. rhabarberinlus).

ILLUSTRATIONS. Singer (1965) pi 13, fig 71; (1958a) fig 23.

126. Marasmius berteroi (Leveille) Murrill, North American Flora 9: 267. 1915. HeliomnYces berteroi Leveille, Ann. Sci. Nat. III. 2: 177. 1844.

126a. Marasmius berteroi var berteroi TYPE. Bertero, from Puerto Rico. Pileus broadly shallowly sulcate-furrowed, campanulate-convex, 8-15 mm broad,

otherwise like the type variety. Lamellae white to cream, not intervenose, subdistant to (mostly) distant, edge concolorous with sides, eleven to sixteen, ascendant or sub- horizontal (oblique), moderately broad to broad, subventricose to 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 . Hymenium like in var major, cystidia occasionally

observed but inconspicuous, mostly not reaching level of sterigmata and basidiomorphous or like basidioles and not differentiated in cresyl blue or phoxine mounts but some elements of the hymenium apparently remaining sterile and showing slightly thicker walls and slight incrustations (cystidioles?); cheilozystidia all hyaline, otherwise like

epicuticular broom cells. Hyphae clearly metachromatic in cresyl blue mounts in the sense that most hyphae are are vinous red as contrasting with the bluish hymenial elements, hyaline, 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 5-9 y, clavate, mostly hyaline to subhyaline but in some broom cells concolorous with setulae and strongly pigmented and then slightly thick- walled, and strongly contrasting in cresyl blue mounts where they are violet (this




type of elements evenly distributed among the other paler broom cells and giving the scalp preparations a mottled appearance), setulae hyaline to golden melleous, 3.5-3.8 X 0.7-1.5 (at base) ,, subacute to acute.

On forest litter adhering to mixtures of sticks and fallen leaves of various tree species rotting in the woods and forests, singly or in closely joined pairs.

MATERIAL STUDIED. PUERTO RICO. "Agaricus fuillus Bertero," Bertero (Herbarium Persoon (L) Type. BOLIVIA. La Paz: Nor-Yungas, Charobamba at 1200-1300 m alt, 30 Jan 1956, Singer B 725 (LIL). ARGENTINA. Salta: Oran, 3.5 km west of Rib Blanco bridge, 17 Mar 1955, Singer T 2310 (LIL); Tucuman: Ciudad Universitaria. 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 Islands is close. The type (FH) differs mainly in the color of the pileus as described.

ILLUSTRATION. Singer (1965) p 14, fig 74.

126b.Marasmius berteroi var major Singer, Sydowia 18: 334. 1965 (ex errore p 249 "var macrus").

TYPE. Singer T 2886, from Argentina. Pileus bright orange fulvous, orange, orange red, rufous ferruginous or intensely

ferruginous (between "feuille morte" and "Eldorado" or between "feuille morte" and "Peruvian br.", "Agate," "Kobe", "henna", 4-A-i2, towards margin sometimes "sunburst"; "Ponce de Leon," ll-L-10, or "Saratoga" (M&P) on center and ribs, sometimes reaching "Maracaibo" or deeper and richer than "caldera"; color often reaching tones deeper and richer than those of Maerz & Paul (such as Villalobos SO, 12 , 8), dried similar to 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, campanulate or conic-truncate with flattened disc, later convex and often umbonate, eventually convex-applanate or with uplifted margin and tending to become obtuse, frequently even with depressed center, 10-56 mm broad. Lamellae cream, pale cream, cream whitish, yellow white, milk white, often becoming pale orange yellow when dried, usually more yellow if older, the edges usually concolorous, rarely few lamellae mostly not intervenose, narrowly discolorous, more rarely sparsely to densely intervenose, all entire or with few lamellulae (through-lamellae 9-19; lamellulae 0-18), ventricose or not, narrow to broad, 3-12 mm broad, sometimes not quite reaching the margin, narrowly adnexed to free, not collariate, spore print pure white. - Stipe reddish brown to chestnut below 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 tapering upwards later sometimes equal, narrowly hollow (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 morphologically or chemically differentiated from the basidioles; cheilocystidia like the epicuticular broom cells but some-



Marasmius 173

times rather scattered and rare, with obtuse, sometimes long, erect setulae, more rarely rather numerous but not making the edge heteromerous (intermixed with basidia and basidioles). Hyphae, all thin-walled in pileus, everywhere pseudoamyloid and with clamp connections, in old specimens many 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 ochraceous ferruginous at least in upper portion which also may be thick- walled; setulae apical and erect, often rather long, especially in type 2 some rather coarse, e g about 1.7 , thick below, but in type 1 usually much more slender, projecting 5-11 .; the two types regularly distributed and scalp appearance mottled.

On branches and trunks fallen to the ground (dicotyledonous trees) and on leafmold of dicotyledons, monocotyledons, conifers, in montane, subtropical, tropical-montane forest, solitary or gregarious but scarcely cespitose (Podocarpus, Gramineae spp, Alnus, Phoebe, etc).

MATERIAL STUDIED. BELIZE. El Cayo, Valentin, 28 Jul 1936, E. B. Mains 3660 (MICH). COLOMBIA. Valle: Buenaventura, Calima, 22 Apr 1968, Singer B 6295 (F), a large-spored form see annotation below. ECUADOR. Napo: Lago Agrio, 17 May 1973, Singer B 7518 (F); Tungurah- ua, Rio Topo, 30 Apr 1973, Singer B 7225 (F). BOLIVIA. La Paz: Nor-Yungas, Rio suapi, 1250- 1300 in alt, 9 Feb 1956, Singer B 1043 (LIL); Rio Yariza, 23 Feb 1956, Singer B 1394 (LIL); Char- obamba, 30 Jan 1956, Singer B 693 (LIL). PERU. Cusco, Paucartambo, Atalaya, Carbon, 700 m alt, Vargas 13519 (LIL). BRAZIL. Santa Catarina: 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); Ciudad Universitaria, 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. Singer T 3073 (LIL); Misiones: Iguazu, Ref. "Yag- uarete ," 29 Feb 1960, R. T. Guerrero 133 (LIL).

The collection from Colombia (B 6295) agrees macroscopically with var major but may key out with Marasmius rhabarberinus rather than M. berteroi because the spores were 12-17 X 3 p, thus larger than usual in M. berteroi var major. Since this collection differs in several aspects including the habitat on wood rather than dead leaves, from M. rhabarberinus, it has been considered a large-spored form of M. berteroi but it may re- quire a special subspecific taxon.

The collection from Ecuador (B 7518) agrees with var major except for the discolorous or partly discolorous (concolorous with pileus) edge of the lamellae and may also be a variety in its own right or subvariety of var major.


127. Marasmius convoluticeps Singer, sp nov Fig 61. TYPE: G. & L. T. Eiten & G. M. Felipe, from Brazil. Pileo dilute brunneo, siccando laete ferrugineo, superficie convoluta gaudente, us-

que ad 15 mm lato; lamellis ad aciem haud discoloribus, confertis vel subconfertis, liberis vel subliberis; stipite atrobrunneo, ad basin strigoso. Sporis 12.5-16 X 3-4 ,; cystidiis 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 convoluted gyrose venose-rugose surface almost all over, glabrous, broadly and obtusely conic-campanulate (much like M. haematocephalus in shape and size) sometimes umbonate, 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




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, inodorous.

Spores 12.5-16 X (2.5-)3-4 u, fusoid, on apex and base sometimes slightly recurved towards the same side, smooth, hyaline, thin-walled, with several small oil

droplets, inamyloid. Hymenium: basidia 4-spored; basidioles fusoid; cystidia none; cheilocystidia like the epicuticular broom cells but subhyaline throughout. Hyphae of pileus trama hyaline, of stipe brown, parallel, thick-walled. Cortical layers: epicutis 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), dermatocystidioid small hairs which are 9-27 X 3.8-7.2 j, hyaline or subhyaline, claviform, more rarely cylindrical, 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. matrisdei which differs, especially with regard to the type of

covering on the stipe.

128. Marasmius matrisdei Singer, Sydowia 18: 338. 1965. TYPE. Singer B 2523, from Bolivia. Pileus when quite fresh beautifully light orange (10-H-7) to ochraceous orange

(11-L-9) in the central portion with light ochraceous orange margin ( l-G-9), soon

fading to melleous brown ("mast color" to "Punjab") in the center, dried "Yucatan" on margin and "Sudan brown" in center, slightly hygrophanous and bleaching-fading parts becoming pale ochraceous when dry (not dried), sulcate on margin over one fifth to one third of the radius both 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 subumbonate to umbonate, campanulate, soon repand, becoming flat-convex, or flat conical, 7-17 mm broad. Lamellae pale cream, close, narrow, adnate. Spore print pure white. Stipe at first entirely pallid white, then ochraceous brown with white apex, entirely very finely pruinate, sometimes with a small socle-like base, usually tapering upwards, about 25 mm long, 0.7-1.5 mm broad; basal mycelium white. Context white or whitish, unchanging, very thin in pileus, inodorous.

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 narrow elements which are irregularly arranged and hyphous but multiseptate (septa close); hymenophoral trama and 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. Cortical layers: epicutis 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), tapering upwards but mostly obtuse, more rarely acute,



Marasmius 175

few elongated to 18 plength, total measurements of broom cells of pileus 24-31 X 7.5-9 p; surface of stipe with numerous broom 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 branchlets of both mono- and dicotyledonous plants in tropical 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. Marasmius neglectus 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 perpaucis et vix distinctis, cystidioliformibus. Ad folia delapsa dicotyledonea in hylaea columbiana. Typus in F conservatus est.

Pileus deep-rusty-red-brown, deeper on center and striae, more rusty-orange-fulvous between striae, dried rather deep ferruginous-fulvous, occasionally slightly striped when dried, with eventually slightly lobulate margin, glabrous, sulcate, conic to campanulate, papillate, 6-9 mm broad. Lamellae light grayish-ochraceous with deep ferruginous (discolorous) edge, broad, subclose (about 16 lamellae and many lamellulae), not intervenose, rounded-subfree. Stipe chestnut below, light grayish ochraceous at the apex, glabrous but not shining, subequal, 16-18 X 0.3 mm; basal mycelium light grayish ochraceous becoming fulvous on drying, radially strigose. Context thin, whitish in the

pileus. 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 epicuticular broom cells but always with hyaline main body. Hyphae of the pileus trama hyaline, thin-walled, with clamp connections, strongly pseudoamyloid. Cortical layers: 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, straight.

On the mid-nerve of dicotyledonous leaves fallen to the ground in tropical rain forest.

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-group but lacks true cystidia. Since only few spores were seen in the type, it may be that the spores reach somewhat larger size in more abundantly sporulating material.

130. Marasmius helvoloides Singer, sp nov Fig 63. TYPE. Singer B 6298 a, from Colombia. Pileo cinnamomeo, debiliter ochraceo-radiato, magis brunneo siccando, sulcato,

convexo, centro umbilicato vel depresso, 2742 mm lato; lamellis albis vel albidis, subconfertis vel distantibus, haud intervenosis, anguste adnexis; stipite atrocastraneo apice pallidiore, ad basin strigoso. Sporis 12-15 X 2.5-3 ,; cystidiis nullus vel inconspicuis




rarisque et cystidioliformibus. Ad lignum putridum in hylaea columbiana. Typus in F conservatus est.

Pileus cinnamon, somewhat striped ochraceous in the depression of 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 narrowly depressed center, 27-42 mm broad. Lamellae white or whitish, edges not discolorous, subclose to distant, not intervenose, rather narrow, narrowly adnexed. Stipe dark chestnut or deep chocolate with paler apex (pale cinnamon e g), glabrous to indistinctly pruinate under 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 inconspicuous and rare, cystidiole-like; cheilocystidia with hyaline main body and pale ochraceous orange to hyaline setulae, otherwise like the epicuticular broom cells. Hyphae hyaline and pseudoamyloid in the trama of the pileus. Cortical layer: epicutis of the pileus hymeniform consisting of broom cells with a main body about 17 X 6-10 p, subhyaline to palest melleous, clavate, setulae 3-6.5 X 1-1.5 ,, acute, narrowly conical, subhyaline to deep ochraceous; Covering of 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 larger species with very variable number of lamellae and should not be confused with Marasmius berteroi var major which differs in colors and no 174, M. helvolus (compare description there).

131. Marasmius graminicola Spegazzini, 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. Lamellae white, with finely or intermittently orange edges, distant (8-9 through-lamellae in most adult specimens), rather broad, all equal, rather broad to broad, not intervenose, adnexed to subfree. Stipe umber to bronze, succineous-umber or rufescent-bister, glabrous and 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 radiating thin filaments, white. Context white in the pileus, thin, inodorous.

Spores (10-)12-13.8 X 3.3-3.7 ,, claviform-boatshaped or subfusoid, smooth, hyaline, inamyloid. Hymenium consisting exclusively of basidia and basidioles; cystidia none; cheilocystidia like the epicuticular broom 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 hymeniform, 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, orange brown, melleous, rarely hyaline, constantly present, subacute, rod-shaped to thin-spinulose.

On dead fallen leaves and stems of trees (e g Populus, Taxodium, Gramineae and other monocotyledonous plants, probably always basically on Gramineae but passing



Marasmius 177

over to other material nearby, fruiting in spring and summer). MATERIAL STUDIED. 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, Singer S 590 (BAFC): La Plata, Spegazzini (LPS), type.

132. Marasmius haediniformis Singer, Bull. Jard. Bot. Bruxelles 34: 363. 1964. TYPE. de Witte 9432, from Zaire. Pileus white, dried whitish (slightly radially striate with dirty pallid) with con-

colorous center, neither in age nor on drying becoming yellow, glabrous, sulcate, (moderately deeply so) to the center, the center often venose, campanulate-convex then con- ves and sometimes with slightly depressed center in age, 18-37 mm broad. Lamellae white with white edge, narrow, distant (10-17 through-lamellae, some lamellulae), often not reaching the extreme margin or reduced to veins there, not intervenose, subfree to adnate. Stipe chestnut to bister, with white apex, glabrous, hollow, equal or tapering 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 margin of 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 rather numerous with the shape of the basidioles but sometimes somewhat irregular, inconspicuous and thin-walled, not or scarcely projecting, 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 epicuticular broom 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 parallel with 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. MATERIAL STUDIED. ECUADOR. Napo, Shushufindi, 240 m alt, 15 May 1973, Singer

B 7446 (F). ZAIRE. Parc National Albert, around Hoysha, 1050 m alt, de Witte 9432 (BR), type. ILLUSTRATION. Singer (1964b) fig 24, B-E.

133. Marasmius proletarius Berkeley & 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 ochraceous cinnamon in the center and gilvous-white on the margin, glabrous, 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, rather narrow, to broad, adnate, more rarely adnexed. Stipe pallid-white, mostly avellaneous to 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 somewhat ascendant on the stipe; white rhizomorphs often present, tomentose. Context white, very thin, inodorous.




Spores 6-9.2(-11) X 2.5-4(-4.8)U, narrowly fusoid 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 epicuticular broom 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. Cortical layers: epicutis 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, hyaline, 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, reminding one of the elements found in Marasmiellus section 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.

MATERIAL STUDIED. CUBA. Wright (K, FH), type. MEXICO. Tabasco: Villahermosa, Parque de la Venta, 1 Aug 1969, Singer M 8783 (F); MA 8785 (F).

134. Marasmius microhaedinus Singer, Sydowia 18: 338. 1965. TYPE. Singer B 25 78, from Bolivia. Pileus hyaline, hygrophanous, dry-fresh white, dried pale ochraceous buff, glabrous,

smooth, obtuse, convex, then becoming concave, 6-11 mm broad. Lamellae white, close, narrow, with some lamellulae intermixed, not intervenose or rarely (when stipe eccentrically attached) somewhat forked-anastomosing, subfree. Stipe chestnut with white apex. macroscopically glabrous to very finely subpruinate (under a lens), equal to very slightly tapering upwards, 26-28 X 0.7-1.3 mm; basal mycelium consisting of a strigosity which radiates from the base over the substratum, the appressed fibers becoming light fulvous when dried, the others remaining white. Context thin, white, inodorous.

Spores 7-9 X 2.8-4 , ellipsoid to ellipsoid-oblong or fusoid, hyaline, smooth, inamyloid. 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 hymenophoral trama hyaline, with clamp connections, stronigly pseudoamyloid. 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 shrubs or herbaceous plants mixed with leaves (of a dicotyledonous tree) fallen to the ground in rain forest and on dead, rotting branches of Legum- inosae.

MATERIAL STUDIED. COLOMBIA. Valle: Rfo Pance near Cali, I May 1968, Singer B 6512 (F). BOLIVIA. Vaca Diez, Riberalta. 8 Apr 1956, Singer B 2578 (LIL), type.

ILLUSTRATION. Singer (1965) pl 15, fig 81

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



Marasmius 179

135a. Marasmius trinitatis var trinitatis TYPE. Dennzis 158, 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 intermediate zone (e g "rust, sorolla br.") or with deep brown-red margin, when very young often entirely "caldera" (brown red), but later becoming golden cinnamon to ochraceous brown (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 depressed in the center, campanulate-convex, then convex-subapplanate, 9-34 mm broad. Lamellae young often cinerous or 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 rarely moderately strongly intervenose in 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 applanate to subconcave on 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 epicuticular elements but entirely hyaline. Hyphae very broad and irregular in the trama of the pileus, some thickwalled, with clamp connections, 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 densely gregarious.

MATERIAL STUDIED. U.S.A. Florida: Highlands County, Highlands Hammock State Park, 30 Mar 1942 Singer F 449 (F). MEXICO. Veracruz, 7 km S of Montepio, Estacion Bio- logica de los Tuxtlas 21 Jul 1969, Singer M 8104 (F). TRINIDAD. Non-Pareil, Sangre Grande, 1 Oct 1949, Deinnis 158 (K), type. BRAZIL. Guapore, Guajara mirim, 10 Mar 1956, Singer B 1821 (LIL). BOLIVIA. Pando: Manuripi, Conquista, 25 Mar 1956, Singer B 2225 (LIL); 24 Mar 1956, B 2166 (LIL), (?).

ILLUSTRATION. Singer (1965) pl 15, fig 86 T.

135b.Marasmius trinitatis var immarginatus Singer, Sydowia 18: 344. 1965. TYPE. Singer B 1883, from Bolivia. Pileus, lamellae, and spore print as in var trinitatis. Stipe ochraceous brown to

chestnut brown with white apex, at first very finely pruinate but glabrescent, otherwise 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. Covering layers: 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-




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. Marasmius olivascenticeps Singer, sp nov Marasinius leoninuzs var olivascenticeps Singer, Sydowia 12: 105. 1958 "ad interim."

TYPE. Singer F 596, from Florida, U.S.A. Pileo roseolo-brunneo ad marginem, fuligineo-olivaceo in 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 conservatus est.

Pileus' fresh pinkish brown on the margin but fuliginous-olive in the center, dried fulvous-brownish with an olive tinge in the center, with sulcate margin, glabrous, smooth in the center, convex, about 20 mm broad or slightly less. Lamellae whitish with olive colored (discolorous) edge, moderately broad, moderately distant or subdistant. free to adnexed. Stipe chestnut, chestnut-black below 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 epicuticular broom cells, also occurring somewhat away from the edge proper, 20-33 X 5.7-6.5 , or spreading above. 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

regular hymenophoral trama all thin-walled. Cortical layers: epicutis of the pileus hymeniform, consisting of broom cells, appearing mottled when seen in scalp preparation because some cells are either melleous hyaline, or golden melleous to brownish melleous because of the pigment of the setulae: main body 25-28 X 5-7.8 u, setulae 4.3-7.8 X 1-1.8 /.

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 discolorous edge and the medium sized spores characterize this 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. elaeocephalus Singer. with olive pileus and distinct cystidia occurs.

137. Marasmius epelaeus 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-rugose either, convex with a shallow umbilicus, 17- 25 mm broad. Lamellae white 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 pileus.

Spores 7.5-8.2 X 3.5 p, ellipsoid-oblong, hyaline, smooth. Hymenium: basidia 4- spored; cystidia none: cheilocystidia like epicuticular broom cells, but all hyaline. Hy- phae of pileus and hymenophoral trama strongly pseudoamyloid. Cortical layers: epicutis 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: Vaca Diez, Arroyo Florida, 22 km west of Rib-

eralta, 7 Apr 1956, Singer B 3047 (LIL), type. This is intermediate between M. trinitatis var trinitatis (from which it differs in

the smaller spores and subdistant lamellae) and M. digilioi (which differs in its smaller carpophores and closer lamellae).


138. Marasmius digilioi 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), glabrous, smooth to distantly furrowed-subsmooth, eventually often becoming long-sulcate, otherwise smooth, or more rarely rugulose in the center, often undulate, with a slight depression or/and a papilla in the center, otherwise convex, but eventually applanate to irregular, 5-16 mm broad. Lamellae (olive-brown) whitish or grayish pallid when young, later between pure white and "rubber," with the edges concolorous with sides or more rarely with pileus, subclose to medium close, narrow to broad, ventricose or not, rounded-free to rounded adnexed, Spore print pure white. Stipe rusty ochraceous brown to deep chestnut below and gradually paler 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 suprahilar depression, 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 epicuticular broom cells or dif-

fering in being all hyaline, or else these broom cells intermixed with ampullaceous to ventricose elements which have sometimes a slight knoblike swelling at the tip, at times branched but not setulae-bearing, 20-27.5 X 5.8-6.5 p, i e cystidiole-like and in this form also occasionally occurring near the edge. Hyphae with clamp connections, pseudoamyloid. 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, sometimes 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 subtropical montane, also subxerophytic premontane and tropical montane forest.

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-




bradita del Mastil, 15 Jan 1950, Singer T 881 (LIL); Tapia, 15 Feb 1951, Singer T 1488 (LIL); Taficillo, 23 Feb 1955, Singer T 2218 (LIL).

139. Marasmius ater 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 lamellarum nullis; cheilocystidiis et elementis epicuticularibus frequenter pigmento intra-

parietali fuligineo instructis, setulis atrosepiaceis vel fuligineis apicaliter appendiculatis. Ad lignumiin Venezuela. Typus in NY conservatus est.

Pileus when dried black, indistinctly sulcate, sometimes somewhat rugulose, 3-4.5 mm broad when dried. Lamellae gray 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 cheilocystidia: cheilocystidia numerous but often intermittent, like the epicuticular broom 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 intraparietal pigment, 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-shaped to conical.

On decorticated, dicotyledonous wood. MATERIAL STUDIED. VENEZUELA. Dto Federal, El Avila, 24 Jul 1972. Dumont et

al VE 5980 (NY), type.

140. Marasmius ruber 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," "mandarine red," "burnt orange," "cadmium orange," "sunkiss," "deep chrome yellow," "Ponce de Leon," "Indian orange," "orange peel," " Cadmium yellow," 2-A- 12 M&P the general sequence being from golden yellow or orange to red as the specimen 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 depressed or umbilicate center, otherwise campanulate, then regularly to irregularly convex and eventually often applanate, 11-50 mm broad. Lamellae "pirate" or "apricot yellow" or "Egyptian red" near edges, sides concolorous with edges or "cadmium orange" or 10-L-9, generally at first more orange and on drying or maturing more red, narrow, tridymous or even tetradymous, not intervenose or with interspaces slightly rugose to slightly intervenose, appearing close where there are many lamellulae and distant or subdistant behind (about 20 through-lamellae in a medium sized specimen of 17 mm diameter), varying from attenuate-free to 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", 1 1-E-7, glabrous to subglabrous, not pilose except at base, equal or slightly tapering upwards, 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, inamyloid. Hymenium: basidia 17.5-19.5 X 4.8-7.5 A, 4-spored; cystidia none; cheilocystidia hyaline, in shape and structure like the epicuticular broom cells. Hyphae of pileus trama and hymenophore hyaline but becoming pale orange all over from a diffusing pigment, thin-walled, sometimes many thick-walled ones intermixed, filamentous but at times very irregular in shape at places, with clamp connections, pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, the whole epicuticular layer developing a cadmium yellow pigment soluble in NH4OH and KOH preparations and diffusing and disappearing eventually by pressure, main body of broom cells cadmium yellow 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 j1 long and 1-1.3 1 diameter at base, spinulose and acute or rod-shaped and obtuse, scalp preparation with 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), paratype.

ILLUSTRATION. Singer (1965) pi 15, fig 22.

141. Marasmius bellus Berkeley, Hook. Jour. Bot 8: 139. 1856. TYPE. Spruce, from Brazil. Pileus cream color, then yellow ("sunrise yellow", 9-G-4 10-F-6 M&P), perhaps

slightly hygrophanous at times, paler between striae and sometimes pallid in elevated parts, dried sometimes "honey yellow" (Ridgway) according to Dennis, in our specimens varying from 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, campanulate then convex, eventually applanate, with or without a small umbo, 13-35 mm broad. Lamellae white to subconcolorous with pileus (e g lamellar ground paler than the pileus, lamellar sides pale cream 9-C-2), subdistant to distant, at times rather strongly intervenose but 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 concolorous with the lamellae, later brown or chestnut, glabrous, smooth, 35-54 X 1-1.2 (-4) mm; basal mycelium fresh, varying from white to light fulvous (10-H-6) dried generally fulvous or partly fulvous (mixed with white fibers), often accompanied by white to fulvous white byssuslike mycelium covering the surface of the substratum. Context thin, white or whitish-pallid in pileus, with a weak odor or more often odorless.

Spores 8-12.7 X 3-4.8 1, oblong or fusoid, often with applanate inner side, smooth, hyaline, inamyloid. Hymenium: basidia clavate; basidioles fusoid, or clavate; cystidia




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 epicuticular broom cells in all regards except that they have at times smaller setulae than the latter. Hyphae and often lower portion of main body of epicuticular elements pseudoamyloid, those of pileus and stipe very strongly pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, 13-32 X 5.5-9 ,, ventricose to vesiculose, generally thin-walled, with apical and often slightly diverging setulae, the latter spinulose, rather thin, subacute, the entire broom cells appearing hyaline or subhyaline when seen individually but in recently collected relatively well pigmented material they are yellowish to chrome yellow when seen in accumulations.

On forest and plantation litter of all kind, such as grass, herbaceous stems, monocotyledonous and dicotyledonous leaves, sometimes attached to small pieces of wood, in coastal montane tropical forest, Amazonas rain forest, etc.

MATERIAL STUDIED. BRAZIL. Amazonas: "Panure" (=Ipanure= Sao Jeronimo), Spruce (K) type; Pernambuco: Recife, Dois Irmaos, Severino Jose da Silva, 10 Apr 1955, comm. Chaves Batista 2224 (LIL). BOLIVIA. Beni: Vaca Diez, virgin forest 3 km below Guayaramerin, 12 Mar 1956, Singer B 1898 (LIL); La Paz: Nor-Yungas, Coroico, 25 Jan 1956, Singer B 669 (LIL).

ILLUSTRATION. Singer (1965), pl 15, fig 84; (1958a) fig 21; see also Dennis (1961) fig 43.

142. Marasmius beniensis 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 smaller when dried. Lamellae white, edge indistinctly concolorous with the pileus, subdistant to subclose (not close or crowded) intermixed with lamellulae which are often crisp or forked, otherwise not intervenose, adnate. Stipe "Al- amo" at base and gradually less deeply rust-ochraceous brown 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; cheilocystidia like epicuticular broom cells. Hyphae of pileus very strongly interwoven and variable in shape and thickness of the wall, pseudoamyloid. Cortical layers: 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 preparation not showing hyaline and colored zones (not mottled!), setulae concolorous with main body, 5.5-7 , long, thin and acute or spinulose and obtuse; covering of stipe almost absent, consisting of inconspicuous, minute, smooth, conic excrescencies 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. MATERIAL STUDIED. BOLIVIA. Beni: Guayaramerfn, 14 Mar 1956, Singer B 2000

(LIL), type. PANAMA. Gatum Lake, 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 larger carpophore of this collection.



Marasmius 185

143. Marasmius bezerrae Singer, sp nov Fig 65.

143a. Marasmius bezerrae var bezerrae TYPE. Singer B 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 arborum dicotyledonearum delapsa in silva tropicali Brasiliae. Typus in F conservatus est.

Pileus light brown to ferruginous (e g 12 J 9) and "rust, sorolla br" or ferruginous (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 marginal portion, convex, obtuse or broadly umbonate, sometimes more applanate in age, 7-17 mm broad. Lamellae pallid to cream, with a broadly and distinctly discolorous (orange to concolorous with some part of the pileus) edge, medium close to subdistant, narrow to moderately broad, not intervenose or in age slightly intervenose, often some forked, adnate, more rarely rounded-adnate. Stipe deep cinnamon brown or chestnut or deep chestnut brown, gradually more pallid at the apex at first, equal, perfectly glabrous, hollow, 15-50 X 0.6-1 mm; basal mycelium moderately abundant but 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-spored ones intermixed; basidioles fusoid; cystidia none; rarely a few cystidioid bodies little differentiated from the basidioles present at the gill edge; cheilocystidia like the epicuticular broom cells. Hyphae of pileus hyaline, all firm- or thin- to thick-walled, moderately strongly but distinctly pseudoamyloid, not dimorphic, radially arranged, interwoven, with clamp connections. Cortical layers: 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 0.4-2 2, obtuse, spinulose, deep brownish melleous, golden melleous or orange, in other cells subhyaline, the scalp view of the epicutis therefore appearing mottled. Surface of the stipe showing a few scattered inconspicuous versiform hairs but no broom cell dermatocystidia or setuloid excrescencies.

On dicotyledonous sticks fallen to the ground in tropical forest. MATERIAL STUDIED. BRAZIL. Pernambuco, Caramagibe, 14 Jul 1960, Singer & Bezerra,

Singer B 3409 (F), type.

143b.Marasmius bezerrae var chitepecensis Singer, var nov Fig 66. TYPE. Guzmzbnz 2800 B, from Mexico. Varietas foliicola lamellis confertis et pileo sublevi insignis. Differs from the type variety only in close lamellae and not or slightly sulcate

margin and growth on dicotyledonous leaves. MATERIAL STUDIED. MEXICO. Chitepec, 30 Oct 1960, Guzman 2800 B (F) type. The relatively broad spores and the strongly discolorous edge of the lamellae

are characteristic.

143c. Marasmius bezerrae var griseoferrugineus (Singer) Singer, comb nov lMarasmius leoninus var griseoferrugineus Singer, Sydowia 18: 337. 1965. TYPE. Singer B 1919, from Bolivia. Differs from the preceding varieties in gray lamellae which are subclose but appear

subdistant when dried, and grayish pallid apex of stipe.




On dead dicotyledonous leaves. MATERIAL STUDIED. BOLIVIA. Beni, Vaca Diez, 3 km below Guayaramerini in virgin

tropical rain forest, 12 Mar 1956, Singer B 1919 (LIL), type. A complete description of this variety has been given by Singer (1965, p 277 as

M. leoninus var griseoferrugineus).

144. Marasmius napoensis Singer, sp nov Fig 67. TYPE. Singer B 7348, from Ecuador. Pileo brunneo, longe profundeque sulcato, 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 conservatus est.

Pileus "cocoa" with "Seminole" (M&P) margin or "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 depressed center when mature, up to 60 mm broad. Lamellae whitish, then (in situ) discolored sordid 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-subfree to adnexed. Stipe deep chestnut below, at first with whitish apex, macroscopically glabrous but under a lens finely pruinate at 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) characteristically ellipsoid, some oblong, ovoid or broadly fusoid, mostly with a slight suprahilar depression, 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 trama hyaline, 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 ,, versiform, ochraceous to brown, fewer hyaline or subhyaline, with thin or thick wall: setulae 4.7-11 X 1-1.5 u, acute, all erect or somewhat spreading but all apical, deep succineous brown or ochraceous brown; 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. Marasmius leoninus 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. Marasmius leoninus 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) margin often 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 established by subhygrophanous zones at times present in the fresh pileus, glabrous sulcate to deeply furrowed-sulcate

(not merely short transparently striate), leaving a small to large disc either smooth or more rarely weakly rugulose-verruculose, convex, then convex with rather narrowly depressed center, often with recurved-applanate to even uplifting margin at maturity, or, on the contrary, with very declivous margin and an applanate zone around an umbilicus or an umbilicate small umbo, eventually mostly entirely subapplanate to concave, 15-42 mm broad. Lamellae pure white, cream white, or light cream, with the edges concolorous with sides, or at least not conspicuously discolorous, narrow to 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, adnexed to adnate to a widened apex of stipe (very variable in gill-attachment), intermixed with lamellulae and sometimes regularly tridymous with entire edge average 12-16 through lamellae. Spore print pure white. Stipe fulvous to very deep chestnut or ochraceous brown, apex at first white pallid or concolorous with the lamellae, later concolorous with base or merely somewhat lighter colored (Aesculus-seed-color, or brownish ochraceous), glabrous or subglabrous, smooth, shining to opaque equal or

tapering upwards, 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 rhizomorphs often observed. Context mostly relatively fleshy in center, thin in margin, tough in stipe, unchanging, taste mild; odor like that of Collybia dryophila or practically absent.

Spores 7-12.3 X 2-4.7 p, the most common length measurements ranging from 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 differentiated from basidia and basidioles inasmuch as the latter vary from transparent to subopaque-refringent; cheilocystidia present or absent, if present like the epicuticular broom cells. Hyphae either all equal and thin- walled, filamentous, strongly pseudoamyloid, with clamp connections, or similar but of two kinds, some thick-walled and even resembling at times a Crinipellis-hair when seen in the Melzer's reagent, very variable in size and shape, the others less strongly pseudoamlyoid, thin-walled and more regularly filamentous, not to very slightly gelatinized where the trama (of the pileus) is dimorphic, hyphae of stipe thick-walled, all strictly parallel. Cortical layers: 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-pedicellate or 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-pigmented than pigmented broom cells present, but more frequently most cells with melleous to succineous-melleous setulae, and a minority with hyaline ones, more rarely some chestnut-brown ones also present; smooth cells generally very few or none; covering layer of stipe either entirely absent, or else consisting of scattered broom cells, often with reduced main bodies and elongated setulae.




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 variable in several respects (1) in the pruinosity of the stipe, (2) structure of the trama of the pileus, (3) the presence or absence of cheilocystidia.

Of the specimens enumerated for this species above, most had the stipe annotated as glabrous and, indeed, under the binocular it appeared still glabrous, and under the microscope no broom cells were discovered away 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 subpruinose and scattered broom cells can be found. The colors of the three collections are similar but not identical, the lamellae vary in density from specimen to specimen of the same collection. Since we cannot find any correlated character we do not think this form to be worth of a varietal name at the present moment.

In Singer (1958a), I attempted to separate, describing them separately, a form with dimorphic and a form with homogeneous pileus trama. The form with the dimorphic trama was our Florida collection F 693. I thought at the time that this character was perhaps correlated with white basal mycelium and presence of cheilocystidia but I have found all transitions from 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 transitional cases between dimorphic and homogeneous trama (as in B 1 718) which I interpret as an indication of incomplete maturity, but the homogeneous forms are often provided with numerous cheilocystidia (B 1831 and F 570). It seems that the dimorphic-cheilocystidiate form is also represented by 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 dimorphic form 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. leotninus cheilocystidia "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 dimorphic.

This means that among the collections which show cheilocystidia, not all have dimorphic trama 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 demonstrate since in some

specimens the 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 dimorphic trama and glabrous stipe. It is uncertain whether this correlation holds any better than others, but I have studied material where the cheilocystidia are scanty which may be interpreted as a transitional form (B 1722); here the stipe is somewhat pruinose. The typical cheilocystidiate form was described by Singer (1958a, p 107, 108).

Consequently, I am not certain whether the degree of pruinosity and the presence or absence of cheilocystidia are useful characters for the infraspecific taxonomy of the Marasmius-leoniiius-complex but as far as the hyphal structure of the trama is concerned, I have thus far found a constant correlation between monomorphic (thin- to firm-walled hyphae only) tramal hyphae and the substratum on fallen leaves on one hand and dimorphic (many thick-walled and a variable number of thin-walled hyphae) hyphal structure with lignicolous habit. Thus I divided the species tentatively into two varieties:

Var leoninlus. Hyphal structure of 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-trama present. Lignicolous.

ILLUSTRATION. Singer (1965) pl 15, fig 90. The following variety is obviously very close to both the preceding but differs

in the fact that this correlation is absent (here both thin- and thick-walled hyphae in the pileus-trama and habitat on dicotyledonous fallen leaves as well as on wood); also the spores are shorter than usual in Marasmius leoninus and the edge of the lamellae is tending to become somewhat discolorous when dried:

145b. Marasmius leoninus var aberrans Singer, var nov Fig 68. TYPE. Singer M 8188, from Mexico. Hyphis et crassi- et tenuitunicatis in tramate pilei, sporis brevioribus, acie lam-

ellarum subdiscolori in siccis nec non habitatione ad folia dicotyledonea delapsa differt. Typus varietatis in F conservatus est.

Pileus dull orange, (ll-E-10 to 12 E 12) dried 11 I 9 (M&P) with paler, even

pallescent marginal zone, glabrous, coarsely deeply rugose-sulcate up to near the center, convex, often somewhat umbonate or slightly umbilicate when dried, 10-30 mm broad. Lamellae whitish, fresh with concolorous (whitish) edge, but on drying edge variably either concolorous or partly to almost entirely pale orangy, slightly ventricose, not intervenose, 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, inodorous.

Spores 7-9.5 X 3.5-4.5 ,, ellipsoid, fewer oblong, without suprahilar depressions, smooth, hyaline, inamyloid. Hymenium: basidia 19-26 X 6-7 u, 4-spored; basidioles fusoid; cystidia none; cheilocystidia like the epicuticular broom cells but relatively more of them, sometimes all, hyaline. Hyphae of the pileus-trama with thick (reaching 1 ,) to mostly thin to firm wall, not gelatinized, with clamp connections, pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform, consisting of broom cells, with hyaline wall but most filled with a dissolved intracellular pale orangy pigment, clavate,




cylindrical or ventricose, 9-19 X 4-12 g, setulae 2-16 X 0.7-2 u, conical, acute or subacute, palest golden to golden hyaline or quite hyaline, or with a more conspicuous in-

traparietal melleous 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 approaches M1. iiapoesis because of its broader spores but differs in having the colors of M. leoninus, glabrous stipe and non-interwoven lamellae.

146. Marasmius floriceps Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 298. 1869. Fig 69.

TYPE. Wright 31, from Cuba. Pileus bright orange-rusty, bright orange or bright red-brown, sometimes orange

with some inconspicuous brown radial stripes and/or orange brown or orange yellow marginal band (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 orange brown becoming "copper br" M&P), glabrous, sulcate excepting the disc, rarely short-sulcate when mature with, more rarely without rugose or venose center, campanulate to 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. Lamellae white to orange-cream, subclose to subdistant, the edges not discolorous or only near margin pale orange when dried, narrow to rather broad, at first somewhat ascendant, then horizontal or almost so, free to rounded-adnexed, intermixed with many lamellulae. Stipe brown to deep chestnut 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 abundant and strigose. white or fulvous to tawny when fresh and dried. Context of the pileus white, very thin, inodorous.

Spores (8-)8.5-13 X (2.7-)3-4.2(-4.8),, fusoid to fusoid-oblong but some sometimes relatively broad and often curved in lower portion if narrow, smooth, hyaline, inamyloid. Hymenium: basidia 16-24 X 6-8 ,, 4-spored, basidioles fusoid; cystidia none

except for a few inconspicuous subulate and hyaline ones at the gill edges among the

cheilocystidia which are broom cells like the epicuticular broom cells but usually all

hyaline or with golden hyaline or orange-hyaline setulae. Hyphae of the trama of the pileus hyaline with firm but not distinctly thickened wall although often slightly swelling in alkali but not dimorphic, filamentous, not gelatinized, with clamp connections, but often many secondary septa present, pseudoamyloid, those of the stipe parallel, + thickwalled, deep yellow. Subhymenium cellular. Hymenophoral trama regular, consisting of filamentous but sometimes relatively broad, thin-walled elements. Cortical layers: epicutis of the pileus hymeniform, consisting of broom cells, their main body 9-13 X 8-11 u, hyaline to ochraceous, scalp preparations appearing mottled 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 excrescencies looking 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

Singer M 8741 (F); 20 Jun 1969, Singer M 8088 (F). COLOMBIA. Valle, Municipalidad Cali, Rio Prance, 1 May 1968, Singer B 6511 (F); Buena Ventura, Calima, 18 Apr 1968, Singer B 6206 (F).

This species is often difficult to distinguish from M. berteroi var berteroi and I. corrugatus var auranltiacus but the characters emphasized in the key should make

it possible to come to an identification in all cases if the distribution of the three species is taken into consideration and the cortical layers of M. corrugatus are duly investigated.

ILLUSTRATION. Dennis (1951b), fig 7 (type).

147. Marasmius cuatrecasasii Singer, sp nov Fig 70. TYPE. Singer B 6517, from Colombia. Pileo dilute ochraceo vel cinnamomeo-brunneolo in centro, pallescente in mar-

gine hygrophano, longe sulcato, 20-31 mm lato; lamellis pallidis, satis angustis, subconfertis vel subdistantibus; stipite cinnamomeo vel castaneo, sub lente subtiliter pruinoso. Sporis 6.2-9 X 2.5-3.7 p; cystidiis nullis vel admodum inconspicuis paucisque. Ad lignum in Columbia. Typus in F conservatus est. Species de scientia botanica colom- biana meritissimo cl. J. Cuatrecasas dedicata.

Pileus light ochraceous brown or cinnamon brownish in the center, bleaching to 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. Lamellae pallid, rather narrow, subclose or in mature specimens mostly subdistant, narrowly adnexed to rounded-adnate or adnate. Stipe dull cinnamon to dark chestnut or dark cinnamon spotted at the apex, macroscopically glabrous appearing subvelutinous when 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 inconspicuous and 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 variable in the thickness of the wall as well as the shape of the hyphal cells but basically thin-walled and radially arranged in the pileus, subparallel in the stipe, strongly pseudoamyloid with clamp connections. Cortical layers: epicutis of pileus hymeniform, broom cells of the Siccus type, main body 10-15 X 6-10 $,, often hand-shaped or subcylindrical to clavate, hyaline, many with and many without setulae, these, where present, rarely hyaline, mostly melleous-ochraceous, obtuse, 4-11 X 1-1.7 1, the broom cells alternating singly 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. MATERIAL STUDIED. COLOMBIA. Valle: Buenaventura, Calima, "Pulpapel," 20 Apr

1968, Singer B 6258 (F); Rio Pance near Cali, 1 May 1968, Singer B 6517 (F), type.

148. Marasmius corrugatus (Patouillard) Saccardo & Sydow, Sylloge Fungorum 16: 54.1902.

Androsaceuis corrugatus Patouillard, Bull. Soc. Mycol. 16: 175. 1900.




148a. Marasmius corrugatus var corrugatus TYPE. Duss, from Guadeloupe. Pileus pinkish orange to reddish cinnamon, (e g 10-C-10 M&P to "Saratoga,"

dried "caramel" mixed with "Alamo" M&P, a narrow disc often with a tan or "burnt umber" shade in both fresh and dried state but this discoloration often missing and, instead, center sometimes more reddish orange than otherwise, smooth to finely striate on margin, otherwise more or less strongly venose or radially rugose, glabrous, convex, often with depressed center in age, 10-47 mm broad. Lamellae white to whitish, narrow, crowded or close, rarely medium close, not intervenose, adnexed, some almost subfree, others almost adnate, in the herbarium usually darkening to red-brown (between "cocoa" and "Arabian brown"). Stipe in primordium white, then fulvous to chestnut with orange ochraceous upper portion, smooth and glabrous, opaque to shining, terete or in larger specimens 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 rather thin, 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 epicuticular broom cells but all hyaline or stramineous-hyaline or 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 basidia and basidioles at least near the stipe. Hyphae of pileus and hymenophoral trama hyaline, thin-walled, sometimes some thick-walled but often very broad, in stipe parallel, all hyphae strongly pseudoamyloid and 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 irregular in 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 fulvous, spinulose, rod-shaped or even fusoid, sometimes forked, mostly simple, acute to obtuse, scalp preparations show mottled surface because of even distribution among the colored setulae and smooth cells of some broom cells with completely hyaline setulae; in age the percentage of the smooth epicuticular cells 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 emerging from the parallel surface

hyphae of the cortical layer; these excrescences inconspicuous, small and scattered, brownish, occasionally fasciculate and appearing to 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.

MATERIAL STUDIED. GUADELOUPE. Basse-Terre, Duss (FH) type. VENEZUELA. Suc- re, NW of 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 geographically and in habitat widely separated from the type may not be simply synonymous; I found neither hairs nor broom cells on the stipe. This collection is only temporarily inserted here.


148b.Marasmius corrugatus var aurantiacus (Murrill) Singer, comb nov Marasmiuts aurantiacus (Murrill) Singer, Sydowia 18: 268. 1965. Gyvmnopus aurantiacus Murrill, Bull. Torrey Club 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), subhygrophanous and bleaching in 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-chestnut or ochraceous brown, glabrous, smooth or slightly finely striate at maturity or in dried caps, but in many specimens more or less distinctly corrugated (radiate-venose-rugose) in the center, conic or campanulate-convex, then applanate and often with uplifted margin and 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 intervenose or very slightly so, about 22 or more through-lamellae. Stipe at first white with ochraceous-pallid or pale orange ochraceous base, later concolorous with the pileus with white apex, becoming ferruginous brown, often canaliculate, otherwise smooth, glabrous or subglabrous opaque and when seen under a lens the young surface is often covered by a thin, scattered pruinosity, glabrescent, equal or very slightly tapering upwards, 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: basidia 4-spored; basidioles fusoid; cystidia none: cheilocystidia like epicuticular broom 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 pseudoamyloid, in trama of stipe and basal mycelium frequently thick-walled, all hyphae with clamp connections. Cortical layers: 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 regularly distributed) 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) ,, characteristically variable in length and also varying from acute and spinulose to rodshaped and 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 generally a small minority). Covering of the stipe consisting of very scattered to rather numerous broom cells similar to those of the pileus but often narrower with long hyaline to golden melleous setulae, those often divergent spreading, sometimes arising almost directly from the surface hyphae 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. Underlying cortical hyphae strictly parallel. Setae, conical excrescencies, etc absent on stipe and even broom cells tending to disappear in old




specimens, but thin-walled, minute, hyaline excrescencies of variable shape often accompanying the broom cells.

On dead wood, generally on very rotten wood of dicotyledonous trees often covered by leafmold or on rotting chips of wood in leafmold, solitary to densely gregarious, almost cespitose; in warm-temperate, subtropical, tropical and tropical-montane forest in the northern hemisphere and in tropical forest in the southern hemisphere. Type on Quercus.

MATERIAL STUDIED. U.S.A. Florida: Alachua County, Planera Hammock, 16 Jul 1938, lMuirrill (FLAS), type; Highlands County, Highlands Hammock State Park, 5 Sep 1942, Singer F 596 (F). COLOMBIA. Valle, Rio Pance, 1600-1800 m alt, Singer B 6945 (F). BRAZIL. Pernambuco: Dois Irmaos, 5 Jun 1960, Singer B 3072 (BAFC). ECUADOR. Napo, Lago Agrio, 8 May 1973, Singer B 7303 (F). BOLIVIA. La Paz. Nor-Yungas, Coroico, 27 Jan 1956, Singer B 676 (LIL). ARGENTINA. Misiones, Camino de San Ignacio a Posadas, km 105, R. T. Guerrero 50, 23 Feb 1960 (LIL).

This variety is similar to lMarasmnius corrugatus var corrugatus which is usually somewhat larger, with slightly broader spores, not growing in cespitose groups and having no broom cells with well developed main body on the surface of the stipe.


148c. Marasmius corrugatus var lacustris Singer, var nov Fig 71. A var aurantiaco cui ceterum simillima haec varietas differt pileo maiore, derma-

tocystidiis stipitis dimorphis. Typus in F conservatus est. Pileus when quite young ochraceous brown or orange but soon, before maturity,

turning partly hyaline or pale orange, hygrophanous, white or whitish to leather-brownish when dry, glabrous, smooth or slightly sulcate to sulcate on margin, not scrobiculate rugose on the disc but sometimes venose almost all over, convex to conic convex, sometimes somewhat more repand with recurved margin, with or without a shallow depression or subumbonate, 25-57 mm broad. Lamellae white, sometimes with orangy ground, edges white, close to crowded, narrow or very narrow, with entire edge, sometimes interrupted with numerous lamellulae, eventually distinctly if not strongly intervenose, when young only rugose in the lamellae-ground, rounded-adnate to 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-oblong to fusoid, smooth, hyaline, inamyloid. 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 epicuticular smooth cells, 15-18 X 7-9.5 , (2) basidiole-like but sterile, e g 20-21 X 4-5 1. Hyphae of the pileus-trama both thin- and thick-walled, hyaline, pseudoamyloid, with clamp connections. Cortical layer: 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 stramineous or golden melleous, acute or obtuse, 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 numerous, or else very numerous from the beginning; covering of the stipe consisting of very scattered minute 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, subcapitate or 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, gregarious but not caespitose. Colombia to Ecuador.

MATERIAL STUDIED. ECUADOR. Napo, Lago Agrio, 7 May 1973, Singer B 7262 (F), Singer B 7422 (F), 11 May 1973, B 7366 (F).

148d.Marasmius corrugatus var portonovensis Singer, var nov Marasmnius portonovensis Singer, Sydowia 18: 272. 1965 nom subnud "ad interim."

TYPE. Rick 576, from Brazil. A var corrugato differt stipite pro ratione magis elongato, pileo minus intense

colorato, cellulis penicillatis epicuticularibus et cheilocystidialbus minus numerosis; hyphis tramatis hymenophoralis manifeste dimorphicis, sporis paullum minoribus. Ty- pus in FH conservatus est.

Pileus "Light Pinkish Cinnamon" and "Pinkish Cinnamon" (Ridgway), between "hazel" and "cigarette" M&P when dried, old fresh specimens flesh color (near "Peach Blow," 1 -B-5 M&P), central portion or entire pileus radiate-rugose, slightly paler between rugosities and margin faintly sulcate in dried material, convex, dried 7-32 mm broad. Lamellae white, 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: basidia clavate, 18-19 X 7 c,, 4-spored; basidioles fusoid; cystidia none; cheilocystidia dimorphic, most like the smooth, epicuticular elements, fewer like the epicuticular broom cells and entirely hyaline, some irregular, sometimes forked, often nodulose or even diverticulate where the subhymenial elements form the external layer of the edge, these hyaline, reminding one of the cheilocystidia of Maras- miellzs. Hyphae: in stipe all hyphae brown and parallel, filamentous, in pileus and hymenophoral trama irregular in shape, not all filamentous, characteristically dimorphic when mature, some thin-walled and moderately pseudoamyloid, others thick- walled like hairs of Crinipellis and very strongly pseudoamyloid, clamp connections present. Cortical layers: epicutis of pileus characteristically consisting of dimorphic elements, type I being clavate-vesiculose and hyaline without setulae and only with occasional asperities or 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 variable in length, (1-)3-8(-35) X 1-2(-3.5) ,, sometimes forked, generally rod-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, versiform hyphal ends, broom cells on stipe absent.

On small branches and large stems in forest litter, also on rotten logs. MATERIAL STUDIED. U.S.A. Florida: Highlands Hammock State Park, 27 Aug 1942,

Singer F 415 (F). BRAZIL. Santa Catarina: Porto Novo, Rick 576 (FH), type. ARGENTINA. Misiones: Bernardo Irigoyen, 27 Feb 1960, R. T. Guerrero 102 (LIL); Iguazu, Refugio "Yaguar- et6," 29 Feb 1960, R. T. Guerrero 114 (LIL).

This is thinner and more elongate, also apparently less intensely orange-chestnut than the type of M. corrugatus and differs from the latter likewise in the rela-




tively small number of broom cells among the setula-less cells in the epicutis and gill edge, in the strongly dimorphic character of the hyphae of the hymenophoral trama and in somewhat smaller spores. It differs from M1. aurantiacus which 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 epicuticular elements of the pileus, and in the color of the pileus.

ILLUSTRATION. Singer (1958a) fig 25. Stirps Cladophyllus

This stirps is intermediate between series Leonini and Actinopodes; it is characterized by the type of stipe-covering, crowded to close and often anastomosing lamellae and often glabrescent stipe; there are two subgroups, one with, one without distinct

pleurocystidia.

149. Marasmius araucariae Singer, Sydowia 18: 333. 1965. TYPE. Singer M 1147, from Argentina. Pileus castaneous-ferrugineous (so deep and rich that nothing comparable was

found in Maerz & Paul) or ferruginous or paler flesh color in age (3-A-11), dried about "rust, sorolla br." or somewhat deeper, glabrous to subvelutinous, slightly radially rugose 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 depressed center, later always convex with depressed center and eventually at timcs irregularly flattened, 15- 30 mm broad. Lamellae pallid-white to pallid-cream, subclose to crowded, in age generally subclose, intermixed with numerous, sometimes crisp lamellulae, occasionally some forked or with transverse ribs on lamellae-sides but not distinctly intervenose even in old specimens, narrow to moderately broad, free to rounded-adnexed. Stipe cinnamon below, cream white above, macroscopically appearing glabrous but under a lens white

pruinate, only in young specimens pruinosity visible without the help of a lens. smooth, equal or tapering upwards, 29-70 X 1.5-2.5 mm; basal mycelium fulvous-whitish. Con- text rather thin, 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 epicuticular elements 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. Cortical layers: 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. Hairs of the stipe surface, e g 50 X 5.5 p, mostly smaller, versiform, usually subcylindric with rounded tip, septate or not, thin-walled, hyaline, numerous.

On very rotten wood and needles of Araucaria brasilienisis in mixed forests of the subtropical-montane type, perhaps also on other substrata.

MATERIAL STUDIED. ARGENTINA. Misiones. Frontera, Gral. Manuel Belgranco, 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. Marasmius cladophyllus Berkeley, Hook. Jour. Bot 8: 138. 1856. Marasmiius polyporoides Murrill, North American Flora 9: 266. 1915. Favolaschia mnagnifica Hennings, Hedwigia 36: 203. 1897.



Marasmius 197

Laschia magnifica (Hennings) Saccardo & Sydow in Saccardo, Sylloge Fungorum 14: 199. 1899. Marasmius magnificus (Hennings) Singer, Lloydia 8: 218. 1945.

150a. Marasmius cladophyllus var cladophyllus TYPE. Spruce 89, from Brazil. Pileus orange fulvous with deep rusty fulvous center or orange ferruginous or a

similar shade (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 neighborhood often "Persimmon" M&P) rarely brown (between "tawny" and "Pekinese" M&P), sometimes "Kis kilim" with "Hispano" M&P margin (intermediate zone 12-D-ll), glabrous, smooth or slightly rugose or reticulate, generally not deeply radially sulcate in the marginal zone unless with cross veins at the same time, more rarely with rugose center and slightly sulcate margin, umbilicate or with more broadly depressed center in age, campanulate to convex, eventually flat, 15-60 mm broad. Lamellae cream, 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 carpophores the whole edge third or half subconcolorous with pileus but paler and the rest of the lamellae sordid (buffish-) pallid, mostly strictly horizontal-linear when mature, but varying from (mostly) narrow to (very rarely) altnost broad, strongly forking and merulioid-anastomosing or at least distinctly intervenose, often appearing poroid, intermixed but lamellulae anastomosing with through-lamellae and 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 reddish brown ("cocoa" to "trotteur tan"), often assuming a blue gray shine from the pubescence which covers it from apex to base, at least when seen under a lens but often glabrescent in age or pubescence becoming obsolete after handling, equal or tapering upwards, 30-112 X 1-4 mm; basal mycelium coarsely tomentose- strigose, mostly fulvous or sordid brown, etc ("Windor tan," "Titian," "alamo"), sometimes with some white fibrils immixed, rarely entirely white. Context white, unchanging, 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;cheilocystidia like epicuticular broom cells but some or more cells with hyaline setulae, often not occupying the entire edge and replaced by hyphal ends and versiform cheil-

ocystidia like those in var glaberripes and var intermedius. Hyphae of hymenophoral and pileus trama hyaline, smooth, in age some with thin, some with thick walls, pseudoamyloid, with clamp connections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, 15-25(-32) u long, main body either hyaline or concolorous with the setulae, setulae either all uniformly orange tan or on some cells hyaline to subhyaline, on others melleous to brownish melleous and sometimes on others even deeper colored, in the latter case scalp preparations giving a mottled appearance, setulae spinulose 3.3-12(-24) X 1.3-1.8(-3.6)ui. Pubescence of stipe consisting of hyaline, thin-walled hairs, these versiform or contorted, often cylindric or ventricose, with broadly rounded apex or forked at apex in a few cells, sometimes even with wavy-nodules outline or diverticula, reminding one of a Ram- eales-structure (Marasmiellus), septate or unicellular, 35-50 X 4-9.8 A (6.8-9.8 A in




middle of stipe of mature specimen), often accompanied by ventricose cells with or without mucro or over some areas replaced by these.

On monocotyledonous and dicotyledonous (but apparently always Angiospermae) wood and leaves, fruits and sticks, solitary or in small groups, not cespitose, in tropical rain forest and in subtropical to tropical forests along the Gulf and Atlantic Coast but not in tropical-montane (Yungas type) forests and not known from northwestern Argen- tina. Known hosts: Leguminosae, Palmae and Heliconia sp.

MATERIAL STUDIED. U.S.A. Florida: Highlands County, Highlands Hammock State Park, 3 Sep 1942, Singer F 521 (F); 26 Aug 1942, F 406 (F). BELIZE: El Cayo, Valentin, 24 Jun 1936, E. B. Mains 3628 (MICH), probably topotype of M. polyporoides; October 1906, Morton E. Peck (NY) type of M. polyporoides. PANAMA. Canal Zone: Barro Colorado Island, 13 Jul 1952, Mar- tin & Welden 7673 (LA); Corundu, 13 Aug 1952, Martin 8360 (IA); Rio Sardinilla, 10 Jul 1952, Martin, Welden & Bouchl 7552 (IA). ECUADOR. Napo, Lago Agrio, 16 May 1973, Singer B 7490 (F). BRAZIL. Para, Belem, 10 Jun 1966, Singer B 4248 (BAFC); Amazonas: Rio Jurua, Bom Fim, Nov 1900, Ule, Herbarium Brasiliense 2699 (FH); Panure (Ipanure (Ipanure = Slo Jeronimo), Sprnce 89 (K) type; Rio Javary, Remate dos Males, 1927, P. Occhioni 15, RB 5, 2, 17 (LIL); Pernambuco: Camaragibe, 6 Jul 1960, Singer B 3109 (LIL); Rio de Janeiro: Angra dos Reis. 30 Sep 1952, Singer B 436 (F); Rio Grande do Sul: Serro Azul 1928, Rick 422 (FH). BOLIVIA. Beni, Vaca Diez, Guay- aramerin, 6 Mar 1956, Singer B 15979 (LIL), 8 Mar 1956, B 1727 (LIL), B 1759 (LIL), 11 Mar 1956, B 1829 9LIL), 14 Mar 1956, B 1978 (LIL), 16 Mar 1956, B 2016 (LIL), 3 km below Guayaramerin in virgin forest, 12 Mar 1956, Singer B 1884 (LIL). ARGENTINA. Misiones: Arroyo Piray Guazli, 21 Feb 1960, R. T. Guerrero 74 (LIL); Bernardo Irigoyen 27 Feb 1960, R. T. Guerrero 68 (LIL).

ILLUSTRATION. Singer (1965) p1 22, fig 117.

150b.Marasmius cladophyllus var glaberripes Singer, var nov TYPE. Singer B 7160, from Ecuador. Stipite glaberrimo recedit. Typus in F conservatus est. This differs from the type variety in glabrous stipe, absence of broom cells on the

edge of the lamellae, slightly different color and larger size. 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. Lamellae radiating but so densely anastomosing that they appear pore-like, pallid with orange ground, later sometimes with partially ferruginous spots, narrow (1 mm), the pores almost isodiametric, adnate. Stipe chestnut, with pallid apex, glabrous (even under a lens), hollow, tapering upwards 67-76 X 1.5-2 mm; basal mycelium strigose, abundant, pallid. Context of pileus white, thin, membranous at margin, inodorous.

Spores (7.8-)8.5-9.8 X 2.5 ,, narrowly oblong-fusoid, smooth, hyaline, inamyloid. Hymenium: basidia 18 X 5.5 p, 4-spored; cystidia none; cheilocystidia consisting of pseudoamyloid 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 variegated as in the type variety, the main body 8-18 X 5-8.5 ,, mostly clavate, setulae acute, rather thin, 4-7 M long; 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. Marasmius cladophyllus var intermedius Singer, var nov TYPE. Singer B 7343, from Ecuador. Differt pileo minore stipiteque subglabro. Typus in F conservatus est. This is intermediate between var cladophyllus and var glaberripes. The pileus is

smaller than in var glaberripes, the stipe is subglabrous and externally beset by minute hypha-like excrescences which are occasionally like hyaline setulae (but singly on a strongly reduced or absent main body); only few broader hairs 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 mucronate, hyaline or browned, 12-27 X 6-18 p, inamyloid; the hyphae are dimorphic, some

strongly pseudoamyloid and 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 material cited for the type variety. It is possible that

some of these materials belong to either var glaberripes or var intermedius I have form-

erly inserted this species with the species of series Actinopodes but the existence of the varieties described here suggests that its place is rather in series Leonini.

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

151a. Marasmius pusio 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 younger, the deeper orange eventually fading to almost pallid in many collections, especially in the marginal zone, dried about light melleous (ll-G/H-6), glabrous, smooth, eventually becoming weakly shallowly and coarsely sulcate and on drying often short-sulcate on margin, not viscid, convex, not umbilicate but in some collections occasionally umbonate, sooner or later becoming applanate, 2-8 mm broad. Lamellae pure white to cream, all equal or more commonly rather regularly didymous or tridymous, rather narrow to very broad, arcuate to horizontal, never ascendant, close to subdistant (the smaller the carpophore the more distant appearing the lamellae and through-lamellae relatively not very numerous: 8-12), free to adnate to a fleshy disc, edges concolorous with sides. Stipe entirely white when quite young, later becoming "Windsor tan" 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, concolorous with lamellae or pileus, macroscopically seemingly glabrous when 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, usually central, more rarely somewhat eccentric, equal, curved or geniculate, 4-26 X 0.1- 0.5 mm, basal mycelium abundant although often indistinct when wet, strigose or tomentose and accompanied by thin white rhizomorphs in some collections, pure white or partly orange-pallid, dried white or partly fulvous-white. Context white, extremely thin, inodorous.




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-ellipsoid to oblong cylindrical, at times with a spur-like bulge on inner side near hilar appendage or with a bulge on the outer side like Canpanella alba, rarely curved, generally with a slight suprahilar depression, thin-walled, hyaline, smooth, inamyloid. 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 basidia and basidioles and only exceptionally tending to be slightly broader and with firmer wall than basidia, fusoid to clavate, e g 30-40 X 6.5-8 p;

cheilocystidia like the epicuticular broom cells, at places crowded, at places more scattered. Hyphae of trama of pileus and hymenophore thin-walled and thin-filamentous, hyaline, varying from moderately strongly but distinctly to very strongly pseudoamyloid, all with clamp connections, hyphae of stipe parallel with each other, always strongly pseudoamyloid. Cortical layers: epicutis of pileus subhymeniform or hymeniform, con-

sisting of broom cells of the Siccus-type, main body 6-35 X 5.5-11 1, hyaline or yellowish-hyaline, simple or branching Ranaria-like, often pedicellate, clavate, vesiculose cylindric-ventricose or mushroom-shaped, pedicel 4-4.5 u thick, setulae either likewise all hyaline 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 branching once 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 accompanied by diverticulate superficial hyphae, more numerous toward 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).


151b. Marasmius pusio var guatopoensis (Dennis) Singer, comb nov Marasmius guatopoensis Dennis, Kew Bull. 15: 95. 1961.

TYPE. Dennis 1103, from Venezuela. Differs from the type variety in broader spores (7.3-11 X 3.3-4.8 u); on fern rachis

and other Pteridophyta remnants but sometimes passing over onto dicotyledonous stems and sticks, usually growing in rows. The stipe is generally short (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. Marasmius pusio 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 rhizomorphs or 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.


151d.Marasmius pusio var graminivorus Singer, 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 generally numerous. 161. M. haedinus.

9. Lamellae medium close to distant; pileus 2-7 mm broad; cystidia few or moderately 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.




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. 166. tenutisetulosus.

3. Pileus 3-15 mm broad. 9. Pileus bright orange. On the leaves of monocotyledonous. more rarely dico-

tyledonous plants. 170. 31. bambulsinuls. 9. Pileus differently colored: bright ferruginous. ferruginous-brown. yellow-

brown, buff color to cinnamon, wood brown or leather brown, cocoa, etc. 10. Pileus ochraceous brown to buff brownish. On rotten grass, or seemingly

on the ground in meadows and grass lawns. occasionally on dung or various rotting herbaceous plants: in the subtropical and tropical-montane regions, also in the subalpine zone. 167. 1I. anomcluhs.

10. Pileus usually deeper colored and in different habitats. 11. Cystidia light fuscous; edge of lamellae discolorous (concolorous with

the pileus). 168. M. phaeocvstis. 11. Cystidia hyaline. sometimes some or many of them pale melleous to

golden melleous, never light fuscous; edge of lamellae concolorous with the sides. occasionally slightly discolorous near the margin of the pileus.

12. Cystidia moderately well differentiated to scarce; pileus larger than 9 mm; spores up to 15.2 p long. not longer (see "16" below).

12. Cystidia well differentiated and usually nutmerous; pileus 2-15 mm broad; spores often longer.

13. Pileus color between "cookie" and "tortoise shell" or 1 3 E 9 (M&P), dried near "Arab" or 13 B 11 in the center and near "Sayal br," "russet" or "terrapin" (M&P), "Hazel" (Ridgway); lamellae white or whitish, distant.

160. 1M. haemtatocepllalus var leucophyllus. 13. Pileus color different; lamellae white but tending to become

pale rusty or brownish or cream if quite distant, otherwise subdistant.

14. Lamellae distant (about 12-13, mostly all equal; if edges distinctly discolorous, see no. 163. AM. hypophaeus), South American species. 169. M. ferrugineuis.



Marasmius 203

14. Lamellae subclose to subdistant. rarely distant, mostly about 15-20 through-lamellae or. if fewer, with some lamellulae inserted among them. Mexico and Gulf region in general. 172. A1. w\ilsonii.

2. Spores smaller than 13.2 , 15. Pileus 13-44 mm broad, rarely smaller and then basal mycelium white when fresh

and mostly also when dried. 16. Cystidia mostly poorly differentiated and/or few on the sides of the lamellae;

color of the pileus between "Cocoa" and "leather brown" (M&P), dried not changing or becoming "auburn" with "pecan brown" striae or "Hazel" (Ridgway); lamellae subdistant; spores 11-15.2 X 2.8-4 p. Amazonas region.

174. M. heelvolus. 16. Cvstidia often well differentiated and numerous; color of the pileus different;

lamellae subdistant and then the disc of the pileus discolorous (bright fulvous or with an olive hue) or else lamellae definitely distant (mostly no more than 12 through-lamellae). Spores smaller than indicated above (up tol3.5, long, mostly shorter).

17. Lamellae white or with a slight cinnamon tinge, subdistant; if there is a discolorous disc area. it is bright fulvous or with an olive ringe around it; dull cinnamon or light pinkish cinnamon; species of the tropical and subtropical forests (Pacific Coast hylaea and Amazonas but north to Mexico at altitudes from 0-1000 m; pileus sometimes with white or pale ocher radial striae or with corrugated center; spores 8-11.7 X 2-3 .

180. LM pseudocorru,gatus. 17. Lamellae not pure white nor with a cinnamon tinge but between whitish

and a very pale cream or orange cream or between whitish and a very pale. dirty gray, decidedly distant; species of the Last slope of the Andes above 1000 m altitude or else spores relatively broader than indicated above ( >3.5 lpbroad); pileus, if discolorous on the disc, not bright fulvous there and without any olive tinge but either "Mohawk" to "Indian red" (M&P) or rusty-spadiceous, otherwise simply paler than the disc region or a bright and deep orange or ferruginous, "burnt sienna" or "Saratoga" or cinnamon to ocher brown, rarely somewhat paler striped and the disc area smooth or scarcely rugulose.

18. Pileus rusty spadiceous without any orange-ferruginous or orange zones or areas; lamellae whitish tending to gray: spores relatively broad (10-1 1 X 4.3-5 uor 7.5-9.7(-11) X 3.5-4.3pu); Amazonian species.

177. 31 hvlaeae. 18. Pileus or lamellae differently colored; spores relatively somewhat nar-

rower; species of the andine or prae-andine regions from Argentina to Colombia, occurring in tropical-montane or subtropical-montane forest above 1000 m altitude.

19. Pileus 13-44 mm broad; spores 9.5-13.2 X 3.5-4.5,; margin of pileus orange to ferruginous. 176. L. varizae.

19. Pileus 6-15 mm broad; spores 8.5-11 X 2.3-4ju; margin of pileus fresh ochraceous brown, dull ochraceous or pale cinnamon.

178. M. praeandinus. 15. Pileus 4-10 mm broad.

20. Basal mycelium white. South American species (see "19" above). 20. Basal mycelium tawny-fulvous. West Indian species growing on Cecropia and

other leaves. 179. M. hinnuileuis. 1. Lamellae crowded, close or subclose. Key III.

Key III

1. Spores (1 1.2-)14-18(-19.8) X 2.3-5.5p. 2. Cystidia poorly differentiated; pileus deep red brown, margin light cinnamon. On

dead wood in rain forest (see 123. M. rubricosus); on dead leaves (see 132. M. nleglectLus).

2. Cystidia very well differentiated; pileus often differently colored. 3. Pileus orange chrome to ochraceous orange; cystidia said to have rounded tips and

yellowish contents; margin of pileus somewhat weakly striate, center smooth; mostly on Bambuseae. 173. 1. dennisii.




3. Pileus often differently colored, with sulcate margin. 4. Pileus 40-48 mm broad; center of pileus radially rugose-venose; growing on

rotten branches in the Amazonas region. 171. M. nogalesii. 4. Pileus5-15(-40) mm broad; center of pileus not or scarcely rugose, often with a

small umbo or subumbonate. Growing on fallen dicotyledonous leaves (but also-on very rotten wood ib leafmold) in the Gulf area (see key 11 " 14").

1. Spores up to 14p in length. 5. Spores 10.5-13.7(-14) X 3-4p; species of tropical woods (not above 1000 m altitude)

with smooth center of the pileus, usually cespitose to fasciculate; pileus ochraceous to orange-rufescent, sometimes pallescent, 11-25 mm broad but sometimes smaller if growing solitarily; lamellae close or subclose. 175. 31. confertus.

5. Spores not more than 13.2 p long and mostly even smaller if pileus ochraceous to orange-rufescent; if growing in tropical forests, the center of the pileus is often corrugated; generally not cespitose or fasciculate. 6. Color of fresh pileus cinnamon, the disc sometimes discolorous; species of the

tropical lowlands in South America (see key II, "7"). 6. Color of fresh pileus not cinnamon.

7. Species of the montane zone of Mexico. Spores 8-13 X 3-4.3 u; lamellae in age with orange edge and scattered to numerous intervenose connections. 182. .1. aztecus.

7. Species of wide distribution; spores 7-11 X 2.3-4p; lamellae even in age not or scarcely intervenose and with the edge concolorous witll the sides or rarely (dried) intermittently orange-spotted. 181. M. spegazzinii.

Stirps Setulosifolius (no. 152-153)

152. Marasmius setulosifolius Singer ex Singer, Sydowia 18: 343. 1965. Marasmins setulosifolius Singer "ad int.," Sydowia 12: 110. 1958. TYPE. Singer B 1649, from Bolivia. Pileus orange-cinnamon, ochraceous cinnamon, rusty ochraceous brown, light brown

("gold pheasant," "harvest," between "tortoise" and "rust, sorolla," 1 l-F-9, 1 l-H-8, 1 1-C-9, 1 l-D-10, 13-B-l 1 M&P), dried "rust sorolla" and sometimes between 12-B-l 1 and "Eldorado" or 4-E-l 2 in center, i e more bright rusty in dried condition than in fresh, glabrous, distantly and sometimes intermittently deep sulcate to sulculate, generally normally long sulcate, center

rugose or smooth, subumbilicate to deeply umbilicate, at times slightly papillate, campanulate to convex, tending to become applanate and/or with uplifted margin 9-30 mm broad. Lame- llae white with the edge region concolorous with pileus or edges like sides (faces) buff-pallid, pale tan or entirely concolorous with the pileus, sometimes only the lamellar ground concolorous with pileus (or slightly paler), not intervenose or very slightly so in larger, mature specimens with a number of lamellulae, narrow to very narrow, rarely rather broad, subfree to

adnexed, rarely subdecurrent. Stipe buffy cinnamon to rusty ("gipsy") or ocher brown when fresh, with white apex, dried somewhat lighter colored, ochraceous, glabrous to slight-

ly pruinate, smooth, generally not shining or weakly so, equal or with widened base and/or with clavate apex 28-40 X 0.7-1.2 mm; basal mycelium tomentose or strigose, often radi-

ally spreading, brownish to cinnamon or white to whitish mixed with fulvous or ochraceous

fibrils, or entirely white. Context thin to very thin, white, unchanging, relatively toulh, inodorous.

Spores 8.2-10.2 X 3.5-4.7 g, ellipsoid to almost ellipsoid-oblong or subfusoid or ob-

long, thin-walled, smooth, hyaline, inamyloid. Hymenium: basidia 20 X 3.5-4 ,; cystidia of the broom cell type numerous all over the edges, sides and interlamellar spaces, 8.5-29 X 15-10 , main body thin-walled to thick-walled, cylindric to clavate, hyaline to melleous, setulae apical, erect to somewhat divergent and often considerably spreading (to 17 j wide), rarely forking, slightly to decidedly thick-walled, subacute to acute, melleous to deep melleous or brownish, 6.5-9.5(-24) X 1-2 (at base) u. Edges not even subheteromorphous. Hy-



Marasmius 205

phae of hymenophoral and pileus-trama hyaline, later with some thick-walled hyphae, rather dense, interwoven, pseudoamyloid, with clamp connections. Cortical layers: epicutis of

pileus hymeniform, consisting of broom cells of the Siccus-type, 15-25 X 9-10 , main body hyaline, thin-walled, cylindric to clavate, pale melleous to melleous, hyaline, setulae melleous to golden ocher brown, (3.5(-6.8-3.5 X 1-1.5 (at base) u, spinulose, acute or subacute; pruinosity of stipe (where present) consisting of broom cells but the main body mostly strongly reduced to virtually absent, setulae brown and up to 24 jlong.

On dead fallen leaves of both Monocotyledones and Dicotyledones (e g Cecropia sp) sometimes also passing over onto small woody sticks, in tropical rain forest.

MATERIAL STUDIED. PANAMA. Rio Sardinilla, 10 Jul 1952, Martin & Welden 7553 (IA). COLOMBIA. Valle, Buenaventura, Calima, 18 Apr 1968, Singer B 6200 (F). BOLIVIA. Beni: Vaca Diez, Gua!aramerin, 7 Mar 1956, Singer B 1649 (LIL) type; 8 Mar 1956, B 1733 (LIL) paratype, 14 Mar 1956, B 1952 (LIL) paratype, B 1966 (LIL) paratype, 3 km below Guayaramerin in virgin forest, 12 Mar 1956, Singer B 1866 (LIL) paratype, B 1885 (LIL) paratype, Ivon, 3April 1956, Singer B 2429 (LIL) paratype.

ILLUSTRATIONS. Singer (1965) pl 20, fig 110; (1958a), fig 27. A large-spored (spores 12.5-14.5 X 3.5-4.5 jL), smaller form has been observed in

Colombia,Valle, Buenaventura, 18 Apr 1968, Singer B 6200 (F). I consider it conspicific with 1l. setulosifolius at present, but if the correlation of spore and carpophore size should turn out to be consistent, this may well be an independent species. There are two palaeotropical species which obviously belong in the same group but have pliciform lamellae. The whole hymenium is here covered by broom cells and basidia. These species are: Marasmius silvestris Singer which is based on specimens considered to be a topotype of Vanrhomburghia silvestris Holtermann (18987) and Marasmius hymenofallax de Seynes (1897) known to me

only from description and illustration (p 18, pl 1, 12-17).

The other neotropical species of thiis stirps is the following:

153. Marasmius pleurocanthus Singer, Sydowia 18: 341. 1965. TYPE. Singer B 1920, from Bolivia. Pileus ferruginous with reddish ferruginous center (center "Monterrey" or "caldera,"

margin "copper brown" or "gypsy," in between "ferruginous"), glabrous, smooth, convex, 13 mm broad. Lamellae white to light grayish livid, with distinctly or indistinctly discolorous (concolorous with pileus) edges, narrow or rather narrow, of three lengths but not regularly tridymous, close to subclose, not intervenose, free to adnate. Stipe chestnut with white apex, glabrous at apex (under a good lens) very finely pruinate, subequal or slightly thicker at base and slightly widened at apex, 60-65 X 1-2 mm, basal mycelium strigose, white, when dried partially very pale fulvous. Context inodorous.

Spores 11-12.5 X 3.8-4.5 pu, oblong, smooth, hyaline, inamyloid. Hymenium: basidia 21.5 X 7 , 4-spored; cystidia of the broom cell type, main body 17-22 X 6.5-12.5 p, hyaline, setulae hyaline to chestnut, somewhat spreading e g 7 ,long, spinulose, obtuse, strictly apical, golden melleous, to melleous-castaneous: cheilocystidia little differentiated, much like the

pleurocystidia but even more like the epicuticular broom cells, more crowded than the pleurocystidia. Hyphae pseudoamyloid with clamp connections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus type, main body 13-17 X 4.5- 7 , subhyaline to pale melleous, often clavate, setulae as in pleurocystidia but sometimes

7 The figures by Holternann (1898 pi 11, 2a-d) are less slender and no spores have been recovered by Holtermann. Donk's collection determined as Valnrhloiblurgia silvestris by him (INDONESIA. Java, Tjibodas, 1600 m alt, Nov 1939 on fallen branches in rain forest, L 9023) was therefore considered the Type of 3Iarasmius silvestris Singer (1952, nom nud).




very long (up to 20 X2 2), the hyaline cells evenly distributed and the surface appearing mottled scalp preparations.

On dead fallen dicotyledonous leaves in rain forest. MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, 3 km below Guayaramerin in virgin forest,

12 Mar 1956, Singer B 1879 (LIL) paratype, B 1920 (LIL), type. ILLUSTRATION: Singer (1965) pl 21, fig 111.

Stirps Oleiger (no. 154-156)

154. Marasmius musicola Murrill, North American Flora 9: 260. 1915. Fig 72. TYPE. Earle 85, from Cuba. Pileus pale fawn colored, darker in the center, dried between "tawny" and "cookie"

M&P, glabrous, somewhat rugulose, with entire margin, convex to subexpanded, 10 mm broad. Lamellae whitish, broad, distant, slightly adnexed. Stipe reddish brown, glabrous. often twisted, shining, 40 X 0.5 mm; basal mycelium present. Context white, very thin.

Spores 16.8-20.5 X 3.2-4.5 p fusoid, smooth, hyaline, inamyloid. Hymenium con-

sisting of basidia, basidioles, cystidioles and cystidia with guttate contents, the guttalae consisting of an oily substance; basidia 4-spored; basidioles fusoid: cystidioles little different from basidioles but mostly clavate and appendiculate; cystidia 30-40 X 10-19 p, hyaline, club-shaped, filled with small oil droplets. Hyphae hyaline, pseudoamyloid. Cortical layers: epicutis of pileus appearing mottled because of accumulations of differetly pigmented broom cells which are arranged hymeniformly.

On banana trash, West Indies. MATERIAL STUDIED. CUBA. Santiago de las Vegas, 17 Jun 1904 F. S. Earle 85 (NY), type. This species has been misinterpreted by Singer (1965) before the type specimen was

carefully studied. Dennis (1951c) thought that M. musicola may be a small form of M. helvous (in his sense which is apparently the same as Marasmius poecilus, no. 118) while my treatment included the non-purple form of M. haematocephahls, both interpretations erronerous because of the characteristic cystidia found in the type specimen of M. musicolh

ILLUSTRATION. Dennis (1951c), fig 13.

155. Marasmius allocystis Singer, sp nov Marasmius allocystis Singer, Sydowia 18: 292. 1965. nom nud.

Type. Singer B 1875, from Bolivia. Pileo in siccis badio-castaneo, 7-12 mm lato; lamellis albis, ad aciem pileo concolo-

ribus; stipite albo, demum atrobrunneo, 18-52 X 0.4-0.8 mm. Sporis 13.7-17.8 X 3.5-4.5

cystidiis contentu oleaginoso conspicuo repletis; cheilocystidiis et cellulis epicutis apicaliter setulosis. Ad folia delapsa. Typus in LIL conservatus est.

Pileus tawny to fuscous ("Cowboy, new bronze, Brussels brown," "Mummy," "Alamo' dried bay to chestnut (between "Mohawk" and "Mandalay" or "cocoa"), glabrous, sulcate almost to the disc, campanulate or conic-applanate, 7-12 mm broad. Lamellae white with

edges concolorous with pileus, broad, distant, about 12-i 5, equal, free or adnexed. Stipe whitish, becoming brown from base upwards, eventually reaching blackish brown, glabrous, smooth, shining, equal, setose 18-52 X 0.4-0.8 mm; basal mycelium abundant, radially strigose or forming a tomentose socle, white but sometimes becoming partly fulvous when dried. Context thin, white, inodorous.

Spores 13.7-17.8 X 3.5-4.5 p, fusoid in frontal view but in side view often with curved hilar portion, hyaline, smooth, inamyloid. Hymenium: basidioles fusoid; cystidia numerous, conspicuous and versiform, cylindric-ventricose to fusoid, broadly rounded or narrowly muc-



Marasmius 207

ronate, 20-55 X 8-12.5 p with conspicuous oily contents (not so strong as in MA. oleiger), hyaline or melleous, opaque, refractive; cheilocystidia exactly like epicuticular broom cells. Hyphae strongly pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus type, main body hyaline, 8-18 X 5-8 p, setulae golden melleous to melleous brownish, some subhyaline, 2-7(-14) X 1.3 ,, needle-sharp-acute or obtuse, spinulose.

On fallen dicotyledonous leaves and moncotyledonous herbaceous stems, perhaps also on other debris.

MATERIAL STUDIED. VENEZUELA. Monagas, S. of Quebrada Seca, 16 Jul 1972, Dumont et al V'E 5173 (NY). BOLIVIA. Beni: Vaca Diez, Guayaramerin, 3 km below village in virgin forest, 12 Mar 1956. Singer B 1875 (LIL) type.

156. Marasmius oleiger Singer, Sydowia 18: 339. 1965. TYPE. Singer B 776, from Bolivia. Pileus "rust, sorolla brown" or slightly deeper colored, varying to "alamo" in the

center and fuscous brown with, perhaps, a slight olive shade (14-J-10) on margin, dried very deep ferruginous to reddish ferruginous, glabrous, sulcate over at least half the radius of the pileus, papillate, campanulate to convex, 5-11 mm broad. Lamellae white to cream avellaneous ("honeysuckle") with "rust, sorolla brown" edges, not intervenose, intermixed with a few lamellulae, more rarely equal, close to medium distant, (17-20 through-lamellae) moderately broad or medium broad, free. Stipe light brown to stramineous buff, glabrous, smooth, equal, setose, 26-50 X 03-0.7 mm; basal mycelium radiating and appressed to substratum, strigose, well developed, white or white to yellow when fresh, partly pale fulvous or fulvous when dried. Context thin, inodorous.

Spores 11.5-16.5 X 2.7-4.3 ,, fusoid when seen in frontal view, hyaline and smooth, inamyloid. Hymenium: basidioles ventricose, to fusoid, apiculate, hyaline without differentiated contents; cystidia 20-55 X 5.5-14 ,, of two types (1) like those of JI. gardleri, opaque and hyaline, (2) gloeocystidia filled with coarse, conspicuous, oily contents which turn deep bright blue in cresyl blue mounts, hyaline to mostly yellow or melleous, versiform: vesiculose, clavate, cylindric, ventricose, subcapitate or mucronate or appendiculate; cheilocystidia like epicuticular elements. Hyphae pseudoamyloid, hyaline to stramineous in trama of pileus, with clamp connections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus type, main body 8-21 X 5-11 , hyaline, clavate or ventricose-vesiculose or cylindrical, golden melleous (like setulae) or more frequently, paler than setulae or even hyaline, with thin wall but sometimes upper portion of main body with somewhat thickened wall; often pedicellate; setulae 3-7 X 1-1.5 (at base) i, golden melleous, subacute, but sometimes some quite acute or obtuse, subcylindric to narrowly conical.

On dead fallen leaves of dicotyledonous trees in tropical montane forest. MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Rio Llolose, 31 Jan 1956, Singer

B 776 (LIL) type, B 7833 (LIL), paratype. ILLUSTRATION. Singer (1965) pi 17, fig 99.

Stirps Megistosporus

157. Marasmius megistosporus Singer, Sydowia 18: 338. 1965. TYPE. Singer B 1971, from Bolivia. Pileus very deep brown on disc, around it a very deep ferruginous (8-J-12 on disc,

otherwise "Sierra, Venetian red"), margin "gold brown," disc smooth, otherwise sul-




cate, glabrous, broadly and shallowly conical with broad obtuse tip, almost convex, about 35 mm broad. Lamellae sordid white, rather broad (up to 4 mm broad), subdistant, not intervenose, rather narrowly adnexed. Stipe chestnut black, glabrous or

subglabrous, subequal, 100-130 X 1.5-2 mm, basal mycelium strigose, sparse to fairly well developed, white, turning partly fulvous on drying. Context thin, inodorous.

Spores 28-37.5 X 3.5-5.5 g, lanceolate-acicular to subclavate, in profile almost Fusarium (conidia)-shaped but only weakly curved, thin-walled, hyaline, smooth, in-

amyloid. Hymenium: basidia 33-39(-48) X 7.5-11 1, clavate to subcylindric, 4-spored; cystidia numerous, 45-85 X 9-15 y, cylindric to elongate-ventricose, with rounded or mucronate tips, or variously appendiculate even with oblique protracted apiculi or forked or with short irregular excrescences, thick-walled, opaque, without conspicuous contents, often distinctly deeper rooting than the basidia and some strongly projecting beyond the level of the sterigmata, others either not deep-rooting or not projecting; cheilocystidia much like the epicuticular broom cells. Hyphae of hymenophoral trama (which is regular) hyaline, strongly vinous pseudoamyloid, with numerous

clamp connections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, main body brown, e g 12-20 X 9-10 1, setulae either light brown or deep chestnut brown (mottled in scalp preparations), subacute, 6.5-11 X 1-3 u.

On rotten wood, in rain forest. MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, Guayaranerin, 14 Mar 1956, Singer

B 1971 (LIL), type. This species is among all Marasmii remarkable because of the enormously long

spores. These it shares with Marasmius megistus Singer from the Congo which has

easily the longest spores known in Basidiomycetes, but differs from Al. megistosporus in the absence of cystidia. Such enormous and at the same time perishable spores would be a handicap to any but a rain forest species where, at least during the larger

part of the year, a constant high degree of humidity is maintained on the substratum as well as in the lowest air-level.


Stirps Haematocephalus (no. 158-160)

158. Marasmius pallescens Murrill, North American Flora 9: 261. 1915. TYPE. Johnston 556, from Puerto Rico. Pileus fresh light lilac or pale red, dried reddish brown, long-sulcate, glabrous,

broadly campanulate becoming slightly umbilicate on drying, 5-8 mm broad. Lamellae white, distant (10-11 through-lamellae only), broad, distant, free to adnate. Stipe avellaneous, fuliginous below, eventually often entirely black, setose, glabrous, equal, 20-32 X 0.2-0.3 mm; basal mycelium abundant, white. Context white in the pileus, very thin in pileus and stipe.

Spores 11-17 X 3.2-4.5 j, tear-shaped to fusoid, hyaline, in the herbarium tending to palest melleous, smooth, inamyloid. Hymenium: basidia 16-20 X 6.5-8 A; cystidia very numerous, 32-43 X 10.1 5.2 j, clavate to ventricose, with obtuse apex or with a broad obtuse mucro, fewer with an appendiculus up to 14 /long, opalescent, hyaline; cheilocystidia like the epicuticular broom cells but all hyaline. Hyphae of the trama of the pileus hyaline, with clamp connections, pseudoamyloid. Cortical layers: epicutis of the pileus hymeniform consisting of broom cells with the main body e g 17 X 8-11.5 p, hyaline, with setulae 3-6.6 X 0.6 ,, pale melleous, interspersed with broom cells with deep melleous setulae (a minority) and epicutis therefore mottled in scalp view.



Marasmius 209

On petioles and leaves of dicotyledonous trees fallen to the ground. MATERIAL STUDIED. PUERTO RICO. Rio Piedras, 18 Aug 1912, J. R. Johnston 556 (NY)

type. VENEZUELA. D. F., Caracas Botanical Garden, 20 Jun 1958, Dennis 1172 (K).

ILLUSTRATION: Singer (1965) pl 18, fig 101; see also Dennis (1961) fig 48. At first sight one might be inclined to consider this species one of the numerous

color variants of lMarasmius haematocephalus but the revision of the type revealed that Dennis was correct in separating this species because of the somewhat smaller spores. However, "authentic" material J. R. Johnston 579 (NY) is probably another species.

159. Marasmius panerythrus Singer, Sydowia 18: 339. 1965. TYPE. Singer B 1609, from Bolivia. Pileus purple red (5-K-4) with paler margin ("copper leaf') becoming at drying

(not dried) "Etruscan red" and eventually "burnt sienna" to reddish chestnut, glabrous, deeply long-sulcate, convex to campanulate, sometimes with somewhat applanate to umbilicate center, up to 20 mm broad. Lamellae lilaceous pink or pink, later whitish with vinaceous or chestnut brown, eventually "burnt sienna" edge, narrow; eventually broad, ventricose, subclose to subdistant (about 17-20), free around a flat brown to ferruginous smooth disc or a brown enlarged apex of the stipe. Stipe at first "rose petal" with

apex colored "ashes of rose", (M&P), becoming black from the base upwards, later becoming dark brown to black all over, dried umber, glabrous, smooth, equal and up to 40 X 1-1.3 mm or tapering upwards to 0.7 mm; basal mycelium sordid white, rarely brownish white, in dried specimens very pale orange fulvous, covering the substratum in strings and tomentose patches (there ochraceous) in the type exuding droplets (but this is probably caused by a parasite). Context thin, white, inodorous.

Spores 13-14.7 X 3.5-4.2 p, fusoid in frontal view, inner side applanate or slightly curved, hyaline, very thin-walled and easily collapsing, smooth. Hymenium: basidia 20-25 X 6-6.5 ,n; cystidia 20-62 X 6.8-9.8 ,, + thick-walled and refractive, fusoid, ventricose, clavate or cylindric, often with irregular appendages, attenuate-obtuse to subacute, moderately numerous to numerous, well differentiated; cheilocystidia like epicuticular broom cells, main body hyaline to more often melleous, weakly pseudoamyloid, 11-18(-20) X 4.5-9.5 p; setulae erect. Hyphae moderately strongly, although distinctly pseudoamyloid, hyaline (in NH4OH and KOH) in trama of pileus and lamellae. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus- type, main body mostly melleous, only in relatively few broom cells hyaline, weakly pseudoamyloid, 11-17(-20) X 4.5-6.5 j, setulae all erect or somewhat spreading, 4-7 (-11.3) X 1.2 2, mostly melleous or golden yellow, rod-shaped or very narrowly spinulose and acute.

On bark and wood of dicotyledonous trees, perhaps also on rotten leaves, in rain forest.

MATERIAL STUDIED. VENEZUELA. Miranda: Los Guayabitos, 8 Jun 1958, Dennis 1011 (K, colored sketches seen). BOLIVIA. Beni: Vaca Diez, Guayaramerin, 6 Mar 1956, Singer B 1609, type; Pando: Manuripi, Sta. Rosa, 22 Mar 1956, Singer P 2147 (LIL), paratype.

This species is so characteristic in every regard, especially its colors, size and brown apical area of stipe, combined with spore size, that I have no doubts about the identity of the material sent to me by Dennis. The descriptive date at my disposal coincide so exactly with ours that they seem to have been taken from the same collection. Dennis does not mention cystidia but has probably not searched for them.

ILLUSTRATION. Singer (1965) pl 18, fig 103; see also Dennis (1961) fig 44.




160. Marasmius haematocephalus (Montagne) Fries, Epicrisis Systematis Mycologici, p 382. 1838.

Agaricus haematocephalus Montagne, Ann. Sce. Nat. Bot. II. 7: 369. 1837. Androsaceus haematocephalus (Montagne) Patouillard, Jour. Bot. 3: 334. 1889. Marasmius rhodocephalus Fries, Nova Acta Regiae Societatis Scientiarum Upsaliensis III 1:

31. 1851. Androsaceus rhodocephalus (Fries) Patouillard, Bull. Soc. Mycol. France 4: 20. 1888. Marasmits semipellucidus Berkeley & Broome, Jour. Linn. Soc. Bot. 14: 36. 1875. Marasmius sanguineus Cooke & Massee, Grevillea 17: 59. 1889. Marasmius atropurpureus Murrill, North American Flora 9: 262. 1915. Marasmius vinosus Beeli, Bull. Soc. Roy. Bot. Beige 60: 158. 1928, non Spegazzini.

160a. Marasmius haematocephalus var haematocephalus TYPE. A. de Saint Hilaire, from Rio de Janeiro, Brazil (type collection not found

but for topotype, see below). Pileus purple red to reddish purple: Hydrangea red (Ridgway) to deep Corinthian

red (Ridgway), "Peony Red" to "Dark Cardinal", 2-I-5 to 2-J-8, "Cuba," "Wallflower" (M&P), Garnet Red, Pompeian Red (Ridgway), margin often lighter or paler and duller colored, e g "marsh rose," "Athenia" or "ember" (M&P), Light Jasper Red (Ridgway), Dragon Blood's Red (Ridgway), center at times reaching as dark as "Ambulance" (M&P) or Haematite Red (Ridgway), eventually all parts slightly paler, glabrous, sulcate over more than half of the radius, campanulate, convex, often with a depression and a minute papilla in the depression or at least umbilicate or with a small umbo, eventually sometimes applanate in the center, 4-21 mm broad. Lameliae at first usually pink to

purplish pink, e g 2-B-3, "Arbustus" or "appleblossom," later pallescent from edges up, often lamellae-ground and faces near ground pink and edge half white, edges not concolorous with pileus when mature except at times very near the margin of the pileus narrow to broad, distant, 8-14 through-lamellae, usually no lamellulae but in some 1-2 lamellulae present, attenuate-free to narrowly adnexed. Stipe full and light umber with apex concolorous with lamellae or "strawberry" or "raspberry" colored, rarely white, later becoming white and eventually concolorous with lower portion of stipe, glabrous, smooth, shining, equal, setose, 16-60 X 0.1-0.8 mm; basal mycelium tomentose or

strigose, usually well developed, sordid pallid or white when fresh, and white to sordid when dried, sometimes a white tomentum and a pale fulvous brownish strigosity present. Context thin, white, inodorous.

Spores 14-21.5 X 3.5-6 ,, clavate-acicular to subfusoid, hyaline, smooth, inamyloid. Hymenium: basidia 29 X 7-7.5 p; basidioles about 20 X 4 p, fusoid; cystidia generally numerous and mostly well differentiated, rarely a minority less well differentiated; 25- 49 X 6-13 ,, clavate, ventricose, ventricose-constricted and appendiculate, more rarely vesiculose, ampullaceous or cylindric, sometimes fusiform, opaque, hyaline to pale melleous, with strongly or somewhat thickened wall, some at times scarcely opaque or refringent and with thin wall, broadly rounded or subacute above, blue in cresyl blue mounts; cheilocystidia like the epicuticular broom cells but generally all hyaline including the setulae, e g 26 X 5.2 u. Hyphae of the trama of the pileus hyaline, filamentous, 3-9 p thick, distinctly pseudoamyloid, with clamp connections. Cortical layers: epicutis of pileus hymeniform or subhymeniform, consisting of broom cells of the Siccus-type, 13-23 X 5-8 1 broad, but some spreading even broader: clavate, more rarely cylindric or vesiculose or ventricose, with erect apical setulae which are hyaline in some and melleous in other broom cells, there being a water soluble (also dissolving in NH4OH) pigment which is pink and dissolves into the medium upon slight pressure on the cover

glass, but this in some collections not visible; setulae, sometimes very fine, 3-6.5 p long,



Marasmius 211

obtuse to acute, main body thin- to thick-walled. On fallen dead leaves and sticks, wooden twigs and rotting pieces of wood, also

on dying outer portions of the bark of living and fallen trees, gregarious or solitary, not cespitose.

MATERIAL STUDIED. BRAZIL. Guanabara, Jardim Botanico, 28 Jan 1961, Singer C 3172 (BAFC), topotype. ARGENTINA. Tucuman, Tafi, Yerba Buena, 12 Feb 1955, Singer T 2157 (LIL); Capital, Garden of Instituto Lillo, 14 Jan 1952, Singer T 1731 (LIL).

In the relatively drier places of the plains and in the Chaco vegetation (subxerophytic woods) we observe a form which differs from the type form only in being very small (and thus resembling MA. tucunmanus), (T 2938 and T 2992); the pilei are 2.5-4.5 nun broad and at least as high. The base is often subinsititious. The epicutis does not contain a visible dissolving pink pigment.

A very similar form is my B 7328 from Ecuador, with the pileus pink (2 I 5 to 2 J 8, M&P) and small, 2.5-4.5 mm), poorly developed basal mycelium on Gramineae, the epicutis in thick layer pink in NH4OH but not visible and epicuticular broom cells as in var rubiciundus which may be identical if it were not for the different pigmentation and habitat.

In the Araucaria forest we observe a form with the pileus a purer purple red ("garnet r." M&P) and very pale colored setulae of the epicuticular broom cells (in mass dull pink, individually melleous subhyaline).

ILLUSTRATION. Singer (1965), pi 18, fig 103; (1958a) fig 33.

160b. Marasmius haematocephalus var purpureomarginatus Singer, Sydowia 18: 337. 1965.

Pileus "garnet red," sometimes paler striped, becoming darker on drying, 9 mm broad, 6 mm high. Lamellae white with pale purple ground, with "garnet red" edge. Cortical layers: epicutis with broom cells alternating with hyaline and melleous setulae, in mass deep reddish brown, no sap exuded. Otherwise like the type variety.

On small woody twigs or on logs of dicotyledonous trees and palms in rain forest.

MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, Guayaramerin, 14 May 1956, Singer B 1967 (LIL), type. Probably same: VENEZUELA. Bolivar: Puerto Zamuro, Jun 1887, Gaillard 52 (IH). ICUADOR. Napo, Shushufindi, 13 May 1973, Singer B 7411 (F), B 7395 (F).

160c. Marasmius haematocephalus var pseudotageticolor Singer, Sydowia 12: 120. 1959. Pileus dark Perilla purple, Indian Purple, Corinthian Purple (Ridgway), sometimes

duller (6-K-2, 7-J-4 M&P) or brighter ("red banana" in center and "rosevale" in marginal zone) or darker ("Liberia" or 56-L-12 in center, "cedar wood" M&P on margin) characteristically striped radially, like Marasmiius tageticolor and some related species of the acystidiate series, with sordid pallid to creamy, white zones mostly alternating with the sulcations, slightly larger than average type form: 8-25 mm broad. Lamellae: pink with white edge half, very distant to distant (6-11, all equal), at times slightly red-edged, free to adnate. Stipe in the darkest forms reaching black below, at times subinsititious. Spores (17.5-)20-23.5(-24.7) X 3.8-5.5 p(slightly larger on an average than in type form). Covering layers: epicutis with broom cells which are mostly hyaline in main body, often melleous to (fewer and not always) brown rather firm setulae, exuding or not a pink sap into the medium.

Otherwise like the type variety. On dead dicotyledonous leaves, sticks and stems.




MATERIAL STUDIED. Rio Sardinilla, 10 Jul 1952, G. W. Martin & A. M Bouche 7549 (F, IA), type. CUBA. Cienfuegos, White (FH). St. Vincent W.I., comm. Massee (as M. tageticolor) (NY). MEXICO. Chiapas, Finca Sospiro, 4 Aug 1969, Singer M 8966 (F). ARGENTINA. Tucuman: Taf', Parque Aconquija, 17 Feb 1955, Singer T 2183 (LIL); Trancas, Tapia, 17 Feb 1951, Singer T 1223 (LIL).

Among the material studied that from Tapia (T1223) is most remarkable for its dark color; it is also the smallest of all collections (pileus 8 mm diameter) and has no soluble pink pigment in the epicutis. The other extreme, the collection from the Amazonas Basin (B 1881) is more red than the rest ("red banana," a bright purplish red), larger than the others (pileus 23 mm diameter) and has likewise no soluble pink

pigment in the epicutis. The other collections have more or less pink pigment dissolv-

ing into the medium (but as always, disappearing in 1-2 minutes) and their size and color is in between the two afore mentioned extremes.

ILLUSTRATION. Singer (1965) pi 18, fig 100 ps: (1961) fig 47.

160d.Marasmius haematocephalus var oenechinus Singer, Sydowia 12: 120. 1959. Pileus vinaceous ("China rose" to "Amaranth purple" M&P), in the center often

dark as "Canyon." Lamellae concolorous, then whitish (bleaching from edge upwards). Stipe umber, light fuscous, eventually reaching blackish near base. Spores 18-22 X 3.5- 4 u. Cortical layers: epicutis of pileus with broom cells showing setulae which are entirely and persistently amethyst-purple to vinaceous; no pink sap diffusing into medium.

Otherwise like type variety. Particularly common on dead leaves of Cecropia. MATERIAL STUDIED. CUBA. Cienfuegos, [White (FH), type. BOLIVIA. La Paz: Nor-

Yungas, Charobamba, 1300-1400 m alt, 30 Jan 1956, Singer B 734 (LIL). ILLUSTRATION. Singer (1965) pl 18, fig 100, Oe.

160e. Marasmius haematocephalus var rubicundus Singer, Sydowia 18: 337. 1965. Pileus red to rose color (not purple) e g "Canna," "Carnelian," "Autumn glory,"

"Egyptian red," "Korea" M&P, 2-14 mm broad. Lamellae pinkish white, later white, interlamellar spaces at first concolorous with the pileus, about 11, free. Stipe at first often pink at apex, later umber all over, color of basal mycelium variable between white and brown.

Spores 15-21 X 4-4.8 ,. Cortical layers: epicutis with broom cells brownish melleous when seen in mass, because of the light brownish melleous color of the setulae; no exudation of pink, soluble pigment in water or NH4OH.

Other characters and habitat as in type variety. MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Charobamba, 1300 m alt, 30 Jan

1956, Singer B 743 (LIL), type, B 737 (LIL) paratype; Coroico, 1700 m alt, 7 Feb 1956, Singer B 945 (LIL) paratype; Beni: Vaca Diez, Guayaranerin, 4 Mar 1956, Singer B 1963 (LIL), paratype; B 1755 (LIL), paratype.

160f. Marasmius haematocephalus var macrocephalus Singer, Sydowia 18: 337. 1965. Pileus red ("hollywood" to 4-1-7 M&P), eventually pallescent to the color of the

proceding variety ("Carnelian"), deeply sulcate to the eventually somewhat rugulose disc, up to 42 mm broad. Lamellae at first with purplish pallid ground, later with white ground, otherwise white all over, distant, broad (4-5 mm), free or subfree. Stipe all white at first, soon becoming chestnut with white apex and brown "Mango") in a very small zone immediately below the lamellae, this being characteristic for the mature specimens, lower (chestnut) zone becoming black at maturity, at least in lower



Marasmius 213

part, up to 40 X 0.5 mm; basal mycelium white, tomentose, in some specimens rather scarce.

Spores 20-21.5 X 4.8-5.3 M. Cortical layers: epicutis of pileus with broom cells marbling the surface in scalp view, some with pale melleous fuscous, some with hyaline setulae, exuding a very short-lived pink pigment dissolved in ammonia medium.

On leaves of Monocotyledonoes in rain forest. MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, Guayaramerfn, 16 Mar 1956, Singer

B 2045 (LIL), type. ILLUSTRATION. Singer (1965), pl 18, fig 100 M.

160g. Marasmius haematocephalus var transiens Singer, Sydowia 18: 337. 1965. Mlarasmius haematocephalhs var transiens Singer, Sydowia 12: 120. 1959, ad int.

Pileus a mixture of rusty brown and dull pink to brown red (e g "chinook" in center, "Etruscan red" M&P on margin or 5-E-12 in center and margin "Jalapa" to "Sonora" or else a mixture of "Spanish cedar," "Indian red" and "Maroon" M&P), rarely some specimens of a group entirely rusty ("chinook") dried colored like M. helvolus (e g 13-J-10/11, i e a dull brown near "sorrel"), 3-12 mm broad. Lamellae white or whitish, 8-11, moderately broad to broad, free or very narrowly adnexed. Stipe white, rarely pale lilac at apex, soon umber below and at maturity mostly black or blackish but light sordid umber again when dried 30-40 X 0.2-0.5 mm; basal mycelium usually abundant, white.

Spores(12.5-) 13-22 X 3.5-4.5 u. Cortical layers: epicutis of pileus 12-16 X 5-11 , main body sometimes thick-walled, hyaline to golden hyaline, setulae all equally golden hyaline to light golden, sometimes many curved, generally thin, 3-5.5 A long.

Other characters as in type variety. On fallen leaves of Pteridophyta, Monocotyledones, Dicotyledones (Driopteris,

Gramineae, Melanostomaceae. etc) also on small woody sticks fallen to the ground. MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Coroico, 1600 m alt, 26 Jan

1956, Singer B 592 (LIL), type. ARGENTINA. Tucumnn: Tafi, Parque de Aconquija, 850 m alt, 1 Mar 1949, Singer T 171 (LIL), paratype.

This variety apparently occurs in tropical Asia since M. senmipellucidus is a synonym of it hlaelmtatocephalus (see Singer 1958a, p 121) and the way it is described by Petch and distributed by Sydow, it fits best in this variety. It is probably widely distributed in the tropics but has been determined under other names, especially M. haematocephalus, without indication of precise colors.

ILLUSTRATION. Singer (1965) pi 18, fig 100 T.

160h.Marasmius haematocephalus var atroviolaceus Singer, Sydowia 18: 337. 1965. Pileus deep violet (48-1-7) or grayish lilac, livid, dried deep reddish chocolate

color ("falcon") in center, margin fuscous-bay (between "coffee" and "trotteur tan" or cinnamon beige, between "Mindoro" and "India spice"), 8-20 mm broad..Lamellae white, at first with lilac interlamellar spaces, rather distant to very distant (10-12), not or very indistinctly intervenose, narrow, free to adnate. Stipe umber with lilac apex or umber with white apex, later black (in dried material again appearing dirty umber) with pale lilac to white apex, sometimes with concolorous apex, 20-45 X 0.2-0.3 mm; basal mycelium tomentose, white, abundant. Extreme apex underneath the lamellae sometimes orange (where var macroceplalus is brown).

Spores 19-22 X 4-4.8 . Hymenium: Cheilocystidia like epicuticular broom cells




but entirely hyaline. Cortical layers: epicutis of pileus with broom cells which have a mottled (in the grayish lilac forms) or uniform (in the deep violaceous specimens) appearance and appear in mass deep brown-red to spadiceous, the individual setulae porphyry brown to fuscous.

On monocotyledonous leaves and on dicotyledonous sticks and branchlets fallen to the ground in rain forest.

MATERIAL STUDIED. BOLIVIA. Pando: Manuripi, Conquista, 25 Mar 1956, Singer B 2189 (LIL) type, B 2241 (LIL), paratype; Beni: Guayaraiierin, 6 Mar 1956, Singer B 1620 (LIL), paratype.

ILLUSTRATION: Singer (1965), p 1 18, fig 100 A.

160i. Marasmius haematocephalus var leucophyllus Singer, var nov. TYPE. Singer B 738, from Bolivia. Pileo neque purpureo neque rubro neque rufo sed opace brunneo. Typus in LIL. Pileus dull brownish (between "cookie" and "tortoise shell" or 13-E-9, sometimes

more light fawn color to pinkish cinnamon (e g "Titian gold" with "Toltec" margin) but without ferruginous or orange or reddish tones, and dried characteristically near "Arab" or 13-B- 1 in small center and near "Sayal brown" or "russet brown" or "terrapin," hazel (Ridgway) in wide margin zone, glabrous deeply sulcate, convex, sometimes slightly papillate or umbilicate, eventually expanding to subapplanate, 5-15 mm broad. Lamellae white or whitish, in older specimens sometimes slightly intervenose but generally well separated, usually equal but sometimes with 1-2 lamellulae, 6-12 through-lamellae, distant, narrow to medium broad, free to adnexed. Stipe umber or deep umber with white apex when young, glabrous, shining, smooth, equal, setose, 15-50 X 0.2-0.4 Inm: basal mycelium white or whitish, sometimes becoming partly brownish when dried, al-

ways well developed. Context very thin except in small center, white, inodorous. Spores 15-20 X 3.5 p, fusoid to clavate, hyaline, smooth, inamyloid. Hymenium:

basidioles fusoid; cystidia well differentiated and numerous as in M1. haelatocephalus: cheilocystidia like epicuticular broom cells but mostly hyaline. Hyphae hyaline, filamentous, thin-walled, strongly pseudoamyloid, with clamp connections. Cortical layers: epicutis of pileus hymeniform with broom cells of the Siccus-type, brownish fuscous when seen in mass but individually with usually hyaline main bodies up to 8 abroad (or spreading to 11 diameter) cylindric, clavate or vesiculose, thin-walled, setulae hyaline to melleous in alternating rows or zones or all pale melleous, medium sized.

On a great variety of monocotyledonous and dicotyledonous hosts, woody sticks and stems, petioles, leaves rotting in forests or plantations, solitary to gregarious, not cespitose.

MATERIAL STUDIED. BOLIVIA. La Paz: Charobamba, 30 Jan 1956, Singer B 738 (LIL), type; Coroico, 1700 m alt, 30 Feb 1956, Singer B 739 (LIL): below Coroico at 1500 m alt, 29 Jan 1956, Singer B 670 (LIL). Also numerous collections from l:orida, Cuba, Mexico.

This variety has been misdetermined by Singer (1965) as MA. izmsicola Murrill but differs in the characteristics of the cystidia. I emphasized in my annotations (1965 p 295) however, that this form is probably a variety of M haemnzatocephalus. It is now introduced as such under a name borrowed from Berkeley & Curtis who however put it under Marasmius helvolus (Cuban collections).

ILLUSTRATION. Singer (1958a), fig 34. OTHER MATERIAL STUDIED (variety not determined). U.S.A. New York: Clyde,

18 Jul 1887, Cook 1538 (BPI). Florida Highlands County, Highlands Hammock State Park, 2 Sep 1942, Singer F 496 (F). BAHAMAS. New Providence, E. G. Britton 612 (NY), type of M. atro- purureus. MEXICO. Oaxaca: Huautla de Jimenez, 1400 m alt, Jul 1956, Singer :Chiapas, Finca



Marasmius 215

Sospiro, 4 Aug 1969, Singer M 8976 (F), near Chontalpa 31 Jul 1969, SingerM 8776 (F); Veracruz: 7 km S of Montepio, Estaci6n Biologica de los Tuxtlas, 20 Jun 1969, Singer M 8064 (F); Puerto de Veracruz, 2 Oct 1962, Yolanlda Ayala (Herbarium ENCB). CUBA. Sanvalle 1868, "misit Wright" (4) (FH), as M. hypophaeus, Wright (as Al. atrorubens) (FH); Sebaruco, 9 Jul 1941, comm. White (I:H); Cienfuegos, Soledad, 9 Jul 1941, L. W. White (BPI). TRINIDAD. Arena Forest 2 Aug 1947, Baker 1488 (K). COLOMBIA. Garz6n, 29 Nov 1939, Garces 0., det. Charles (BPI); Buenaventura, Singer (COL). VENEZUELA: Sucre, Rio Media, 15 Jul 1972, Dumont et al VE 5127 (NY). BRAZIL. Amazonas, road from Humaita to Labrea, 29 Nov 1966, Prance, Pena & Ramos 3461 (F).Minas Gerais, Agricultural College lands, Localidad Vicosa, 5 Mar 1930, Ynes Mexia 4429 (UC). ECUADOR. Napo, Lago Agrio, 9 May 197 3, Singer B 7328 (F). BOLIVIA. La Paz: Nor-Yungas, La Cienaga, E. R. de la Sota, comm. & det. Singer B 1067 (LIL); Suapi, 1250 m alt, 9 Feb 1956, Singer B 1045 (LIL); Charo- bamba, 1200 m, 30 Jan 1956, Singer B 740 (LIL). ARGENTINA. Misiones: Frontera, General Man- uel Belgrano, 19 Apr 1957 Singer M 1090 (LIL); Salta: Guemes, Quebrada de Yaquiasme, 800 m alt, 7 I eb 1957, Singer T 2938 (LIL), T 2956 (LIL); Tucuman: Capital, Parque Avellaneda, 22 Mar 1949, Singer T 241 (LIL); Jardin del Instituto Miguel Lillo, 14 Jan 1952, 20 Feb 1955, T 2191 (LIL); Buru- yacii in subxerophytic woods, 19 Feb 1957, Singer T 2992 (LIL); Tafi, Parque Aconquija, 800 m alt, 12 Mar 1949. Singer T 222 (LIL); Dique de Escaba, 31 Mar 1957, Singer 30671 (LIL). Also material from tropical Africa (see Singer 1964b p 377) and Asia.

The material from New York is probably adventitious.

Stirps Haedinus (no. 161-162)

161. Marasmius haedinus Berkeley, Hook. Jour. Bot. 8: 135. 1856.

161a. Marasmius haedinus var haedinus TYPE. Spntce 31, from Brazil. Pileus whitish or white all over, sometimes at first pale buff but soon pallescent,

mostly becoming pure white at maturity, often with a small central area remaining buffish-ocher reaching "topaz," glabrous, distantly and widely sulcate, campanulate to campanulate-convex, later more repand, convex, with or without an umbilicus or a papilla, 10-23 mm broad. Lamellae white, sometimes buffy white, distant with (3-)8-10 through- lamellae and several lamellulae, the smaller ones often with only 1-3 lamellulae, narrow to broad, sometimes some not reaching the margin, attenuate-free or subfree, not intervenose. Spore print white. Stipe reddish-brown or chestnut but reaching blackish brown or chestnut black and then sometimes becoming paler when dried, but apex at first white or buffy white, glabrous, equal or slightly attenuated upwards, sometimes with a widened base or a socle-like base, hollow, 13-20 X 0.3-1 mm; basal mycelium scarce or more often abundant, palest fulvous or more often white. Context white, thin to very thin in the pileus, inodorous or almost so.

Spores 10-14.5 X 2.8-4 , mostly 12-13.5 X 3-3.7 p, fusoid to clavate, smooth, hyaline, inamyloid. Hymenium: basidia 17-18 X 5.5 ,, 4-spored; basidioles fusoid; cystidia numerous though not crowded, well differentiated, 20-37(-50) X 6.5-16.5 (-21) u, fusoid, ventricose, even subvesiculose or irregularly cylindrical, often wavy with apical constrictions or with a subapical constriction, sometimes the majority, sometimes very few appendiculate with irregular apical appendages, hyaline, with the wall 0.4-1.5 , thick;cheilocystidia like epicuticular broom cells. Hyphae in the trama of the pileus filamentous, thin-walled, but in mature specimens some thick-walled (wall to 2 ,) ones present, hyaline, with clamp connections, but sometimes some secondary septa present, all pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells which are entirely hyaline or the setulae (especially in the youngest stage and in the center of the pileus) palest golden melleous, main body 11-22 X 4.5-12%, setulae 3.5-7 X 0.7-1.7 p, all strictly erect or somewhat diverging but always apical, conical, acute.




On dead fallen leaves or small sticks of both monocotyledonous and dicoty- eldonous hosts, in tropical-montane as well as tropical lowland rain forest and plantations.

MATERIAL STUDIED. TRINIDAD. N of Arima, 25 Sep 1949, Defnnis 56 (K). COL- OMBIA. Valle, Buenaventura, San Joaquin, 19 Apr 1968, Singer B 6228 (1:), 22 Apr 1968. Singer B 6280 (F); Quinta La Flora between Cisneros and C6rdoba, 23 Apr 1968, Singer B 6322 (F). BRAZIL. Para, Estancia Pirelli 7 Jun 1966, Singer B 4212 (BAFC); Amazonas, "Panure" (= Ipanure, now Sao Jer6nimo), Spnuce 31 (K). type. BOLIVIA. La Paz: Nor-Yungas, Coroico. 1700 m alt, 27 Jan 1956, Singer B 590 (LIL).

Illustration: Singer (1965) pi 18, fig 102.

161b.Marasmius haedinus var minor Dennis, Trans. Brit. Mycol. Soc. 34: 421. 1951. TYPE. Dennis 57 A, from Trinidad. Pileus only 5-10 mm, lamellae slightly less distant. Otherwise like the type var-

iety (see also Dennis 1951a where a full description and a colored figure, 1, 19 fig. 10, is given). Trinidad.

Dennis (1951a) describes also a var minrlor forma brevispora Dennis which is said to differ in spores 8 X 3.5 p. I suspect that this is, if cystidiate, the following species or, if acystidiate, Marasmiius microhaedin,us Singer (no. 133). I have not studied it.

162. Marasmius splitgerberi (Montagne) Singer, Sydowia 18: 353. 1965. Fig 73. Agaricus splitgerberi Montagne, Ann. Sci. Nat. 11. 18: 241. 1842. TYPE: Splitgerber 1280, from Surinam. Pileus white, sometimes a very pale buff in the center, glabrous, somewhat to

rather strongly sulcate, conical-convex or convex, with or without a small papilla, even-

tually flattened around the papilla or umbilicate when dried, 2-9 mm broad. Lamellae white with white edge, broad or medium broad, medium close to distant (ten to twenty through-lamellae. 0-several lamellulae), not intervenose, free or subfree. Stipe ochraceous yellow or fulvous to reddish umber, fuscous or chestnut, with white apex, ? shin-

ing, smooth, glabrous, subequal or equal or with a slight bulb or disc at the base, 10-25 X 0.1-0.3 mm, bulb if present up to 1 mm across: basal mycelium very scarce to abundant, consisting of radial, white fibrils covering a small zone around the base. Context

very thin, white in the pileus, inodorous.

Spores (7-)9-12.5 X 2.7-4 , fusoid or fusoid-clavate, smooth, hyaline, inamyloid. Hymenium: basidia 16-22 X 4.8-8 p, basidioles fusoid: cystidia varying from fairly numerous (but not crowded) to very scarce and often restricted to few cells among tle

cheilocystidia, (17-)18-33.5 X (5.5-)7-9.5 ,. versiform, mostly fusoid to clavate or ventricose to subvesiculose, rarely with a broad mucro appendage or lateral bulge, often many or all not projecting beyond the basidia and basidioles, hyaline, often somewhat opalescent; cheilocystidia like the epicuticular broom cells. Hyphae of the trama of the pileus hyaline, somewhat unequal, with thin or up to 0.8 , thick wall, pseudoamyloid, with clamp connections. Cortical layers: epicutis of the pileus lhylleniform, consisting of broom cells, main body 12-27 X 5-10 u, clavate, broadly fusoid or pear-sliaped, hyaline: setulae 2.5-6.2 X 0.2-1 p, mostly relatively thin, either straight or curved, some forked, also hyaline.

On fallen monocotyledonous and dicotyledonous leaves, petioles and small stems. Known host: Cecropia sp.

MATERIAL STUDIED. TRINIDAD. Non-Pareil, Sangre Grande, 11 Oct 1949, Dennis 164 (K). SURINAM. Paramaribo, 1837, Splitgerber 1280 (L) type. COLOMBIA. Valle: Buenaventura, San Joaqufn, 22 Apr 1968, Singer B 6283 (F). BRAZIL. Pernambuco: Camaragibe, 6 Jul 1960, Singer B 3108 (BAFC). BOLIVIA. Beni Vaca Diez, Guayaramerin, 6 Mar 1956, Singer B 1595 (LIL).



Marasmius 217

The specimens with few or no pleurocystidia were formerly determined Marasmius subrotula by Dennis (195la) and myself (1965) but this latter species differs in larger spores and more pigmented pileus. Marasmius splitgerberi differs from M. microhaedinus, even if there are but few cystidia, in the spores reaching only 9 p in length in the latter species which is also characterized by close lamellae.

Stirps Ferrugineus (no. 163-169)

163. Marasmius hypophaeus Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 298. 1869. Fig 74.

TYPE. Wright, from Cuba. Pileus dried rusty brown or rusty orange, not pallid-striped but at first unicolor-

ously "leafmold" (M&P), later "henna" in the center and along the ridges of the sulcate region, between ridges reaching "Mandalay" (M&P), broadly and rather shallowly sulcate over 2/3 to 5/6 of the radius, glabrous, shallowly conical and subumbonate but at first often conical and papillate, eventually more obtuse and more repand, even with umbilicus 7-17 mm broad. Lamellae pale grayish cream or cream, with at least partially discolorous (rust-brown) edge, on drying tending to become whitish to brownish with more distinctly discolorous edge, subdistant to more often distant (11-14 through- lamellae) narrow to rather broad and even ventricose, not intervenose, all equal or with few lamellulae, narrowly adnexed to rounded-free. Stipe rusty brown to chestnut brown, becoming gradually deep chestnut below, with at first cream colored or whitish pallid apex, the whole stipe tending to become light to deep umber in the herbarium, glabrous, equal or tapering upwards, sometimes with slightly broadened base, 8-43 X 0.8-1.5 mm: basal mycelium scanty to rather abundant, whitish to yellowish, sometimes developing a himantium-like pellicula on the substratum. Context very thin in pileus, inodorous.

Spores (12-)14.5-21.5 X 3-5.5 p, mostly 16-20 X 3-5.2p, fusoid with the inner side somewhat concave in lateral view, at times slightly curved with long-effilate lower portion, smooth, hyaline, inamyloid. Hymenium: basidia 25-30 X 6-8.5 P, 4-spored; cystidia 20-60 X 6-13 p, rather numerous, sometimes very numerous and crowded, versiform, often clavate or subulate, fusoid-ventricose, cylindrical or broadly ventricose, acute or more fre- quentlv broadly rounded at the apex, but occasionally or frequently with a subapical or apical appendage or mucro, well differentiated andrefringent-opalescent,without differentiated contents, hyaline: cheilocystidia like the epicuticular broom cells. Hyphae of the pileus-trama hyaline, many with somewhat firm to slightly thickened wall, with clamp connections, pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform, consisting of broom cells with a main body 8-23 X 5-10p, extremely versiform but often clavate, with the wall often thickened in the upper portion, entirely subhyaline or with the upper portion golden melleous to ochraceous brown, setulae pale brownish ochraceous or deep ferruginous, 1-14(-20) X 0.7-2.2 p, epicutis often mottled in scalp view.

On leafmold and rotten wood, also on accumulated herbaceous stems, gregarious. MATERIAL STUDIED. CUBA: 1857, Wright 129 (FH), proposed lectotype. MEXICO. Mex-

ico, Tla(l)nepantla, rubbish heaps near the village, 27 Jul 1958, A. May iVah (no number), (Herb- arium, Politecnico, Mexico). COLOMBIA. Valle, Buenaventura, Calima, 20 Apr 1968, Singer B 6262 (F), 24 Apr 1968, Singer B 6355 (F), Identity somewhat doubtful. ECUADOR. Napo, Shushufindi, 15 May 1973, Singer B 7454 (F).

This species is by no means the same as Marasmius haemzatocephalus as Murrill (1915) thought, even though the color of the fresh pileus is not known with certainty. Dennis




describes and illustrates a species under the name of Marasmiuis ferrugineus which is tawny when fresh and has tawny gill edges (Dennis, 195 Ia, p 424, pi 19, fig 3). The discolorous edges of the lamellae would tend to indicate that M. ferrugineus sensu Dennis (195 la) is indeed the species described here as M. hIvpophaeus. We may then conclude that the color of the pileus when fresh is tawny (near "old bronze" M&P) and not red (blood-red- rufous) as described by Berkeley & Curtis and assumed to be true by Murrill. It may be that other collections of this species are indeed red when fresh (I have not studied any others) but then these were confused with M1. hiypophaeus as interpreted here on the basis of Wright 129 (FH, no number on specimen). This specimen then is here proposed as lectotype of A. hypophaeus. Otherwise there would be no valid name for the species we are describing above.

Marasmius Ilhpophaeus in this sense is closer to AM. ferrugineus than to l. haemnato-

cephalus and differs in the much more distinctly discolorous (rust-brown) edges of the

lamellae, slightly larger size and perhaps the colors of the pileus. Marasmius hvlpophaeus differs from M. mointagnealnus by discolorous edges of the

lamellae and the smaller size of the carpophores and their colors.

164. Marasmius guzmanianus Singer, sp nov Fig 75. TYPE. Guzmlzl 2601, from Mexico. Pileo badio, plus minusve 54 mm lato; lamellis flavido-albis. subdistantibus; stipite

85 X 5 mm, mycelio basali albo. Sporis 13.5-16 X 3-4.5 ;: cystidiis numerosis; cellulis

epicuticularibus dimorphicis, aut aureo-succineis et crassitunicatis aut hyalinis et tune

frequenter absque setulis. Ad lignum putridum in Mexico. Typus in Herbario Instituti

Polytechnici Mexicani conservatur. Pileus dried near "Mohawk" (M&P) i e a deep reddish brown with paler spots

around the umbonate center, glabrous and slightly rugose when dried, about 54 mm broad when dried. Lamellae yellowish white when dried, with not discolorous edges, medium broad to rather broad, subdistant, intermixed with lamlellulac, strongly anastom-

osing-intervenose, narrowly adnexed. Stipe dark brown below, lighter brown at apex when mature and dried, glabrous, hollow, subequal, 85 X 5 mm; basal mycelium white. Context white in the pileus, fleshy, relatively thick.

Spores (8-)13.5-16 X 3-4.5 p, mostly about 14.5-15.5 X 3.5-4 , fusoid to cylindric oblong, in profile mostly with applanate inner side, hyaline, smooth, inamyloid. Hy- menium: basidia 4-spored; cystidia numerous, e g 28 X 9 u versiform, some appendiculate, more or less opaque, hyaline, with homogeneous contents: cheilocystidia like the

epicuticular broom cells but entirely hyaline, only very few of them pigmented. Hy- phae of the pileus trama hyaline, with clamp connections, very strongly pseudoamyloid; those of the stipe yellow in KOH, some thick-walled, parallel with each other. Cortical

layers: epicutis of the pileus hymeniform, consisting of broom cells, main body 8-18 X 5-14 ,, clavate to subvesiculose. entirely or in the upper portion with characteristically thick wall which is yellow to golden succineous, often lobed, setulae 2-6 X 0.3-2(-3) P,

rod-shaped or tapering upwards to a subacute to obtuse tip, golden-succineous, often

staggered, sometimes undulate or forked; among these broom cells there are many com-

pletely hyaline cells with low obtuse setulae or without any setulae (entirely smooth). Covering of the stipe at the apex of the latter consisting merely of a few occasional setulae not more than 6 X 2.2 p, rising directly from the subjacent hyphae of the rind of the stipe.

On rotten wood (probably Abies) among mosses and earth, in the montane zone of Mexico.



Marasmius 219

MATERIAL STUDIED. MEXICO. Mexico, La Marquesa, Parque Nacional Miguel Hidalgo, Sept 1960, 2800 m alt, Gtuznan 2601 (ENCB) holotype, (F), isotype.

This species is characterized by the structure of the epicutis of the pileus and the large size as well as by the habitat. It is named for the collector.

165. Marasmius montagneanus Singer, nom nov Marasmius montagnei Singer, Sydowia 18: 339. 1965, non Bataille (1919). TYPE. Singer B 2089, from Bolivia. Pileus fresh rarely unicolorous but striped or with a star-shaped design, sometimes

orange ferruginous ("mosque") in center, otherwise ochraceous and ferruginous or golden ochraceous brown and pallid, or tawny to cinnamon and pallid ("cornhusk," "Antique gold," "Titian gold," 12-F- 11, 13-8 C-10 M&P), dried "Saratoga, "gold brown," "Sudan brown" (M&P) or Kaiser Brown (Ridgway) or "toltec" (M&P) in marginal zone and "Peruvian brown," "gold pheasant," "raw sienna" to "gold brown" in central portion, the pallid stripes where present in fresh material turning "Inca gold" (M&P) dried, glabrous, distinctly sulculate or sulcate, smooth in center, campanulate convex, eventually irregularly applanate, with obtuse center or with an umbilicus and sometimes with a minute papilla in the umbilicus, the margin tending to become uplifted, (10-)13-28 mm broad. Lamellae white to cream, with the interlamellar spaces often sterile and then yellowish brown ("harvest") colored mostly all simple, but in some mature specimens some forked, intermixed with lamellulae or all equal, in the first case often somewhat intervenose but not strongly so in most specimens and only in mature caps, narrow, the througl-lamellae spaced in large distances and therefore extremely distant but if strongly internixed sometimes appearing rather subdistant, free to decidedly adnexed. Stipe fuscous to chestnut-black with at first white apex which darkens to "Inca gold" and eventually becomes concolorous with base, glabrous, smooth, slightly shining, filamentous-subsetaceous, equal or 20-66 X 0.3-2 mm; basal mycelium well developed, strigose or tomentose or mostly both, white when fresh, more rarely sordid fulvous-whitish, strigose portion becoming partly fulvous in many dried specimens. Context very thin, white, unchanin,g, inodorous.

Spores 14.5-21.5 X (2-)3-4.7 p, circular to clavate or oblong applanate inside when seen in profile, convex outside, hyaline, smooth, inamyloid. Hymenium: basidia 19-30 X 5.5-9 p, mostly near the lower limits and only those developed after most basidia have been formed, reaching larger sizes, 4-spored, sometimes a few 2-spored (among those formed late); basidioles fusoid; pseudoparaphyses sometimes noticeable, 20-32 X 9.3-10.5

n, mostly clavate; cystidia varying from poorly to well differentiated, numerous, 27-43 X 5.5-11 ,, well visible in KOH and Melzer, versiform, most frequently basidiomorphous or ventricose, sometimes subcapitate or appendiculate at apex, generally broadly rounded, rarely acute, sometimes cylindric or ampullaceous, hyaline, somewhat opaque, deep-rooting or not, usually not strongly projecting; cheilocystidia intermixed with basidioles and cystidia like epicuticular broom cells, but generally entirely hyaline or with melleous setulae, with main body 14-18.5 X 6.7-7 u. Hyphae hyaline, thin-to firm-walled, often swollen (to 25 u across), pseudoamyloid with clamp connections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, main body of broom cells generally smaller than that of the cheilocystidia, generally concolorous with setulae i e either hyaline or melleous (and therefore scalp preparations mottled) rarely the zone around the setulae with deeper (chestnut colored) pigmentation, setulae all erect, 2.7-1 1.5 X 1-1.5 p, rodshaped or spinulose-elongate or pencil-shaped, hyaline, in

other cells melleous or golden melleous, not strictly or consistently acute.




Generally on all kinds of leaves in rain forest leafmold. MATERIAL STUDIED. FRENCH GUYANA. Cayenne, Leprieur 990 (PC), Lcprieur 1023

(PC). BOLIVIA. Beni: Vaca Diez, Guayaramerin, 17 Mar 1956, Singer B 2089 (LIL) type: 3 km below Guayaramerin in virgin forest, 12 Mar 1956, Singer B 1889 (LIL); La Paz: Nor-Yungas, Rfo Yariza, 16 Mar 1956, Singer B 1180 (LIL); Charobamba, 20 Jan 1956, Singer B 738 (LIL).


166. Marasmius tenuisetulosus (Singer) Singer, comb nov Marasmiius graindisetulosus var tenuisetulosus Singer, Bull. Jard. Bot. Bruxelles 34: 380. 1964. TYPE. Vandervst 4015, from Zaire. Pileus rusty-tawny, striped radially paler tawny between striae, dried often more

orange ferruginous in part (e g "Agate" M&P) glabrous, deeply sulcate over 4/5th of the radius, center often appearing rugose when dried, convex, with depressed center when mature, sometimes subumbonate in the depression, 8-28 mm broad. Lamellae whitish, the edges not discolorous, medium broad to rather broad, distant, more rarely subdistant (13-16 through-lamellae, few to many lamellulae), subfree, rounded-subfree. Stipe very deep chestnut brown with white apex, tending to umber in the herbarium or blackening on drying, equal or tapering downwards, 15-30 X 0.5-1 mm: basal mycelium stri-

gose, fulvous-tawny to pallid. Context of the pileus white, very thin, inodorous.

Spores 14.5-19 X 3-4 j, fusoid-oblong, smooth, hyaline, inamyloid. Hvmenium: basidia 20-24.5 X 5.7-9 ,u, 4-spored; cystidia numerous, 30-64 X 8.2-11.5 A, versiforn, often clavate ventricose or fusoid, rounded above or with subacute apex or appendage, opaque or thick-walled, sometimes constricted in the middle, hyaline, with homogeneous contents; cheilocystidia like the epicuticular broom cells. but hyaline. Hyphae of the pileus-trama hyaline, thin- to firm-walled, not thick-walled, often very irregular in sliape and sometimes swollen to 25 p broad, with clamp connections, pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform, consisting of broom cells with the main body 5-16.5 X 3.3-11.5 A, tawny orange, in some hyaline with the wall thick or thin or thick above and thin below; setulae 5-7.5 X 1-1.7 p, erect and straight, acute or subacute, some with rounded tip, few hyaline, most ochraceous brown or tawny orange, scalp preparations appear more or less mottled.

On rotting wood of dicotyledonous trees in tropical forests. MATERIAL STUDIED. COLOMBIA. Valle: Buenaventura, Calima, 24 Apr 1968, Singer B

6354 (F). ZAIRE. Kasai: Kikwit, Vandervst 4015 (BR) type. As I have indicated before (1964b, p. 319) this fungus is extremely close to M.

montagneanus It differs mainly in the more frequently acute or subacute setulae of the

epicuticular broom cells, in the broader lamellae and in the habitat on wood rather than leaves.

ILLUSTRATION. Singer (1964b) fig 28.

167. Marasmius anomalus Lasch in Rabenhorst, Klotzchii Herbarium Vivum no. 1806. 1854 (non Peck, 1872).

Androsaceus epodius var imicrosporns Maire, Bull Soc. Mycol. France 24: Ivi. 1908. Marasmius epodius var microsporus (Maire) Kuhner, Botaniste 25: 101. 1933. Marasmius coprophlilus Spegazzini, Bol. Acad. Nac. Ci. C6rdoba 29: 122. 1926. TYPE: Lasch, from Germany. Pileus deep buff, ochraceous brown with a cinnamon shade, somewhat deeper, al-

most rusty brown in the small disc area, paler towards the margin (13-D-9 to 14-B-11), pallescent in age (to "auteuil"), eventually pallid or cream on margin, glabrous, at first smooth, later sulcate, otherwise even or somewhat rugulose, campanulate to convex, then



Marasmius 221

convex to applanate, center eventually often depressed or umbilicate, sometimes somewhat irregular, 5-9(-15) mm broad. Lamellae white, more rarely cream, with concolorous (with sides) or cinnamon colored edges, distant, 10-20, mostly 14-17 through-lamellae, 0-1, rarely 2 lamellulae, rarely slightly intervenose, rather broad to broad, more or less ventricose, rounded-free, sometimes with an indistinct collarium (which is not quite free from apex of stipe or imperfect). Stipe concolorous with pileus or darker in the middle portion, at apex at first white, at base often reaching blackish, gradually becoming darker umber brown from base upwards, e g "clove," glabrous and smooth, slightly to strongly sining, equal, setose, seemingly insititious but rarely without any mycelial tomentum when seen under a good lens in dry condition, 13-50 X 0.3-1.5 mm; basal mycelium rarely absent, usually distinctly conspicuously strigose to pubescent-tomentose, whitish. Context very thin except at the very disc, unchanging, inodorous.

Spores (10-) 12-16.7 X 3.3-5.5 u, clavate-elongate or cylindrical-subfusoid, smooth, hyaline, inamyloid. Hymenium: basidia 21-30 X 6-7.5 A, clavate, 4-spored; basidioles fusoid; cystidia 30-83 X 8-14 ,, versiform, often fusoid, clavate or cylindric, sometimes upper portion subulate but not acute, mostly broadly rounded at tip, not or not strongly opaque-refrigent, not or not strongly projecting in most cases but sometimes very deep-rooting, often very slightly incrusted; cheilocystidia like epicuticular broom cells. Hyphae strongly pseudoamyloid, with clamp connections. Cortical layers: epicuticular layer of pileus hymeniform, consisting of broom cells of the Siccus-type, these 16-17 X 5-8.8 ,, clavate or ventricose, sometimes shallowly bifurcate, hyaline below but the setulae golden melleous or deep buff, among the broom cells few basidiomorphous smooth cells present, at least when caps are old.

On grass leaves or roots, culms and generally rotting herbaceous rests or dung, on the earth in meadows, usually gregarious, but not cespitose.

MATERIAL STUDIED. GERMANY. Driesen, Lasch (FH) type. Bei Schwetzingen, on Ainarantllus retroflexus 6 Aug 1937, collector unknown (FH), as Agaricus (Collybia) calopus. ITALY. Trentino, Bresadola (BPI). ARGENTINA. Tucuman: Taft del Valle at 2100 m alt, 13 Jan 1950, Singer T 870 (LIL), 10 Feb 1961, T 1182 (LIL); Cordoba: Alta Gracia, Bruch 67 (LPS) type of M. coprophilus. CHILE. Santiago: Estaci6n Experimental, potrero near Bodega de Vinos, May 1966, Lazo (SGO).

This is not a typical neotropical species but enters the American subtropics at least in the Southern Hemisphere.

Marasmius anomalus is characterized by the peculiar color of the pileus, the small size and the medium large spores.


168. Marasmius phaeocystis Singer, sp nov TYPE. Guerrero 61, from Argentina. Pileo rubido-brunneo vel ferrugineo, 5-10 mm lato; lamellis albidis acie pileo con-

coloribus; stipite subatro, mycelio basali strigoso, sparso, albo. Sporis 15-18 X 4.5-4.8 ,; cystidiis numerosis, contento oleaginoso destitutis sed neque hyalinis nec melleis sed dilute fuscis. Ad ramulos dicotyledoneos in silva subtropicali Argentinae. Typus in LIL conservatus est.

Pileus "cocoa," dried nearer "Alamo" with "montella" disc, i e ferruginous, glabrous, deep and long sulcate with venose-rugose disc, convex, some almost umbilicate or umbonate, some more campanulate when young, some more applanate when old, 5-10 mm broad. Lamellae pallid whitish with the edges concolorous with the surface of the pileus, distant (12-15), equal, free (sometimes in part leaving a very narrow, rusty zone around the apex of the stipe), medium broad to rather broad. Stipe almost




black when fresh, blackish brown to deep umber when dried, glabrous, smooth, 25-39 X 0.2-0.9 mm; basal mycelium strigose, not extensive, white. Context thin, inodorous.

Spores 15-18 X 4.5-4.8 ,, hyaline, thin-walled, subclavate, smooth, inamyloid. Hymenium: basidia 25-29 X 6.3-8.3 p; cystidia 27-59 X 5.5-9 p, fusoid or clavate, rounded to subacute at tip, rarely apex incrusted with a rusty granular-resinous incrustation, without a conspicuous oily content but wall rather thick to very thick and of opaque aspect, deeper rooting than the basidia, light fuscous (not hyaline or melleous!) except rarely in a hyaline apical zone, smooth; cheilocystidia like the epicuticular elements

although never thick-walled. Hyphae thin-walled and hyaline in the pileus and the hymenophoral trama, pseudoamyloid, with clamp connections. Cortical layers: epicutis of pileus consisting of a hymeniform layer of broom cells of the Siccus-type, mostly hyaline in the main body with golden melleous setulae, more rarely setulae also hyaline or main body more or less golden, the latter thin-walled, rarely thick-walled, 9.5-21 X (3-)5-8 1, setulae thin-rodshaped or spinulose, obtuse to subacute, 2-7 X 1- 1.5 M, few occasional epicuticular elements smooth (without setulae).

On fallen dead twigs of dicotyledonous tree in subtropical forest, perhaps also on leaves, gregarious.

MATERIAL STUDIED. ARGENTINA. Misiones: Arroyo Piray Guazui, 24 Feb 1960, R. T. Guerrero 61 (LIL), type.

This species was misinterpreted by Singer (1965), as being identical with MIarasmtius paucifolius Murrill. However, the type of that species turned out to have hyaline cystidia and other colors; see also under Marasmius ferrugineus no. 169).


169. Marasmius ferrugineus (Berkeley) Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 297. 1869.

Agaricus ferrugineus Berkeley, Lond. Jour. Bot. 2: 630. 1843, non Persoon ex Secretan (1833). Marasmius paucifolius Murrill, North American Flora 9: 262. 1915 (= var gardneri). Marasmius gardneri Singer, Sydowia 12: 114. 1958 (= var gardnleri).

169a. Marasmius ferrugineus var ferrugineus TYPE. Gardner, from Brazil (Berkeley Herbarium, K). Pileus bright ferruginous-fulvous (e g between "copper lustre" and "spice," dried

between "gold pheasant" and "Sudan brown" or "burnt almond"), glabrous, long sulcate, campanulatc-convex, 3-11 mm broad, rarely broader. Lamellae pallid to pale brownish, edges not discolorous, not anastomosing but with an occasional vein connecting two of of (9-13) through-lamellae, all equal, free to adnate, not collariate, distant. Stipe cinereous-umber to umber-fuliginous, glabrous, smooth, shining, setose, equal. 11-30 X 0.3 mm; basal mycelium distinct to sparse, tomentose-filamentose, palest brownish-white, causing the base to appear slightly thickened. Context very thin; odor none.

Spores 15-21 X 2.8-4.3 p, fusoid to clavate or acicular sometimes in some collections tending to becoming 2-3-celled when over-mature, hyaline, smooth, inamyloid. Hy- menium: basidia clavate, basidioles fusiform; cystidia numerous, 28.5-57 X 7-17 Au, cylindric or ventricose, often irregularly shaped, more rarely clavate or ampullaceous, at times mucronate or appendiculate opaque, refringent, hyaline to pale melleous (in dried material, KOH or NH40H) sometimes all hyaline; cheilocystidia similar to the broom cells of the epicutis. Hyphae hyaline and thin-walled and pseudoamyloid in trama of pileus, all with clamp connections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, main body up to 6.5 u broad, hyaline or pale golden,



Marasmius 223

setulae in some cells golden hyaline to pale golden melleous, in others melleous brown or succineous, thus causing a mottled effect in scalp preparations, all vertical and narrowly conical, 4-5.5 1 long.

On dead pieces of bark and dead branches of dicotyledonous trees and shrubs in tropical forest.

MATERIAL STUDIED. BRAZIL. Minas Gerais. (Berkeley set at K), holotype; Amazonas: Panure (=Ipanure, Sao Jeronimo), Spruce 139 p p (K). BOLIVIA. La Paz: Nor-Yungas, Carmen Pampa, 19 Feb 1956, Singer B 1270 (LIL).

169b.Marasmius ferrugineus var gardneri Singer, var nov TYPE. Gardner, from Brazil (Hooker Herbarium, K). Sporis minoribus. Ad frustula lignea. This differs mainly by the slightly smaller spores. The pileus often becomes some-

what darker ("cocoa") on drying and the habitat is on small woody sticks, leaf petioles and veins.

Spores 13.5-18.3 X 3.2-5 ,, i e shorter on an average and relatively broader than in the type variety.

MATERIAL STUDIED. PUERTO RICO. Rio Piedras, 6 Sep 1912, Johnston 580 (NY), type of M. paucifolius. BRAZIL. Minas Gerais, Gardner, Hooker Herbarium set at (K), type; Rio de Jan- eiro, Angra dos Reis, 1 Oct 1952, Singer B 437 (LIL).

Marasmius poecilus Berkeley is apparently a mixed collection which was singled out from Spruce's collection by Berkeley because of the supposed lack of a basal mycelium but Berkeley indicates himself that possibly under this name are included two

species. One part of the collection, as pointed out earlier by Singer (1958a) is represented by the carpophore in the FH set of Spruce 139, a carpophore with the surface of the pileus turned upward, slightly larger than the others, better preserved and with numerous spores present (20-21.8 X 2.8-3 j) and this, since it has numerous pleurocystidia like those of M. ferruggineus, is here interpreted as belonging to var ferrugineus. As for the rest of the carpophores, I did not attempt to determine them, but suppose that they are difficult to place; a guess was made (Singer, 1965) that they might be M. bambusinus. The K set of the collection contains among other material a dead leaf with five

carpophores, one loose. This is designated as the holotype of M. poecilus and it was considered to be MA. tageticolor until it turned out that M. tageticolor should be divided into two species, one based on the type of M. tageticolor and the other on the lectotype of M. poecilus This has been described under no. 117.

Marasmnius paucifolius is, according to the data obtained on the type specimen, the same as AM. ferrugineus var gardneri. Dennis (1951 a) also identified it with M. ferrugineus. The type specimen is annotated by the collector as being "dark red" or "bay." This would indicate that Johnston mixed up specimens belonging to M. haenuatocephalus but there is no indication now that originally red carpophores are among the type co- lection, inasmuch as Murrill knew M. haematocephalus well, although in a very broad sense-and omitted "dark red" from the type description. The pileus is now about "cocoa" M&P. With this color and the spore size (14.7-17 X 3.5-5 p) it could hardly be M. haenzatocephalus but the spore size fits best in var gardneri of M. ferrugineus.

As for the type assemblages of Agaricus ferrugineus themselves, they are evidently not fully homogeneous either and can be separated into a relatively large- and narrow- spored set at one hand and the shorter-spored set on the other. Since M. gardneri Sing- er was originally based on the Hooker set, which, at least the specimens checked, belongs to the shorter-spored group, I have used this name on the varietal level in order to set it




apart from the Berkeley Herbarium set which Dennis, to whom I am obliged for assistance in this matter, considers the set from which the holotype of M. ferruginleus var ferrugineus should stem. These latter specimens, at least those from which Dennis and I took our preparations, belong to the long-and-narrow-spored form. Additional collections of each of these varieties did not show more than occasionally somewhat browned gill edges near the margin, not a consistently discolorous gill edge such as we find characteristic for M. ferrugineus sensu Dennis (195 la, non 1970 which refers to our IM. ferrugineus) and our M. hypophaeus (see there no. 163).


170. Marasmius bambusinus (Fries) Fries, Epicrisis Systematis Mycologici p 385. 1838. Agaricius batmbusinus Fries, Linnaea 5: 507. 1830. TYPE. Beyrich, Brazil (not located, see notes below). Pileus orange-rufescent to orange ferrugineous or ochraceous orange to bright or-

ange ("mosque," "Monterrey," "chinook," "talavera," "carrot red," "bittersweet orange," "capucin orange" or 3/4-A-12, 12-A-ll, l-E-10 M&P) eventually pallescent, dried more brown ("talavera," "Arab," "Alamo," 13-C-12, "Amber glow"), glabrous, sulcate when mature but sometimes only slightly so or even quite smooth, hemispherical then convex and eventually applanate, obtuse, sometimes umbilicate, 2-11 mm broad, rarely reaching 11 mm. Lamellae varying from pure white to light cream, at first very narrow, later narrow to rather broad and with 0 to 3 lamellulae intermixed, these at times venose, distant at maturity, 6-13 through-lamellae present, sometimes forked but not intervenose, free to adnate and very indistinctly collariate in some specimens, usually clearly non-collariate. Stipe brown to blackish, at first with white apex, dried sordid umber to umber, glabrous and smooth, mostly shining, setose, equal, 6-25 X 0.1-0.4 mm; basal mycelium mostly distinctly showing, more rarely becoming obsolete, minutely fibrillose or strigose or tomentose, white or whitish, at times arising from a corticioid white mycelial patch. Context very thin, inodorous.

Spores 13.8-22 X 3.5-5.5 1, mostly 16.4-20 u long, clavate to fusoid-oblong, hyaline, smooth, inamyloid, subplane on inner side when seen in profile. Hymenium: basidia 22-25 X 7-8.3 u, 4-spored; cystidia 14-55 X 8-12.5 , moderately numerous to very numerous, distinctly differentiated, versiform, often ventricose-subampullaceous or vesiculose-subclavate, clavate or ventricose-subclavate-elongated, etc, opaque to refringent to subopaque, with rather thin to thick wall, mostly quite hyaline, broadly rounded at tip; cheilocystidia either quite like the epicuticular broom cells or differing from them only in being entirely hyaline, rarely very few, mostly numerous. Hyphae filamentous, hyaline and thin-walled in trama of pileus, pseudoamyloid, with clamp connections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus- type, either all subhyaline to golden hyaline and uniformly colored in scalp preparations or mottled because entirely melleous broom cells alternate with entirely hyaline cells, main body 11-21 X 4-9 , but sometimes spreading to + 17 ,, setulae 2.5-5.5 u1 long, cylindric-rodshaped, pencil-shaped or long-spinulose, thin, especially in young caps, sometimes somewhat curved and wavy, firmer and straight setulae mainly where the broom cell is pigmented.

On culms and leaves of monocotyledonous plants, more rarely on leaves of dicotyledonous plants, frequently on Musaceae, and Gramineae; known hosts: Musa, Chus- quea, Panicunz, Protium.

MATERIAL STUDIED. VENEZUELA. Bol'var, road from El Dorado to Sta. Elena, 5 Aug 1972, Dumont et al VE 7375 (NY). COLOMBIA. Valle, Buenaventura, La Brea, 24 Apr 1968,



Marasmius 225

Singer B 6345(1 ). BRAZIL. Pernambuco, Camaragibc, 6 Jul 1960, Singer B 3112 (BAFC), Singer B 3112a (1 ), 6 Jul 1960, Singer B 3119 (I ). BOLIVIA. Beni: Vaca Diez, Guayaramerin, 3 km below the village in virgin forest, 12 Mar 1956, Singer B 1926 (LIL); La Paz: Nor-Yungas, Charo- bamiba 1200-1 300 in alt, 30 Jan 1956, Singer B 735 (LIL), B 686 (LIL); Coroico, 7 Feb 1956, Singer B 944 (LIL); Road frolm Coroico to Rio Llolosa, 12 Fe:b 1956, Singer B 1106 (LIL); Carmen Pampa, 17 Iel) 1956, SinLger B 12255 (LIL).

My intcrpretation of Marasmius bainbusinus is based on the original description. We htave not studied the type and do not know whether it is preserved and if so, in what state it is. The original diagnosis describes it as having "lamellis paucis aequalibus adnatis" whicli is quite correct for our species. In Epicrisis, probably because of a lapsus plumae, Fries transcribes "lamellis adnatis paucis aequalibus venosis albis." Whether the venose lamellac are authentically observed or merely added by error, does not matter too much since young specimens, precisely those that may be compared with insititious Micena, do have very narrow lamellae which, observed on a specimen that had been

pressed flat according to the methods of preparation at the time in use, will easily appear venose and we know other cases where mycologists even more experienced in tropical agarics thant Fries in 1830, have committed the same error in their diagnoses. Having studied a large number of marasmioid agarics in and around Brazil on bamboo (and otlher monocotyledonous leaves), I cannot think of any other Marasmius which would fit this description and only in Marasiniellus do we have similarly colored (although less

brightly so) small carpophores occurring on that host.

Fries' species was redescribed and illustrated only by Bresadola (1929), as far as I am aware, and his material came from China rather than the neotropics. The color of this species is much less bright than Beyrich's ("luride fulvellus vel subochraceus" rather than rufescens), has small spores and apparently insititious stipe. This specimen (WU) is difficult to determine but it seems to be a Marasmiellus rather than a Marasmius, in fact perhaps the same as a small Marasmiellus sometimes occurring on bamboo in the neotropics. It cannot be proved at the moment which of the two interpretations is correct but if the original Beyrich material does not exist or is not in good condition, it seems that the inter-

pretation based on neotropical material ought to have preference inasmuch as it agrees with the entire description of 1830 including the color of the pileus.


171. Marasmius nogalesii Singer, Sydowia 18: 339. 1965. TYPI. Singer B 2558, from Bolivia. Pileus brown ("tiffin" to "oak") in the center and pale argillaceous ("walnut taffy")

on the margin with a rosy shade at least in parts and then becoming "raw sienna" in the center with the margin "papyrus", slightly hygrophanous in center, in dried condition

rusty ochraceous brown ("kis kilim" and "Arab") on margin and orange rusty ("Titain") in center, sulcate over most of the pileus, the center rugose-venose, campanulate, then

flattened-gibbous or subumbonate, 40-78 mm broad. Lamellae white, moderately broad

(broader than illustrated in M. dennisii), subclose to medium distant, about 17 through- lamellae in a medium-sized cap, edges not discolored, free to adnexed. Stipe chestnut with white apex, later chestnut all over or blackish-chestnut below, glabrous although at extreme apex sometimes lightly pubescent, smooth, 75-104 X 1.5-5 mm; basal mycelium white, in part turning light fulvous when dried in some carpophores. Context inodorous.

Spores 15-17 X 3.5-4 ,, clavate and slightly curved, hyaline, smooth. Hymenium: basidia 16.5-19 X 6.2-7 p, 4-spored; Cystidia 31-49 X 7-10 ,, very numerous, versiform, usually slightly attenuate from broadest portion which is in lowest third, often with con-




strictions or with appendage or mucro, hyaline, or stramineous, smooth, thick-walled and refrigent. Hyphae of hymenophoral trama regularly arranged, they and the hyphae of the trama of the pileus strongly pseudoamyloid, with clamp connections, 2.5-21 u

broad, walls up to 1.2 p thick but also often thin. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, 13-14.5 X 8-14 p, spreading sometimes to 18 u broad with pale fuscous brown main body and melleous brown setulae, the latter 6-8.5 X 1.4 ;, cylindric to slightly attenuated upwards, obtuse.

On rotten branchlets in tropical rain forest. MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, Arroyo Florida, 32 km east of Riber-

alta, 7 Apr 1956, Singer B 2558 (LIL), type. This is a large and conspicuous species of the appearance of M. amazonicus yet

without the purple color and the spots of the latter and clearly cystidiate. ILLUSTRATION. Singer (1965) pl 18, fig 104; pl, fig 105.

Stirps Siccus (no. 172-173)

172. Marasmius wilsonii Murrill, North American Flora 9: 261. 1915. Fig 76. TYPE. Wilson 297, from Puerto Rico. Pileus ochraceous-brown to orange-ferruginous brown (e g "amber br." M&P), paler

(pale bay brownish but often drying dark to "Mandalay" M&P) on the margin and frequently also in radial stripes between the sulci of mature material, dried "Ginger" M&P, "cookie," burnt umber" in the darker areas, on margin and between the sulci much paler, occasionally with small rusty spots, everywhere smooth at first, later becoming sulcate over two thirds to three quarters of the radius, glabrous, campanulate or conical, then convex, eventually often narrowly depressed in the center and sometimes subumbonate in the depression but at times remaining campanulate even when mature and then obtuse or with a small, low umbo up to 8 mm high, 5-11 mm broad. Lamellae pure white when fresh, edge also white or sometimes somewhat brown-discolorous exclusively near the margin of the pileus, subclose to subdistant, rarely distant, (e g 15-20 through-lamellae and 0-4 lamellulae), not intervenose, at first rather narrow, eventually rather broad and often ventricose, adnexed to rounded-subfree or free. Stipe at first light brownish with white apex, becoming chestnut or even dark chestnut below, again lighter brown when dried, glabrous and shining, smooth, equal, often long 20-72 X 0.2- 1 mm; basal mycelium both fibrillose and rhizomorphoid, or only consisting of radiating fibrils, white. Context of the pileus white, thin, inodorous.

Spores 11.2-15.5(-19.7) X (2.5-)3.5-4(-5);, fusoid, smooth, hyaline, inamyloid. Hymenium: basidia 23-32 X 6.5-9 1, 4-spored, few 2 -spored; cystidia distinct and rather numerous, deep-rooting, 23-62 X 5-13.7 u, either clavate or ventricose and then comparatively broad, or fusoid and then relatively narrow, with rounded-obtuse or acute tip, or with 1-2 apical appendages, some constricted near the middle, hyaline or subhyaline with or without very finely granular hyaline contents, more or less opaque, with firm to thick walls; cheilocystidia taken from the middle zone of the lamellae like the epicuticular broom cells but the great majority entirely hyaline or subhyaline and all thin-walled. Hyphae of the pileus-trama hyaline, with thin to firm, never thick wall, with clamp connections, pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform, consisting of broom cells, main body 6-25 X 5-9 ;, hyaline and thin-walled, some golden-melleous or melleous brown and then often thick-walled in mature material (wall in this case up to 2 p thick), setulae 3-8.5 X 0.5-1.8; , strictly erect, golden melleous or melleous brown,



Marasmius 227

in a few broom cells deeper brown, subacute or acute, scalp preparations appearing mottled.

On dead fallen dicotyledonous leaves, rarely small twigs, in the northern parts of the neotropics. Known hosts: Bursera, Quercus.

MATERIAL STUDIED. PUERTO RICO. Luquillo Mountains, Jul 1902. P. Wilson 297 (NY) type. MEXICO. Morelos: NW of Tepoztlan, 1800 m alt, 18 Jul 1969, Singer M 8498 (F), 1850 m alt, 2 Jul 1969, Singer M 8266 (F); Chiapas, road from Bochal to Tuxtla de Gutierrez near Soyalto, 2 Aug 1969, 1400 m alt, Singer M 8827 (F).

This species is close to M. ferrugineus as was also inferred by Dennis (1951a) but the main difference between the two species resides in the less distant lamellae and the precise colors of M. wilsonii whose older pilei are not smooth but sulcate and the lower portion of the stipe may be as dark as that of M. ferrugineus.

Some of the supposed color forms of the temperate North American M. siccus would key out with lM. ferrugineus or M. wilsonii; however, they have somewhat larger carpophores and spores than M. wilsonii and also more distant lamellae.

173. Marasmius dennisii Singer, Sydowia 18: 335. 1965. Marasnlzius dennisii Singer, Sydowia 12: 117. 1959 (ad int., nom nud). TYPE. Deinnis 42 A, from Trinidad. Pileus orange chrome to ochraceous orange, somewhat feebly striate to convex,

then expanded, sometimes slightly depressed, 15-50 mm broad. Lamellae pale buff, rather narrow (narrow according to illustration but said to be up to 4 mm broad in very large specimens, subclose ("moderately spaced"), not intervenose or only at the end of the lamellulae, tridymous, attenuated at each end, adnexed. Spore print white. Stipe dark purplish brown below, white at apex, smooth and polished throughout, hollow, cylindrical, long, 1-2 mm thick. Context of pileus white, thin.

Spores 14-17 X 3-3.5 p, tear-shaped. Hymenium: cystidia irregularly cylindric with rounded tips and yellow contents, not protruding above the level of the basidia; cheilocystidia like epicuticular broom cells. Cortical layers: Epicutis formed by broom cells with a main body of 8-10 u diameter, setulae yellow, about 7-9 X 1 ,u.

On dead leaves and litter of plantations, apparently mainly on bamboo. MATERIAL STUDIED. None. The description is adapted from Dennis (1951a, p

419, as l. leoninus). This is apparently a rare form which has come to a luxuriant development in bam-

boo plantations of Trinidad. ILLUSTRATION. Dennis (1951a), pl 19, fig 1.

Stirps Helvolus

174. Marasmius helvolus Berkeley, Hook. Jour. Bot. 8: 136. 1856. TYPE. Spruce 139, from Brazil. Pileus between "cocoa" and "leather brown", when dried not changing color or

becoming "auburn" with "pecan brown" striae (between elevation of sulcai) or hazel (Ridgway) mostly striped in this manner or palest cinnamon even when fresh, glabrous, deeply sulcate, subumbilicate, convex, then applanate, often subumbonate or with flattened disc, 9-31 mm broad. Lamellae white or whitish with ochraceous ground color (interlamellar spaces) edges not concolorous with pileus in many dried specimens and darkening beginning at edges, the ochraceous ground color often reaching up to half the breadth of the lamellae (leaving the edge portion whitish) medium broad, subdis-




tant, adnexed to adnate, sometimes to a widened apical disc of the stipe. Stipe somewhat variable in color, fresh chestnut with ochraceous pallid to pale ochraceous brown apex, then umber to liver brown, dried light tan to light fuscous, glabrous, shining, smooth, mostly equal but sometimes with enlarged base or with discoid enlarged apex, 15-32 X 0.5-1.5 mm: basal mycelium white, tomentose, often forming a small socle at base, not extensive but well developed. Context thin, inodorous or with the mawky odor characteristic for some species of Cortinarius (alboviolaceus, etc).

Spores 11-15.2 X 2.8-4 ,, fusoid-clavate to subcylindric-oblong, hyaline, smooth, inamyloid. Hymenium: basidia 15-30 X 6-7 ,, 4-spored; cystidia present but often few on sides, more numerous (among cheilocystidia) on and near edges, 19-25 X 5-10.2 u, hyaline to very pale melleous, not striking, versiform, sometimes with geniculate base, opaque and thick-walled, in general neither projecting nor deep-rooting, most frequently clavate to fusoid-ventricose, often variously appendiculate; cheilocystidia in dried material like epicuticular broom cells, main body either hyaline or pale brownish, setulae 4-7 X 1.3 p, brown or many or all cheilocystidia entirely hyaline. Hyphae hyaline to pale melleous in the trama of the pileus, mostly hyaline and thin-walled but eventually some with slightly thickened walls, in cresyl blue mounts a heterogeneous hyphal sys- tem becoming evident which is metachromatic (pinkish) while the other hyphae become pale blue, often irregular in shape, with clamp connections, pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, broom cells of Siccus-type, main body hyaline or melleous to deeper cinnamon brownish, cylindric or clavate, sometimes constricted, 8.2-11.7 X 5.5-10.3 ,, setulae 3-8 X 1-1.4 ,, either hyaline or on other cells brownish melleous or cinnamon brownish, rod-shaped and obtuse to spinulose and acute, offering a mottled surface in scalp preparations.

On dead leaves, leaf petioles, small fallen twigs and woody parts of small vines, apparently generally on dicotyledonous hosts, solitary or gregarious.

MATERIAL STUDIED. BRAZIL. Amazonas: Panure (SSo Jeronimo) Spruce 139 (K), type. ECUADOR. Napo, Shushufindi, 13 May 1973. Singer B 7412 (F). BOLIVIA. Pando: Manuripi, Conquista, 25 Mar 1956, Singer B 2235 (LIL), 26 Mar 1956, Singer B 2282 (LIL).

Marasmius helvolus is often similar in colors to M. helvoloides (see no. 130) which should be compared. AMarasmnius helvohls has somewhat inconstant cystidial characters and may be looked for in series Leonini where it is also keyed out.


Stirps Confertus

175. Marasmius confertus Berkeley & Broome, Jour. Linn. Soc. Bot. 14: 35. 1873. TYPE. Thwaites, from Ceylon (not studied). Pileus ochraceous, orange ochraceous, orange or orange rufescent, sometimes bright

orange-ferruginous, especially in center and in dry (not dried) condition since it is at times slightly subhygrophanous (but not fading on drying!) e g 10-1-7/10, "Saratoga," "orange rufous," at times (pale forms or pallescent), orange pallid, glabrous sulcate over half of the radius or more, with smooth center, with a small depression in the center and/or subumbonate, 8-25 mm broad, mostly about 11-15 mm. Lamellae white, then becoming cream, often with edges concolorous with pileus from margin part way to the stipe but not always so, rather narrow or narrow, about 22, intermixed with lamellulae, close or subclose, free to adnate. Stipe ochraceous brown to chestnut, base often umber to almost black and eventually sometimes becoming entirely brown-black, lighter above, with white apex at first, dried light umber to tawny, glabrous, smooth, shining, equal or very



Marasmius 229

slightly tapering upwards, hollow, 30-80 X 0.5-2 mm; basal mycelium strigose and tomentose, wet watery pallid to ochraceous whitish, dried white with fulvous tawny tips or all brownish fulvous; whitish to ochraceous rhizomorphs very frequently present. Context of pileus thin, white, inodorous.

Spores 10.5-13.7 X 3-4 p, fusoid-oblong, applanate inside when seen in profile, hyaline and smooth, inamyloid. Hymenium: basidia e g 21 X 5.5-7.5 p, 4-spored, sometimes 2-spored; cystidia 24-35 X 4-9 y, hyaline to melleous, moderately numerous to numerous, opaque, smooth, versiform, e g ventricose-fusoid, cylindric or clavate, ampullaceous, often mucronate or appendiculate in some collections, well differentiated; cheilocystidia golden hyaline to hyaline with more often hyaline than melleous setulae, otherwise like the epicuticular broom cells, in some collections rather scattered, in others crowded. Hyphae of trama of pileus and hymenophoral trama hyaline, some with distinctly thickened walls, others thin-walled, broad or narrowly filamentous, strongly pseudoamyloid, with clamp connections, subhymenium consisting of small elongated and small subisodiametric elements, hyaline. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, main body hyaline to golden hyaline, 8-15.5 X 6-11 ,, setulae numerous per cell, 3.5-6.5 X 1-2.2 (at base)j , alternating subhyaline to golden hyaline and melleous to tawny, acute to obtuse of subacute, rod-shaped to spinulose, surface mottled in scalp view.

On termite nests (woody material with termite holes) on rotting wood in forest, on monocotyledonous remainders and generally forest debris with woody particles, rarely solitary, usually characteristically cespitose to fasciculate, in tropical-montane and in tropical rain forest.

MATERIAL STUDIED. VENEZUELA. Miranda: Guatopo, 800 m, 23 Jun 1958, Dennis 1107 (K,v.i.). PERU. Loreto: Maynas, Rio Nanay, 28 Nov 1958, Lowy 367a (LIL). BRAZIL. Guapore, Guajara mirim, 10 Mar 1956, Singer B 1785a (LIL), B 1785 (LIL). BOLIVIA. Beni: Vaca Diez, Guayaramerin, 7 Mar 1956, Singer B 1631 (LIL); La Paz: Nor-Yungas, Las Cienagas, 10 Feb 1956, E. R. de la Sota, comm. & det. Singer B 1068 (LIL).

This differs from the preceding species in smaller spores, distinctly sulcate pileus, smaller average size and often cespitose habit.

The Venezuelan collection was compared with the type by Dennis. Petch redescribed AI1. conifertus exactly the way we observe this species in South America. He indicates as synonyms M. chondripes Berkeley & Broome and M. hemibaphus Berkeley & Broome.

ILLUSTRATION. Singer (1965) pi 19, fig 106; Dennis (1961) fig 44.

Stirps Pseudocorrugatus (no. 176-179)

176. Marasmius yarizae Singer, Sydowia 18: 345. 1965. TYPE. Singer B 1369, from Bolivia. Pileus deep reddish chestnut or red-brown in the narrow disc area, ("Mohawk,"

"Indian red"), around it ferruginous ("burnt sienna," "Saratoga") and in the marginal zone dull light orange to saturated orange (ll-H-4 to "Eldorado"), sometimes entirely colored in the color of the disc area or on the contrary never with any chestnut to brown central area, eventually generally ferruginous ("ferruginous") all over, shallowly, later deeply sulcate to the disc area, the latter smooth, umbilicate, campanulate to convex, 13-44 mm broad. Lamellae white, cream white, orange cream, edges usually concolorous with pileus, ground also sometimes concolorous with pileus (where hymenium does not bridge over the interlamellae spaces) medium broad to broad at stipe at ma-




turity but narrow near the margin, distant to very distant, 10-12 through-lamellae but lamellulae also present, adnate to the fleshy disc or merely adnexed. Stipe "cocoa brown" to chestnut with white to pallid apex when mature, in young stages concolorous with the pileus with white apex, glabrous or slightly pruinate, glabrescent, equal, 30-63 X 0.5-1.5 mm, basal mycelium sometimes arising from a basal socle, fluffy-tomentose to strigose, white, strigosity partly especially at tips tending to become fulvous, usually forming gilvous-white rhizomorphs. Context white in pileus: odor none.

Spores 9.5-13.2 X 3.5-4.5 p, hyaline, smooth, tending to germinate on the lamellae. Hymenium: basidia 22-36 X 4.8-7.5 ,, clavate, hyaline, 4-spored; cystidia 25-50 X (4-)5-9 p, moderately crowded to rather numerous, cylindric to clavate or ventricose, differentiated clearly from the basidia by being opaque and/or rooting deeper than the basidia and basidioles, often subcapitate or appendiculate, hyaline, thin- to thick-walled: cheilocystidia: Siccus-type broom cells, much like the epicuticular broom cells, main body hyaline, 18-25 X 4.5-10.7 jp, often cylindric or cylindric clavate, beset with slightly diverging but strictly apical setulae, these hyaline to very pale golden hyaline, more rarely some cells with golden melleous setulae, short (5 4) to very long (20 pu), spinulose, acute or subacute, narrow (1.2-1.5 y diameter at base). In the hymenium sometimes hyphal pegs present, very scattered, hyaline, of interwoven filaments variable in size and shape. On edge occasional transition between cystidia and cheilocystidia (smooth cheilocystidia) observed. Hyphae in trama of pileus irregularly shaped arld somewhat interwoven, of different diameter and some very short, some pale stramineous, some with moderately thickened walls, but most filamentous, hyaline and thin-walled, all pseudoamyloid, with clamp connections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, main body 13.5-17 X 7.5-10 U, sometimes shorter, hyaline, mostly some golden ones intermixed, versiform, often vesiculose, setulae either all erect or some slightly spreading-divergent, strictly apical, either all golden hyaline to hyaline or more often

many golden melleous to even melleous brown ones on certain cells, spinulose, acute, subacute or obtuse, some strongly elongated, 3-22 X 1.2-1.6 p, Pruinosity of stipe con-

sisting of broom cells with often strongly reduced main body (main body up to 10 X 4 1) and setulae often spreading-subdivergent, often very long (up to 25 p long), subacute to acute, the whole broom cell mostly brownish.

On rotten fallen dicotyledonous branches, rotting leaf mold mixed with rotten wood, on petioles of dicotyledonous trees in tropical montane forest.

MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas. Rio Yariza, 16 Feb 1956, Singer B 1194 (LIL), 23 Feb 1956, Singer B 1369 (LIL) type; Carmen Pampa 2000 m alt, 26 Feb 1956, Singer B 1489 (LIL).


177. Marasmius hylaeae Singer sp nov Fig 77.

177a. Marasmius hylaeae var hylaeae TYPE. Singer B 7342, from Ecuador. Pileo ferrugineo-spadiceo, sulcato, 25-30 mm lato: lamellis intermittenter spadiceo-

ferrugineo-marginatis, siccando dilute grisellis, distantibus; stipite brunneo, 47 X 2 mm, mycelio basali albido. Sporis 10-11 X 4.3-5 1; cystidiis numerosis; cheilocystidiis sparsis ubi acies lamellarum colorata est. Ad ramum putrescentem in hylaea aequatoriana. Ty- pus in F conservatus est.

Pileus rusty spadiceous with lighter brown margin, dried deep red-brown (uniformly like "Java" but much deeper glabrous), sulcate over two thirds of the radius, about



Marasmius 231

25-30 mm broad. Lamellae not pure white but a very pale sordid grayish with intermittently narrowly discolorous (rusty spadiceous) edge when seen dried under a strong lens, broad, regularly intermixed with lamellulae (didymous, but lamellulae of variable length), not intervenose, distant (14 through-lamellae) rounded-subfree just reaching a brownish disc around the extreme apex of the stipe. Stipe brown, at first with pallid apex, glabrous but under a strong lens appearing slightly pruinose from the broom cells, equal, hollow, 47 X 2 mm; basal mycelium whitish. Context thin.

Spores 10-11 X 4.3-5 #, ellipsoid-oblong, smooth, hyaline, inamyloid. Hymenium: basidia 20-26 X 7-7.5 ,, 4-spored; cystidia rather numerous to very

numerous, 18-35 X 5.5-7 A, most opalescent, fusoid-subventricose or ampullaceous, some

appendiculate or mucronate, with mostly thickened wall, inamyloid; cheilocystidia scattered and present only where the lamellae-edges are discolored, like the epicuticular broom cells. Hyphae of the pileus hyaline, thin-walled, with clamp connections, pseudoamyloid. Cortical layers: epicutis of the pileus hymeniform, consisting of broom cells, main body 7-14 X 4.5-8.2 ,, mostly clavate, subhyaline to mostly brown, setulae 3.5-13 X 1-2(-3) #, spinulose, brown, some paler melleous, with firm walls, acute to obtuse: covering of the stipe consisting of rather numerous broom cells but many with strongly reduced main body and long, often spreading or forking setulae which are mostly acute and reach 25 ,uin length, all brown.

On rotting branch of dicotyledonous tree. MATERIAL STUDIED. ECUADOR. Napo, Sacha 4 (Sacha Norte), 10 May 1973, Singer

B 7342 (F).

177b.Marasmius hylaeae var hypsizygus Singer, var nov TYPE. Singer B 7340, from Ecuador. Differt carpophoris minoribus et habitatione in arboribus vivis. Typus in F. Pileus rusty spadiceous, dried between "Mandalay" and "Java" M&P, sulcate over

two thirds of the radius or more, and dried somewhat paler striped in the sulci, glabrous, about 10-15 mm broad. Lamellae not pure white but sordid whitish with discolorous (color of the pileus) edge, distant, not intervenose, adnexed or subfree from a widened apex of the stipe which shows a brownish ring when dried. Stipe brown, at first with whitish apex, appearing glabrous under a lens and brown dotted from the broom cells, equal or subequal, hollow, 25-30 X 1-1.5 mm; basal mycelium tomentose, whitish to

pale fulvous. Context whitish, inodorous. Spores (few seen) (7-)7.5-9.7(-11) X 3.5-4.3 p, ellipsoid or ellipsoid-oblong, smooth,

hyaline, inamyloid. Hymenium: basidia 23-25 X 6.5 ,, 4-spored; basidioles fusoid; cystidia 25-43 X 4.5-6.5 p, fusoid to clavate, often with constriction(s) at the apex which is frequently appendiculate or capitulate, with rather thin wall, sometimes somewhat opalescent, numerous, hyaline; cheilocystidia making the edges heteromorphous, dense, like the epicuticular broom cells. Hyphae of the pileus-trama hyaline, thin- to mostly firm- walled, with clamp connections, pseudoamyloid. Cortical layers: epicutis of the pileus hymeniform, consisting of broom cells, the main body hyaline to brownish, 10-23 X 6-11 L, thin-walled, versiform, setulae 5-10 X 1-2 1, melleous to succineous brown, subacute or obtusate, rodshaped to attenuate toward apex; covering of the stipe consisting of numerous broom cells, these often accumulated in bunches, main body often strongly reduced, 2-23 X 2-11 , hyaline to mostly brownish, thin-walled, versiform, setulae often strongly elongated and forked, subacute to obtusate, brown, 5-22 X 1.5-3 ,.

On living trees, lignicolous, solitary. MATERIAL STUDIED. ECUADOR. Napo, Sacha 4 (Norte), 10 May 1973, Singer B 7340

(F), type.




178. Marasmius praeandinus Singer, Sydowia 18: 353. 1965. Fig 78. TYPE. Singer T 3668, from Argentina. Pileus ochraceous brown to light cinnamon with deeper cinnamon to chestnut

center and sometimes dull ochraceous extreme margin, dried e g 12 F 9 on margin and "burnt sienna" (M&P) in center, glabrous, deeply sulcate up to the 1.5-2 mm wide smooth disc zone or over six seventh of the radius of the pileus, hat-shaped, with the margin tending to be recurved, papillate or subpapillate, 6-15 mm broad. Lamellae white to sordid- white-cream-pallid, medium broad to relatively broad, with entire edges which are concolorous with the sides or vaguely browned when dried, often ventricose, distant (10-12 through-lamellae and some lamellulae), not regularly inserted, sometimes somewhat anastomosing-confluent, but more frequently simple, not intervenose, very narrowly adnexed to free. Stipe chestnut, dried often more fulvous, with white apex, smooth, glabrous, somewhat shiny above, subequal or tapering downwards, 20-38 X 0.5-1 mmn basal mycelium whitish-pallid, dried tending to become fulvous-whitish to fulvous, strigose, connected with a surrounding pulverulent appressed (to the substratum) mycelium and white to fulvous rhizomorphic strands. Context of the pileus white or whitish, thin, inodorous.

Spores 8.5-11 X 2.3-4 y,, fusoid-oblong to fusoid, smooth, hyaline, inamyloid. Hy- menium: basidia 16-31 X 5.8-6.5 ,; basidioles fusoid; cystidia 20-42 X 4-8.3 ,u, rather to very numerous, generally deeper rooting than the basidia and with firm (0.2-0.4 pu thick) wall, few opalescent-refringent, not projecting or not much projecting beyond the apex of the basidia, versiform, mostly fusoid, ampullaceous, cylindrical and at the apex irregularly appendiculate or narrowed into a monilioid apex, hyaline; cheilocystidia like the epicuticular broom cells but hyaline to fulvous-hyaline or palest ochraceous. Hy- phae of the regular hymenophoral trama and the pileus-trama hyaline, filamentous, but in places strongly swollen, with clamp connections, pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform, consisting of broom cells which are ochraceous- melleous, setulae with firm wall, 4-11 X 1-1.8 , mostly subacute to acute, few obtuse.

On woody matter, e g bark of living dicotyledonous trees or on small fallen sticks in subtropical as well as tropical-montane forest.

MATERIAL STUDIED. COLOMBIA. Valle, Municipalidad Cali. Saladito, about 1900 m alt, 15 Apr 1968, Singer B 6163 (F). BOLIVIA. La Paz, Nor-Yungas, Rio Yariza, 23 Feb 1956, Singer B 1407 (LIL). ARGENTINA. Tucuman, San Javier, west slope of the Sierra de San Javier at 1000 m alt, 8 Mar 1951, Siuiger T 3668 (LIL) type.

179. Marasmius hinnuleus Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 297. 1869. Fig 79.

TYPE. Wright 155, from Cuba. Pileus ferrugineous-cinnamon to fulvous, dried "terrapin" M&P or "cinnamon

rufous" (Ridgway), glabrous, sulcate, convex and often somewhat truncate, 4-10 mm broad. Lamellae white with white, not discolorous edge, tending to brownish when drying, narrow to broad, not intervenose, distant, adnexed. Stipe dried umber to brown below with white apex at least at first, setose, glabrous, equal, 25-35 X 0.2 mm; basal mycelium well developed, strigose, fulvous. Context of pileus white, very thin.

Spores (7.7-)9.7-13.5 X 3.3-4.5 p, mostly 9.7-10.3 X + 3.8-4 ,, fusoid to oblong, smooth, hyaline, inamyloid. Hymenium: basidia 27-28 X 5-5.3 p, 4-spored; basidioles fusoid; cystidia numerous, 20-34 X 6.5-11 ,, ventricose to fusoid or subclavate, often apically appendiculate: cheilocystidia like the epicuticular broom cells but often scarce and entirely hyaline or few with bright melleous setulae. Hyphae of the pileus-trama hyaline, with clamp connections, pseudoamyloid. Cortical layer: epicutis of pileus hy-



Marasmius 233

meniform, consisting of broom cells, main body 11-22 X 4.3-7 y, clavate, hyaline; setulae 2-6.5 X 0.5-1 ,, mostly acute or obtuse narrowly conical, erect, hyaline, golden-hyaline, sometimes some brighter melleous ones on some cells.

On dead leaves in Cuba. Known hosts: Cecropia palmata, Hibiscus sp. MATERIAL STUDIED. CUBA. Wright 155 (FH) type; on "Paritium" (Hibiscus, Malvaceae)

(FH); Sebaruco, Soledad, 9 Jul 1941, White 846 (FH, BPI). In view of the rather pale color of most of the epicuticular broom cells the color

of the fresh fungus might be slightly paler than described above from dried material.

180. Marasmius pseudocorrugatus Singer, Sydowia 18: 340. 1965. TYPE. Singer B 2049, from Bolivia. Pileus a dull cinnamon (11 B 8 to 13 H 10 or 13 1 1) with or without an olive

tinge in the center, eventually often paler cinnamon, dried similarly colored but perhaps somewhat duller and deeper, the center at times tending to a more rusty ochraceous brown ("amber brown"), glabrous, radiately venose in the center or all over, but also sulcate over at least four fifths of the radius, occasionally with some white portions which disappear on drying, convex, soon with somewhat depressed center, 20-62 mm broad. Lamellae white or whitish, generally with a slight cinnamon tinge, especially when old or dry, eventually becoming pale cinnamon like the pileus, edges not distinctly discolorous fresh but sometimes developing a very narrow discolorous line (lens!) on drying, rather narrow to rather broad (3-7 mm broad) not intervenose in smaller or younger caps but sometimes intervenose in old, large specimens, medium distant, subclose to subdistant (usually about 14 through-lamellae) intermixed with a number of lamellulae, adnate to free. Stipe brown to very deep chestnut, glabrous, smooth, subequal, 40-75 X 2.2-2.7 mm; basal mycelium abundant, strigose, mainly white but often some cinnamon to fulvous portions present; whitish to pale fulvous rhizomorphs also often present. Context thin, inodorous.

Spores (7.5-)8-11.7 X 2-3.5 p, cylindrical to fusoid-ellipsoid, some with somewhat curved hilar end, thin-walled, hyaline, smooth. Hymenium: basidia 21-25 X 4.5-7.5 , 4-spored; cystidia usually moderately numerous to numerous, well differentiated, 35- 50 X 6.5-11 p, versiform, cylindric, ventricose etc, often constricted or appendiculate, otherwise broadly rounded, sometimes mucronate or capitulate, with moderately thickened walls, hyaline; cheilocystidia mainly hyaline, otherwise like epicuticular broom cells. Hyphae of trama of pileus interwoven, hyaline, thin- to somewhat thick-walled, some narrow, some broad (in older specimens up to 35 , in diameter), pseudoamyloid, with clamp connections. Cortical layer: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, 10-17 X 5-10 p, main body remarkably variable in its development, sometimes strongly reduced, hyaline to golden melleous or in other caps all golden melleous, in the first case mattled when seen in ammonia or KOH in scalp preparations, setulae usually concolorous with the main body, more rarely golden melleous on a hyaline main body, 2-13 X 1.2-2.8, (at base), sligihtly tapering upwards in most cells and acute to obtuse (or all obtuse) at the tips.

On all kinds of debris, often on dead wood or on bark of dicotyledonous trees in tropical and subtropical forest and plantations.

MATERIAL STUDIED. MEXICO. Sierra Costera, region of San Agustin Loxicha, 850 m alt, 27 Jun 1958, Guzman 1869 (ENCB); San Luis Potosi, near Tomasopo, 750 m alt, 11 Sep 1967, Sanchez 132 (ENCB). PERU. Loreta: Maynas, Rio Nanay, 23 Oct 1958, B. Lowy 423 (LIL) (?). BOLIVIA. Beni: Vaca Diez, Guayaramerin, 16 Mar 1956, Singer B 2049 (LIL) type, 14 Mar 1956, B 1985 (LIL); Pando: Manuripi, Conquista, 24 Mar 1956, Singer B 21 73 (LIL).





181. Marasmius spegazzinii Saccardo & Sydow, Sylloge Fungorum 14: 117. 1899.

Fig 80. Marasmius balansae Spegazzini, Revista Argent. Hist. Nat. 1: 102. 1891, non Patouillard

(1890). Marasmius floridanus Murrill, Bull. Torrey Club 67: 149. 1940.

TYPE. Balansa 4284, from Paraguay. Pileus orange to orange ferrugineous (10 H 9/10, "Marathon," 3 G 12, eventually

perhaps "terra cotta" to "ferruginous"), dried pinkish-ochraceous to fulvous-rubescent (e g 12 D 9) perhaps reaching "Arabian brown," glabrous, smooth and little striate in the extreme margin or vaguely sulculate all over for a long time but apparently eventually more strongly sulcate, conic to campanulate, than convex, often with flattened disc or margin or a small, low umbo, 10-33 mm broad. Lamellae white to pale cream, at first narrow, late r medium broad, in the largest specimens reaching 4.5 mm, intermixed, close, or subclose, not intervenose or with scattered rare anastomosing veins, simple, free, or rounded-adnexed. Stipe chestnut-umber, later often chestnut with pallid apex and blackish chestnut base, slightly shining and glabrous, smooth, equal or slightly thickened at base or both apex and base, 14-60 X 1-3 mm; basal mycelium tomentose to strigose and in the latter case somewhat radiating, white and remaining so or sometimes becoming slightly fulvous shaded on drying; whitish rhizomorphs inconspicuous but often formed. Context white, tough, inodorous, thin or moderately thin.

Spores 7.3-11 X 2.3-4.1 p, hyaline, smooth, inamyloid, fusoid to cylindric. Hy- menium: basidia 17-29 X 4.8-7.5 , 4-spored; cystidia rather numerous to very numerous, not or not much projecting in most collections and not very conspicuous in all collections but undoubtedly differentiated, 27.5-52 X 6.5-12.5 p, versiform ventricose, clavate or ampullaceous, sometimes with one or 2 constrictions, subcapitate, more opaque than basidia, somewhat indefinitely thick-walled, often some appendiculate more strongly differentiated and more numerous in age; cheilocystidia subhyaline, otherwise like epicuticular broom cells, but often intermixed with smooth, clavate cells (e g 9.5-5.5 p) and small, irregular bodies. Hyphae of trama of pileus eventually rather broad and somewhat thick-walled, in younger specimens more thin-walled ones present or all thin-walled, the thick-walled ones more strongly pseudoamyloid, all with clamp connections, their diameter reaching 21 A. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus type, 14-22(-42) X 3-10p, main body hand-shaped or Clavariella-shaped but also often clavate to vesiculose pedicellate or cylindric, hyaline to concolorous, thin- or some thick-walled, with setulae e g 6-24 A high, at the base 1-2.3(-2.7) , broad, rarely a few up to 40.5 A long, pale golden melleous or golden melleous, rarely a few hyaline, either the majority acute or subacute and a minority with rounded tips or vice versa.

On wood and bark of dicotyledonous trees and on woody sticks of various Dicotyledones in tropical and subtropical forest, rarely on dicotyledonous leaves fallen to the ground.

MATERIAL STUDIED. U.S.A. Florida: Alachua County, Murrill (NY) isotype; holotype of M. floridanus (FLAS) COSTA RICA. Ocampo 516 (F). PERU. Loreta: Rio Nanay, 28 Oct 1958, B. Lowy 447 (LIL). BOLIVIA. Beni: Vaca Diez: Guayaramerfn, 7 Mar 1956, Singer B 1636 (LIL), compared with type, B 1693 (LIL), 8 Mar 1956, B 1723 (LIL, cf observations below). PARAGUAY. Guarapl, fall 1884, Balansa 4284 (LSP, no. 2689),, holotype, (LSP, no. 2689 bis), syntype. ARGEN- TINA. Misiones: Bernardo Irigoyen, 24 Feb 1960, R. T. Guerrero 91 (LIL).

We have assumed that the larger specimens with long and distinctly deep sulcate pileus and somewhat deeper colors are more mature forms of older specimens of the same species since there does not seem to exist any other appreciable difference. The smooth specimens



Marasmius 235

agree perfectly with the type which is evidently not fully mature since there are only few spores in it. Yet, there are also, as often in Marasmius, individuals which are actively and abundantly sporulating without having reached the characters of a fully mature or old carpophore. Specimens characterized by larger size and sulcate pileus and studied by me are those from Peru: B 1723 from the Beni, the one from Argentina and the one from Florida.

Marasmius spegazzinii differs from most cystidiate species of this section in its small spores.

Mlarasmliuls floridalus Murrill was originally compared with M. berteroi by Murrill and therefore thought to be acystidiate inasmuch as an acystidiate species (M. leoninus) has actually been found in Florida but the type is clearly cystidiate and identical withM. spega- zzinlii.

Marasmzius virginianus Singer from the temperate zone of North America is similar but differs in habitat, somewhat smaller spores and slightly larger size and distinctly intervenose lamellae, especially near the periphery of the pileus. See Singer (1958a, p 112).

The following species, also very close to M. spegazzinii and M. virginianus differs in habitat, somewhat larger spores, distinctly though not extensively intervenose and at the edges discolorous lamellae.


182. Marasmius aztecus Singer, sp nov TYPE.Guznlun 6148, from Mexico. Pileo laete ochraceo-brunneo vel laetissime ferrugineo-aurantiaco vel aurantiaco, levi

vel breviter sulcato dein profunde sulcato, centro levi vel ruguloso, 17-30 mm lato; lamellis prope marginem pilei frequenter discoloribus ad aciem, ex integro per acies anguste discoloribus in siccis et demum interdum aurantiac'o-fimbriatis; stipite 40-160 X 1.5-2.5 mm. Sporis 8-13 X 3-4 ,, cystidiis paullum vel bene evolutis. Ad quisquilias in silvis montanis mexicanis. Typus in Herbario Instituti Polytechnici Mexicani conservatus est.

Pileus a bright ochraceous brown to brightest rusty orange or orange (dried varying from "Sudan br." to 3 A 12, if orange mostly with some copper br. but rather inconspicuous stripes, if "Sudan br." often but not always with much paler margin or with much darker ("bark") zones or areas, all M&P), varying from smooth to relatively short sulcate in young to deeply sulcate in large old specimens, glabrous with smooth to more or less rugulose center, 17-30 mm broad, rarely broader. Lamellae white or cream whitish near the stipe but often discolorous near the margin of the pileus in young, entirely but narrowly (lens!) discolorous edge in dried old specimens, the edges in older specimens often but not constantly conspicuously orange fimbriate, rather narrow but mostly medium broad to rather broad in the marginal (outer) third, didymous- to tridymous, young not, mature distinctly tough, mostly not extensively intervenose, close or subclose to medium close, with about 14-15(-18) through-lamellae, but not distant because of the many lamellulae, adnexed to subfree. Stipe chestnut, later dark chestnut with at first white apex and sometimes becoming almost black at base, smooth and glabrous, hollow, 40-106 X (0.8-)1.5-2. 5 mm; basal mycelium abundant, strigose, whitish to pale fulvous. Context white in pileus.

Spores (7.7-)8-13 X (2.5-)3-4,u, fusoid, smooth, hyaline, inamyloid. Hymenium: basidia 19-28 X 5-7.5 A, 4-spored, occasionally some 2-spored; cystidia moderately numerous to numerous, poorly to well differentiated but always constantly present on the sides of the lamellae, 19-40 X 3.5-1 1 u, versiform, often ventricose to clavate, but also ampullaceous or cylindrical, often with attenuated but not acute apex, mostly slightly deeper




rooting than the basidioles, best seen in Melzer's reagent but inarnyloid, somewhat opaque with indistinct inner outline of the often rather thick wall (up to 1 ,), not strongly projecting beyond the sterigma-level, in some collections some cystidia appendiculate, all hyaline and without differentiated contents bodies; cheilocystidia like the epicuticular broom cells, with hyaline main body and hyaline to mostly light golden orange setulae, in addttion where the edges are fimbriate, the excrescencies consisting of thick-walled hyphae originating in the hymenophoral trama, these hyphae filamentous, densely fasciculate, the terminal cell often transformed into a broom cell but with very small or no main body and pale orange setulae. Hyphae of the pileus-trama hyaline, with clamp connections, with thin, firm or thick wall (1 I), strongly pseudoamyloid; hymenophoral trama regular, hyaline, consisting of hyphae which are either all thin- to firm-walled or with some thick-walled ones intermixed, not gelatiiized, also pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform, consisting of broom cells, main body 7-15 X 4-15 p, versiform, with a wall 0.2- 0.4 p thick and lyaline, fewer with at least the upper portion concolorous with the pigmented setulae; setulae 2-7.5(-13) X 0.5-1.5(-2.5)t if hyaline, but up to 22 X 3 Aif pigment (pale golden to golden mellcous) attenuated to an obtuse, more rarely subacute tip, in very few cells occasionally absent; scalp preparations usually mottled because of the alternating groups of pigmented and less or not pigmented broom cells.

On all kinds of debris (small pieces of wood rotting in the leafmold) or on rotting trunks in the oak-pine woods and oak woods of the iionitane forests of Central Mexico.

MATERIAI STUDIED. MEXICO. MCexico, near Pefla Blanca, SE of Valle de Bravo, 2100 in alt, 26 Sep 1967, (;Guzma 6148 (EINC'3) type, (1I), isotypcs; Morelos. 5 klm W of Tepoztlan, 1900 m alt, 13 Jun 1967, A. l.opcz (;onzlatez 119 (INC1B). I. Piedras 108 (I:N('CB).

This species should not be confused witih Marasmrius pscudocorrngatus Singer (no. 180) which differs in the color of the pileus and slightly smaller spores and occurs only in forests below the montane zone and under 1000 m altitude.

Marasmius sect Sicci subsect Siccini ser Actinopodus Singer, ser nov Elementis setoideis praesentibus. Setoid bodies present on pileus or stipe or lamellae, or else on several of these sur-

faces; cystidia present or absent on the lamellae. TYPI SPIECIES. Marasnius actimlopus Montagie.

Key to the Species of Series Actilopocles

1. Setae or setoid cystidia or long hairs absent from the cortical layer of the pileus and tlic lamellae as well as from the stipe where the covering consists of ordinary broom cells or small setulac of these or of thin-walled, hyaline, cystidiforml elements, not longer tihan 50 p and with obtuse apex. 2. Stipe covered merely by broom cells, tlhe main body of these often obliterated and

the setulae normally short, not strongly elongated and setose: stipe usually glabrescent or only minutely pruinate-subpubescent when fresh and young (if cystidia are present on the sides of tile lamellae and the setulac of tlhe stipe extraordinarily elongated, see "7" and "15" below), see series Leonini.

2. Elements of stipe covering not in form of broom cells; lamellae crowded to subclose. 3. Pileus white when fresh. 1 84. lM. personatus. 3. Pileus orange to red brown wlien fresh.

4. Lamellae strongly anastomosing and partly or entirely pore-like or at least very strongly and consistently intervenosc; cystidia indistinct or absent. 150. M. cladophyllus.

4. Lamellae only very slightly or not intervenose but may be at times partly forked or crisp. 5. Cystidia indistinct or absent; on conifers (otherwise compare series A if

hairs very inconspicuous). 149. Al. araucariae. 5. Cystidia present; not on conifers. 183. AM. chrvsohlepharis.



Marasmius 237

1. Setae or setoid cystidia or long (over 50) hairs present on pileus and/or lamellae and/or stipe. 6. Stipe covered by long (over 50, at least many of them) hairs and sometimes these

also present on the pileus very rarely the lamellae, but these hairs, at least in the middle and the apex of the stipe, not or not strongly pigmented i e hyaline or light stramineous, with thin to thick wall. 7. Lamellae subdistant or moderately distant.

8. Spores more than 9.5p long (see "10" below). 8. Spores up to 9.5 p long; center of pileus often rugose or venose.

9. Pileus orange or orange-tawny, 6-10 mm broad; setulae of the epicuticular broom cells mostly yellow, orange yellow or melleous, some hyaline; spores 7.5-8.5 X 3-3.5p; on leaves and small twigs. 185. M. rubroflavus.

9. Pileus deeper rufescent-purple. broader; setulae of the epicuticular broom cells brown; spores 8.3-9.3 X 3.8-5 p on wood. 186. Al. glaucopus.

7. Lamellae crowded to subclose. 10. Spores reaching larger size than 10.5pu; pileus 420 mm.

11. Pileus white or orange (see "3" above). 11. Pileus not so colored.

12. Pileus ochraceous to ochraceous brown; many or all of hairs of stipe obtuse.

13. Most of the hairs of the stipe with obtuse tip, few subacute; spores 11-15.8 X 3.5-4.2p; transition zone between basal mycelium and stipe covering tending to brown; Florida, West Indies and Mexico.

187. AI. bahamensis. 13. Most of the hairs of the stipe acute, subacute, fewer obtuse; spores

11-14 X 3-4 p; hairs of stipe and basal mycelium white to stramineous, the latter tending to fulvous when dried; like South America and Africa. 188. A. actinopus.

12. Pileus chestnut, reddish-ferruginous or purple red; most or all hairs of the stipe acute. 189. M. atrorubens.

10. Spores not longer than 10.5 p; pileus decidedly over 20 mm in diameter. 14. Pileus white when fresh; epicuticular elements partly with setulae partly

like the broom cells but without setulae; cheilocystidia like the epicuticular broom cells or cystidia pseudoamyloid. 190. M. pseudoniveuts.

14. Pileus cinnamon when fresh; epicutis composed of broom cells only; cheilocystidia different from the epicuticular broom cells. 191. M. heterocheilus.

6. Setae or setoid cystidia present in the epicutis of the pileus and/or on the lamellae and/or on the stipe, in the latter case at least many of them are pigmented and thick-walled and acute, the setoid cystidia of pileus and lamellae likewise pigmented, rarely hyaline; setoid bodies sometimes forming transitions to broom cells but then main body often strongly reduced and setulae very large. 15. Spores 12-21 X 4-5 p; stipe pubescent; setae of stipe like large setulae of broom

cells. 3-5.5 , broad at base. 192. M. spiculosus. 15. Spores smaller or stipe glabrous.

1 6. Lamellae subdistant or distant. 17. Most setae or setoid cystidia partly or entirely brown (see '21" below). 17. Setoid bodies hyaline. 194. M. flanmmans.

16. Lamellae close or subclose. 18. Setoid cystidia absent on the lamellae. 195. M. echinatulus. 18. Setoid cystidia present on the sides of the lamellae.

19. Setoid cystidia or setae brown, at least many of them. 20. Smooth broom-like bodies present in epicutis, in addition to

normal broom cells and setae. 196. M. venezuelanus. 20. Smooth broom-like bodies absent in epidermis, epicuticular cells

either normal broom cells with setulae or setae of the epicutis in form of setulae-bearing broom cells.

21. Pileus 25-67 mm broad, cream buff to cream gray. 197. M. jalapensis. 21. Pileus 10-26(-34) mm broad, deeper colored. 193. 1M cohaerens.

19. Setoid cystidia hyaline (see "14" above).

183. Marasmius chrysoblepharis Singer, sp nov Fig 81. TYPE. Singer M 8201, from Mexico. Pileo aurantiaco, siccando ferruginascente, glabro, nonnihil sulcato vel sublevi, con-




ico dein convexo, 8-14 mm lato; lamellis albis vel aurantiaco-pallidis, ad aciem aurantiacis ve] ferrugineo-aurantiacis praesertim in siccus, confertis vel paene confertissimis; stipite ferrugineo-castaneo, dein obscuriore, pubescente vel velutino, apice albo vel aurantiaco-pallido, 35-43 X 0.5-0.8 mm; mycelio baseli abundante, albido. Sporis 9.3-13 X 3.3-3.7 ,; cystidiis numerosis; stipite pilis cystidiiformibus numerosis 9-80 X 4.11 /j, pleruinque cy- lindraceis ornato. Ad folia arborum dicotyledonearum in Mexico. Typus in F conservatus est.

Pileus orange, in age deep orange yellow when fresh, dried "rust, sorolla br" (M&P), glabrous, smooth, then shallowly sulcate or subsmooth, at first conical, then convex, eventually convex-applanate with slightly depressed center, obtuse, more rarely somewhat umbonate, 8-14 mm broad. Lamellae white to orange-pallid, the edges discolorous (pale orange to rusty orange) especially after drying, rather narrow, close to almost crowded, rounded-adnexed, adnexed or subfree. Stipe rusty-chestnut, then deep chestnut with the base eventually almost black, dried still deep brown, the apex white or orange-pallid, entirely pubescent or velutinous, equal or very slightly tapering upwards, 35-43 X 0.5-0.8 mm; basal mycelium abundant, whitish, often tending to fulvescent. Context of pileus white, thin, inodorous.

Spores 9.3-13 X 3.3-3.7 y, fusoid but attenuated in upper third or broadened in lower, acute or subacute above, smooth, hyaline, inamyloid. Hymenium: basidia 17-21 X 5.5-7.5 p, 4-spored; cystidia numerous, 15-22.5 X 6-8.5 u, fusoid or ventricose, more

rarely clavate, some with small apical appendage which is often oblique, sometimes constricted in the middle, opaque and the inner outline of the mostly thick wall not clearly visible in most cases, hyaline, easily overlooked because they are little larger than the basidioles, best seen in cresyl blue mounts; cheilocystidia like the epicuticular broom cells, making the edge hetermorphous. Hyphae of the pileus-trama hyaline, thin- to moderately tlhick-walled, with clamp connections, pseudoamyloid. Hyphae of the stipe yellow, sometimes tending to olive in the rind layer, parallel with each other, with clamp connections. Cortical layers: epicutis of the pileus hymeniform, consisting of broom cells, main

body 9-16 X 5-8 u, versiform, thin-walled, hyaline or orange-golden; setulae 3-5.5 X 0.5-1.2 .u, apical-erect, some golden, some golden-hyaline and scalp preparations therefore appearing mottled; covering of the stipe consisting of cystidiform hairs, these numerous or even

dense, 9-80 X 4-11 , most cylindric with rounded-obtuse apex, some more ampullaceous because of a swelling in the basal portion and also obtuse, few with apical or lateral ob-

lique appendage, often flexuous, often with a clamped septum near the middle, subhyaline or yellowish, with firm or slightly thickened wall (0.3-0.9 p in diameter), inamyloid.

On leaves of dicotyledonous trees in Mexico. Known host: Platainus occidentalis. MATERiAL STUDIED. MEXICO. Veracruz, Ttilapa, banks of Rio Aserradero, 24 Jun 1969,

Singer M 8201 (F), type; Chiapas, between Finca Sospiro and El Pozo, 4 Aug 1969, Singer M 8960 (F).

184. Marasmius personatus Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 297. 1869. TYPE. Wright 10, from Cuba. Pileus white, drying yellow, closely striate towards the margin, otherwise smooth,

convex, eventually depressed, 6-10 mm broad. Lamellae cream, edge concolorous, crowded, of three lengths, adnexed. Stipe fulvous or dark brown, minutely pubescent (glabrous according to original diagnosis), with white apex, about 40-45 X 0.8-1 mm; basal mycelium strigose, white, associated with a patch of white mycelium over the substratum.

Spores 10-14 X 2.5-3.7 u. Hymenium: cystidia 20 X 6 p, fusiform, only their tips protruding above the basidia (Dennis); cheilocystidia like epicuticular broom cells. Hyphae



Marasmius 239

of the hymenophoral trama strongly pseudoamyloid. Cortical layers: epicutis of pileus formed of hyaline broom cells 10-12 p across with short rounded hyaline to golden-yellow setulae; covering of stipe made up of hyaline hairs which are cylindrical, obtuse, often thickened at base, about 40 X 4 u.

On sticks. MATERIAL STUDIED. CUBA. Wright 10 (FH), type. VENEZUELA. D.F., Caracas Botanic

Garden, 6 Sep 1958, Dennis 1022 (K).

ILLUSTRATION: Dennis (1961), fig 49.

Stirps Actinopus (no. 185-189)

This group is characterized by the presence of long hairs ( > 50O ) on the stipe but these not distinctly setoid nor are there usually setoid cystidia on the pileus or lamellae but non-setal dermatocystidia or long hairs present on the pileus in some species.

185. Marasmius rubroflavus (Theissen) Singer, Lilloa 22: 326. 1951. Fig 82. Mlarasmius nummularius var nrbroflavus Theissen, Broteria 8: 57. 1909.

TYPE. Theissen, from Brazil (holotypus not found). Pileus orange-tawny, sometimes with purple red center, testaceous or orange when

dried, towards margin generally slightly paler than in the often rugose center, sulcate in the marginal half, pubescent from hyaline hairs but glabrescent, campanulate-convex, then convex, slightly umbonate and later sometimes flattened or depressed around the umbo, eventually subapplanate, 6-10 mm broad. Lamellae white, the edges white, but towards the stipe often slightly pale orange (under a lens), some forked, not intervenose, rarely eventually slightly intervenose, didymous, subdistant, somewhat ventricose, adnexed or free. Stipe orange-tawny or light fulvous below, white or whitish above. eventually becoming chestnut at the base, pubescent, hollow, equal, 13-50 X 0.5-0.7 mm; basal mycelium strigose-tomentose or discoid-membranous, sometimes attached to the white rhizomorphs which are equally pubescent as the stipe, white, on drying ochraceous-fulvous. Context in the pileus white, very thin, inodorous.

Spores 7.5-8.5 X 3-3.3 p, oblong, smooth, hyaline, inamyloid. Hymenium: basidia

4-spored; cystidia none; cheilocystidia like epicuticular broom cells. Hyphae of the pileus- trama hyaline, with clamp connections, with thin walls, pseudoamyloid. Hyphae of the

stipe-trama hyaline to yellow or (in base) often brownish, parallel with each other, with

clamp connections, strongly pseudoamyloid. Cortical layers: epicutis of the pileus hymeniform, consisting of broom cells and hairs, the former with a main body e g 11 X 7.5 pI, hyaline, setulae hyaline or more often yellowish-orange or melleous, acute, medium sized; hairs like those of the stipe; covering of the stipe consisting of moderately dense, short to more often long hairs, 20-150 X 3 p, filamentous but often broadened at the base (to 7 p, across and rarely broadened at the apex to 5.5 p) with constantly obtuse- rounded tip, with thin, more rarely somewhat thickened wall at least in the lower portion, reaching 0.5-1 , in diameter, hyaline or yellowish-hyaline, simple, rarely forked.

On small sticks and leaves of dicotyledonous trees, gregarious, in tropical and subtropical forest.

MATERIAL STUDIED. MEXICO. Veracruz, 7 km S of Montepio, Estacion Biol6gica de los Tuxtlas, 20 Jun 1969, Singer M 8084 (F). BRAZIL: Rio Grande do Sul, Theissen leg. & det. (FH), authentic.




186. Marasmius glaucopus (Patouillard) Saccardo & D. Saccardo, Sylloge Fungorum 17: 40. 1905.

Anidrosaceus glaucopus Patouillard in Duss, Enumeration Champignons, p 43. 1903. T-YPE. Duss 594, from Guadeloupe. Pileus deep purplish rufous, dried spadiceous, appearing subvelutinous, center rugose-

venose, plicate-sulcate, with incurved margin, campanulate-convex, 20 mm broad. Lamel- lae dark purple with glaucous pruina, intermixed, rather distant, broad, not intervenose, attenuate. Stipe deep reddish fulvous, strongly pruinose all over, equal, hollow, 30 X 3 mm; basal mycelium present.

Spores 8.3-9.3 X 3.8-5 1, ellipsoid, sometimes tending to be tilda-shaped when seen in profile, hyaline, smooth, inamyloid. Hymenium: Basidia 27-30 X 6.5 1, 4-spored; cystidia about 36 X 8.7 ,, thick-walled and somewhat opaque, fusoid ventricose with obtuse apex. Hyphae of trama of pileus pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, these about 25 j long (with setulae) base about 4.5 , across, hyaline, setulae often spreading to about 19 p but strictly apical, 6.5-10.3 j,long, spinulose, mostly simple but some forked, brown.

On rotten wood of Chtrysophyllunm. MATERIAL STUDIED. GUADELOUPE: Pointe-Noire, interior forests, 1902, Duss (FH)

type. According to Patouillard the pruina of the apex of the stipe is cinereous. The

anatomy of the pruina has not been established. ILLUSTRATION. Singer (1958a) fig 35.

187. Marasmius bahamensis Murrill, North American Flora 9: 265. 1915. Fig 83. TYPE. Britton 611, from the Bahamas Islands, New Providence. Pileus ochraceous brown or ochraceous yellow, rarely with a slight orangy hue

("umber brown" with "buckthorn brown" margin, Ridgway) when fresh and wet, drying to "Clay Color" (Ridgway) or "Yucatan" (M&P) with "Pinkish Buff' (Ridgway or "topaz' (M&P) margin, dried and rewetted becoming brown, smooth or slightly sulculate, macroscopically glabrous, convex, eventually more applanate, 7-11(-15) mm broad. Lamellae cream to pale yellow-buff ("Light Ochraceous Buff," Ridgway) close, often ventricose, narrow to medium broad, not intervenose, rounded-free or narrowly adnexed. Stipe brown with the apex concolorous with the lamellae and the base pale melleous to ocher brown, hirsute-tomentose on blackish ground, merely pruinate-pubescent above, smooth, equal, sometimes with broadened base, 22-50 X 0.5-1 mm; mycelium white but passing into the brown tomentum of the lower portion of the stipe, but even the white basal mycelium proper tending to become fulvous in parts on drying. Context: pallid to brownish more pallid in the pileus, more brownish in the stipe, rather tough, without distinctive odor.

Spores (11-)14-15(.15.8) X (3.5-)3.7-4(-4.2)jj, fusoid in frontal view, often somewhat club-shaped sometimes slightly curved and like Fusariunm spores, smooth, hyaline, inamyloid. Hymenium: basidia 17-25 X 6-8.2 u, 4-spored; basidioles fusoid, sometimes remaining sterile and then often with capitate apex (cystidioles); true cystidia none. Hyphae hyaline in the pileus-trama, with clamp connections, elongated but relatively broad, with thin (0.1-0.3 , thick) wall, pseudoamyloid. Cortical layers: epicutis of the pileus hymeniform, of broom cells, these with a main body 8-20 X 5-10 f,, versiform but often subisodiametric, hyaline to golden hyaline, setulae 2-5 X 1-1.5 u, these and often also the uppermost portion of the main body golden melleous; hairs of the stipe 15-120 X 4-9(-10),i, with often tapering but at the tip rounded apex, rarely subacute



Marasmius 241

unicellular, with thin to thick wall, rather stiff, hyaline but stramineous towards the base.

On fallen leaves of Quercus, Nectandra, Ficus and other dicotyledonous trees in South Florida and the West Indies, west to Mexico.

MATERIAL STUDIED. U.S.A. Florida: Dade County, Matheson Hammock, 9 Dec 1942, Singer F 1597 (FH). BAHAMAS. New Providence, at Lake Cunningham, 8 Sep 1904, E. G. Bri- tton 611 (NY), type. MEXICO. Tabasco, Villahermosa, Parque de la.Venta, 1 Aug 1969, Singer M 8784 (F).

The species appears to be extremely close to Marasmius actinopus Montagne (no. 188) and might yet be relegated to varietal status. The main differences are the smaller spores and the often acute hairs of the following species.

188. Marasmius actinopus Montagne, Ann. Sci. Nat. IV. 1: 112. 1854. TYPE. Leprieur, from French Guyana. Pileus dull ochraceous yellow to dull olive brown ("Samovar" or Buffalo" or

"Antique bronze" M&P) or pallid in the center with clay colored, bronze brown or olive- ochraceous margin ("clay," "bronze," "desert," "centennial brown") glabrous, sulculate over at least half of the radius when mature, otherwise smooth, hemispherical, later convex, 14-16 mm broad. Lamellae yellowish white, adnate or adnexed, close or subclose. Stipe melleous to reddish brown, hirsute, 25-50 X 0.3-1 mm; basal mycelium pallid to fulvous, strigose. Context thin, white, inodorous.

Spores 11-14 X 3-4 ,, clavate-subfusoid, smooth, hyaline. Hymenium: basidia 15. 5 X 5.5 u, 4-spored; cystidia 15-36 X 5-13 u, not always present but frequently, many sterile bodies basidiole-like (cystidioles!) but some slightly opaque and somewhat thicker- walled; cheilocystidia like epicuticular bioom cells. Hyphae: subhymenium consisting of very short, irregular, small cells, appearing subcellular. Trama of pileus hyaline, of thin- walled and a few slightly thick-walled hyphae, some of them rather voluminous, rapidly and strongly pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, main body hyaline, 10-17 X 4.5-8 ,, cylindric or clavate, setulae often spreading and subdivergent but strictly apical, 4-11 X 1-1.5 (at base) p, in some cells hyaline in others golden melleous therefore surface appearing mottled in scalp preparation; surface layer of stipe consisting of dermatocystidioid hairs, 20-130 X 5-17 A,, with often enlarged base, often opaque, thin- to thick-walled (wall up to 1.3 p

diameter) subulate to cylindric, obtuse or acute at tip, subcapitate once or twice at times.

On dead woody twigs, leaf petioles and leaves of Dicotyledones in tropical rain forest and tropical-montane forest, usually gregarious but not strictly cespitose. From northeastern South America to Bolivia.

MATERIAL STUDIED. FRENCH GUYANA. Leprieur (K) type (part of type conserved in the Berkeley Herbarium). VENEZUELA. Caracas: Rfo Chacaito, 18 Nov 1949, Dennis 397 (K); Parque Nacional El Avila, 24 Jul 1972, Dumont et al VE 5867 (NY). BOLIVIA. La Paz: Nor-Yun- gas, Charobamba, 13 Feb 1956, Singer B 1121 (LIL). 20 km NE or Yambao, 470 m, Louis 15216, 15241 (BR); Lemfu, Vanderyst 699 (part) (BR).


189. Marasmius atrorubens (Berkeley) Berkeley, Hook. Jour. Bot. 8: 137. 1856. Agaricus atrorubens Berkeley, Lond. Jour. Bot. 1: 138. 1842. Marasmius castaneus Montagne, Ann. Sci. Nat. IV. 1: 109. 1854. Marasmius portoricensis Murrill, North American Flora 9: 262. 1915 (according to Dennis). Marasmius jamaicensis Murrill, North American Flora 9: 261. 1915.




189a. Marasmius atrorubens var cystidifer Singer, var nov TYPE. Singer B 1700, from Bolivia. Differt praesentia cystidiorum numerosorum. Typus varietatis in LIL conservatur. Pileus red ("autumn glory," "pomegranate purple," "garnet r" M&P) becoming

gradually reddish ferruginous or orange ferruginous ("terra cotta," "mosque," 5 E?G 12,

"ferruginous," "Indian red") especially on margin or often chestnut to deep reddish chestnut and remaining so in the herbarium ("Coromandel") the reddish tones disappearing through drying out "in situ" but if dried rapidly and immediately while still wet and red, the red maintains itself in the herbarium well preserved, dried e g between "Liberia" and "Carbuncle" and those parts that are "terra cotta" fresh, becoming "Alamo," glabrous at first smooth, then striate over one sixth to one half of the radius and eventu-

ally long sulcate in many specimens, sometimes papillate but more often obtuse, convex, more rarely shallowly conic, 6-13 mm broad. Lamellae yellowish-white to more or less

yellowish or buffish ("Leghorn" to "Nankeen", 9 C 2), then often reaching orange-cream, with the edges concolorous with pileus or with sides of the lamellae, narrow (0.8-1.5 mm

broad), not intervenose, subclose to close or mostly almost crowded, rounded-adnexed or more often adnate. Stipe chestnut brown to light cinnamon (12 B 8, "wild honey," "russet brown," "toast " to "cinnamon" or "cinnamon brown") but apex varying between concolorous with lamellae and "ferruginous," dried tan to lighter straw-yellow and even fresh often appearing paler than ground color because of the dense hirsute pilosity all over which is hyaline to pallid stramineous, apex at first sometimes pure white

(if lamellae yellowish white), equal or slightly and gradually tapering upwards, 22-45 X 0.3-1.2 mm; basal mycelium abundant, strigose, yellow to brown ("cinnamon brown," 12 F 7, "russet brown" M&P) more rarely brownish pallid or light buff to stramineous, not pure white, sometimes green from algal populations. Context inodorous.

Spores (12-)14-20.5 X 3-5 1u, hyaline, fusoid or clavate-fusoid, inner side usually more or less applanate, thin-walled, overmature spores frequently thick-walled, smooth, inamyloid. Hymenium: basidia 18-22 X 4.8 p, 4-spored; cystidioles 22-27 X 6-1 Ou, ventricose to fusoid, with conical subacute apex or mucronate, with very finely granular, hyaline incrustation and/or contents in many of them, hyaline to stramineous-hyaline, thick-walled and refractive, numerous; cheilocystidia like epicuticular broom cells but

frequently less deeply colored because of more numerous hyaline to melleous setulae, intermixed with basidia and cystidia. Hyphae of trama of pileus and hymenophoral trama hyaline, of the stipe stramineous to pale melleous, all pseudoamyloid and with

clamp connections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type, 16-29 X 4-11 u, main body about 9-22 u high, ventricose-vesiculose to cylindric-clavate, also strongly club-shaped, in some specimens all concolorous with setulae or at least melleous, in others all hyaline or hyaline and melleous mixed, depending on density of pigment and age (older specimens more pigmented) and color (deep red in herbarium specimens more pigmented) setulae 4-7 X 1.2-1.4 (at base) u, rod- shaped or spinulose (in variable proportion) in accumulation appearing melleous brown to deep red brown, individually varying in some specimens from hyaline to melleous in others from light golden melleous to melleous brownish and in these intensities regularly distributed so that the surface appears mottled-marbled in scalp preparations, obtuse to acute. Pilosity of stipe consisting of erect, stiff hairs or setae, these numerous, but not forming a palisade, 23-230 X 6.5-14 u, pale melleous to melleous or melleous stramineous, generally unicellular, rarely with one or two secondary septa, simple, subulate to lanceolate, needle-sharp acute to subacute, smooth, thick-walled below or



Marasmius 243

everywhere (wall 1-1.7 ,), not or very weakly pseudoamyloid; at the base of these three is a second type of element present; short and broad dermatocystidia, these 20-

70(-100) X 8.5-13.5 j, hyaline to stramineous, smooth, with broadly rounded tip, varying from scattered to numerous and from vesiculose to ventricose.

On dead dicotyledonous leaves in tropical rain forest, generally solitary, more rarely in small, not crowded groups. Known host: Cecropia sp.

MATERIAL STUDIED. BOLIVIA. Beni, Vaca Diez, Ivon, 3 Apr 1956, Singer B 2478 (LIL); Guayaramerfll, 8 Mar 1956, Singer B 1700 (LIL) type B 1748 (LIL), 9 Mar 1956, B 1780 (LIL), B 1824 (LIL), 11 Mar 1956, B 1828 (LIL), 14 Mar 1956, B 1980 (LIL), B 1980a (LIL). (Some of these have not been checked for the type of cystidioles and may belong to var atrorubens; B 1700 has those as described above.)

189b. Marasmius atrorubens var atrorubens This agrees in most characters with var cystidifer but Dennis (1951b) describes

the material as having no pleurocystidia.8 Acystidiate forms do indeed occur and are cited below. None of these was ever found to have red pileus when fresh but it is russet-ferruginous, orange-ferruginous when fresh and a deep red brown when dried.

MATERIAL STUDIED. SURINAM. Hostmann 297 (K, Hooker Herbarium, PC), holotvpe, (K. Cooke Herbarium) syntype . GUADELOUPE. Duss 1765 (FH) as AM. castaneus Mont. l .) VENEZUELA. Miranda, Rfo Cupira near El Bachiller, 5 Jul 1972, Dumont et al VE 3938 (NY). ECUADOR. Shushufindi 15 May 1973, Singer B 7465 (F), B 7461 (F). 13 May 1973, B 7400 (F).

Dennis (1951b) also mentions a cystidiate form which would correspond to our var cystidifer but is said to have also brown setae on the surface of the pileus. It occurs in Ceylon.

The following variety has no cystidia but differs from the type variety in the presence of hairs on the surface of the pileus:

189c. Marasmius atrorubens var dumontii Singer, var nov TYPL. Dumont et al VE 4877, from Venezuela. Differt absentia cystidiorum et praesentia pilorum longorum in epicute. Typus in

NY. Pileus dried deep chestnut-ferruginous to deep badious-castaneous, unicolorous, long

and deep sulcate, glabrous (macroscopically), subconical to convex, 4-5 mm broad. Lam- ellae white with white edges, sometimes in a zone 1 mm wide around the stipe with brown edges, close or subclose, (e g 16 through-lamellae), narrow, not intervenose, adnexed. Stipe stramineous, villous-hirsute, all over from pallid or palest stramineous hairs, most densely hairy towards the apex, equal, 16-18 X 0.2 mm; basal mycelium pale tawny to fulvous when dried, abundant. Context thin.

Spores 12.5-20 X 3.3-4.5 , narrowly fusoid to cylindric-oblong, some above slightly curved, smooth, hyaline, mamyloid. Hymenium: basidia 22 X 8 u, 4-spored; basidioles fusoid; cystidia none; cheilocystidia like the epicuticular broom cells, but most of them

8I am not certain about the presence or absence of true cystidia in the type specimens. However, my collections seem to have cystidioles which are very difficult to distinguish from basidioles and are thin-walled.

9Mixed collection (one specimen different from M. atrorubens).

?Mixed collection (?) "Tantot blanc tantot rouges," said to occur on "sarments, sur tiges de Panicum maximum et brindilles de paille", what is left in the envelope is on dicotyledonous leaves; spores not recuperated.




either entirely hyaline or with very pale stramineous setulae. Hyphae of the trama of the pileus hyaline, most filamentous but some (especially near the epicutis) swollen and short- celled, thin- to firm-walled but not distinctly thick-walled, with clamp connections, pseudoamyloid. Cortical layers: epicutis of the pileus hymeniform, consisting of broom cells, these with a main body 8-20 X 6-9 A, cylindrical to clavate or subvesiculose, stramineous or hyaline or melleous-ochraceous to (rarely) spadiceous, with firm, more rarely rather thick wall; setulae 3-8 ilong, spadiceous or golden yellow, few hyaline or stramineous, erect to oblique-spreading, rodshaped to narrowly conical, acute or obtuse. Aside from these, particularly in the marginal zone of the pileus but not constantly so, some thick- walled hairs present like those of the surface of the pileus but reaching up to 450 p in

length, all inamyloid, entire, wall 1-2 p thick, hyaline to stramineous, about 6-7 p broad and tapering to an acute tip; covering of the stipe consisting of numerous hairs, at least 27-170 p long, 6-11 A broad, acute, the shorter ones often obtuse, thick-walled below (wall there 1-1.3 A thick), most entire, few at base or at apex bifurcate, stramineous or (fewer) hyaline, inamyloid.

On woody branches and dead dicotyledonous leaves, Venezuela. MATERIAL STUDIED. VENEZUELA. Sucre, Trail from Los Pocitos to Rio Grande Arriba,

13 Jul 1972, Dumont et al VE 4877 (NY) type.

190. Marasmius pseudoniveus Singer, Sydowia 18: 340. 1965.

190a. Marasmius pseudoniveus var pseudoniveus TYPE. Singer B 1648, from Bolivia. Pileus hyaline, hygrophanous, in fresh condition becoming white or in places very

slightly cinnamon-pallid or ocher, dried darkening even if well dried and becoming orange cinnamon in marginal region or "tortoise" color, the extreme margin reaching 13 K 10 M&P (ochraceous brown near raw sienna) while the center and the ray-like extensions from there along the furrows either remain white or become light ochraceous yellow ("Inca gold"), glabrous, sulculate to deep sulcate over up to four fifths of the radius, otherwise rugulose-uneven to rugose-venose, campanulate, later convex with an umbilicus or without it, the marginal region tending to become uplifted and the pileus hat-shaped, 22-45 mm broad. Lamellae white, not intervenose, very narrow to narrow, subclose to almost crowded, adnate. Stipe entirely white in very young specimens, soon chestnut and gradually lighter colored upwards with white apex, later brown all over, finely but

distinctly and consistently pubescent-pruinose all over, hollow, equal, 30-45 X 1-1.5 mm; basal mycelium tomentose to strigose, abundant, white, drying to light fulvous. Context thin, white, inodorous or with a slight aromatic odor.

Spores 8-10.3 X 2.7-4 A,fusoid to fusoid-oblong, hyaline, smooth, inamyloid. Hy- menium: basidia 19-20 X 5-6 1, clavate; cystidia none; cheilocystidia like epicuticular broom cells. Hyphae hyaline, thin-walled, with clamp connections, strongly pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of vesiculose to vesiculose-

clavate-pedicellate cells, some with, some without setulae, both becoming thick-walled in

age, the setulose ones recalling broom cells of the Siccus type with a main body of 8-28 X 6.5-27 p, hyaline to brownish hyaline, wall when thickened up to 1.5 p in diameter, setulae 4-23 X 1-7 (at base), , cylindric or subcylindric, or conic, erect and apical, 2-many, obtuse to needle-sharp-acute; smooth epicuticular cells corresponding to those of the section Globulares, 13-28 X 8-27 ., hyaline, in scalp preparations often appearing angular because of mutual compression, at first about equal in number as compared with the broom cells but in mature and overmature specimens mostly much more numerous than



Marasmius 245

the broom cells; covering of stipe consisting of numerous but not crowded dermatocystidioid hairs or setae which are 20-65(-100) X 4.5-9 u, entire or with one to three setuloid or sterigmatoid apical erect excrescences, cylindric or ventricose, rarely clavate, apex obtuse to rounded, rarely subacute, hyaline to stramineous or melleous, thick-walled (wall about 2 u thick), few occasional hairs forked in middle or below or even twice forked.

On leafmold and very rotten wood under dicotyledonous trees in rain forest. MATERIAL STUDIED. COLOMBIA. Valle: Buenaventura, San Joaquin, 180 m alt, 22 Apr

1968, Singer B 6273 (F). BOLIVIA: Beni: Vaca Diez, 7 Mar 1956, Singer 1648 (LIL) type; 3 km below Guayaranerfn in virgin forest, 12 Mar 1956, Singer B 1889 A (LIL), paratype.

ILLUSTRATION: Singer (1965) p1 23, fig 122.

190b. Marasmius pseudoniveus var amylocystis Singer, var nov TYPE. Singer B 6297, from Colombia. Differt cystidiis pseudoamyloideis. Typus in F conservatus est. Differs from the type variety in pseudoamyloid cystidia. Pileus white, dried white with very pale buffish center and/or margin, glabrous,

slightly sulcate, with smooth center, convex, about 22 mm broad. Lamellae white, subclose or close, narrow, not intervenose, adnexed to subfree. Stipe ochraceous brown below, white above, hyaline pilose-pubescent all over, equal, 30 X 1.5 mm; basal mycelium strigose, abundant, white. Context white in the pileus, thin, inodorous.

Spores (7-)7.7-8.7 X 2.5-3 ,u, oblong, smooth, hyaline, inamyloid. Hymenium: basidia (11-)16-17.5-24 X 4-5.8 p, 4-spored; cystidia both on edges and sides rather numerous, 13.5-37 X 2-7 i, thick-walled, often bifurcate or even subramose, with subacute to obtuse attenuated apex, often widened in the lower portion and there ventricose but also subulate to subcylindrical, hyaline, pseudoamyloid at least in the larger upper portion. Hyphae with clamp connections, filamentous, thin-walled, pseudoamyloid. Cor- tical layers: epicutis of pileus hymeniform, consisting of broom cells, broom cell-like cells without setulae, and dermatocystidia like the cystidia of the hymenium; broom cells with ventricose-subclavate to vesiculose, main body hyaline to palest stramineous, setulae where present up to 7 X 1.3 u, concolorous with the main body. Covering layer of the stipe consisting of hairs 16-90 X 4.5-10 p, cystidiform hair- to seta-like, below usually but not always ventricose, strongly pseudoamyloid excepting sometimes the base and rarely the tip, with rounded, rarely subacute tip, some bifurcate above, some with a lateral bulge or wavy-constricted all over, hyaline, thick-walled, wall 0.5-1.5(-2) A thick.

On rotting wood. Colombia. MATERIAL STUDIED. COLOMBIA. Valle: Buenaventura, Calima, 22 Apr 1968, Singer B

6297 (F), type.

191. Marasmius heterocheilus Singer, Sydowia 18: 337. 1965. TYPE. Singer B 2076, from Bolivia. Pileus cinnamon ("Maya" to 12 A 11 M&P), glabrous, smooth (not striate or sul-

cate or corrugate), convex and at maturity with flattened center or with a slight depression around the subumbonate disc reaching over 50 mm in diameter. Lamellae pale cinnamon with pallid edges, crowded, many forked, narrow, with an occasional anastomosing vein but not typically intervenose, sinuate-attingent to sinuate-subfree, dried appearing subdecurrent; where eaten by insects or snails the lamellae regenerate with a structure like the one of the surface of the pileus. Stipe brown with paler apex, entirely distinctly finely pruinate, tapering downwards, reaching over 8 mm in length, about 4 mm in diameter at apex and 2.5 mm at base, basal mycelium over a thick whitish cottony




mycelial mat, white, tomentose on base of stipe. Context white, inodorous.

Spores 6-6.3 X 3.5-4 ,, ellipsoid, hyaline, thin-walled, inamyloid. Hymenium: basidia 20-25 X 6.5 A, 4-spored, clavate; cystidia on sides of lamellae none; cheilocystidia not like the epicuticular broom cells but characteristically polymorplious, mostly clavate and simple, much more rarely cylindric or clavate with one to four apical sterigmatoid mucros ( e g 4 j long), often irregularly contorted and with small appendages at the tip, rarely branching into two to three branchlets, if simple about 20-28 X 5.5-7 p, hyaline or subhyaline. Hyphae hyaline, strongly pseudoamyloid with numerous clamp connections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the

Siccus-type, cylindric or clavate, often hand-shaped or Clavariella-shaped, with short to long obtuse setulae. Elements of the stipe exactly like the cheilocystidia but more often clavate-subcapitate or knob-shaped (reduced) and in the latter case e g 4.8 X 4.5 u, if subcapitate capitulum 4.8-5.5 , in diameter and constriction about 3 , in diameter, hyaline to stramineous, often shorter than the cheilocystidia, numerous and in places crowded but not palisadic.

On rotting foliage and small wooden sticks in rain forest. MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, Guayaranerfn, 17 Mar 1956, Sinzger

B 2076 (LIL) type. ILLUSTRATION. Singer (1965), pl 23, fig 121.

Stirps Cohaerens

This group of species (no. 192-197) shares the pigmented setae or setoid cystidia on the pileus and/or the hymenium with the mostly temperate species Marasmius cohaerens; in one species these bodies are all hyaline but then the lamellae are subdistant and the pileus strongly pigmented (this species, Marasmius flammans Berkeley, forms the transition to stirps Actinopus, coming closest to Marasmius pseudoniveus var amylocystis, which differs in close or subclose lamellae and white pileus).

192. Marasmius spiculosus Singer, Sydowia 18: 343. 1965. TYPE. Singer B 1206, from Bolivia. Pileus rusty to deep reddish brown ("alamo," "cocoa" M&P) on margin sometimes

brown ("cognac"), at first smooth and more rusty, later (especially when dried) up to four fifths of radius sulcate glabrous to finely velutinous (under a lens) and sometimes with fimbriate margin, glabrescent, convex to campanulate or almost conical, 6-15 mm broad. Lamellae sordid cream-grayish, pallescent, rather narrow or medium broad, subdistant to almost crowded (12-13 or more through-lamellae) intermixed with numerous

lamellulae, adnate. Stipe light brown to brown ("bure" to "oak" M&P) later chestnut at base, eventually concolorous with pileus or darker, dried stramineous to reddish brown, finely pubescent-velutinous all over, sometimes microscopically glabrescent, especially through handling and in age, equal, 2248 X 0.8-1.2 mm; basal mycelium forming a in somewhat larger pileus, cream white to pale buff lamellae and broader cystidia.

Spores 12-21 X 4-5 ,, hyaline, smooth, inamyloid. Hymenium: basidia 23-24 X 7.5-8.2 ,, 4-spored, hyaline, clavate, mostly strangely thick-walled and reminding one of the cystidia when seen without sterigmata; basidioles fusoid; cystidia numerous, 29- 36 X 7-9 p, hyaline, sometimes deeply rooting, versiform, often ventricose with flattened

capitulum (in the manner of Gloiocephala cystidia and dermatocystidia) thick-walled and

opaque; cheilocystidia none but edge densely beset with setae which are hyaline or col-



Marasmius 247

ored, inamyloid, much like the setae occurring on the surface of the pileus; sides of lamellae without setae. Hyphae with clamp connections, pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type and setae; broom cells with main bodies subhyaline and setulae short to medium long, moderately thick at base, light golden melleous in some, subhyaline in other cells, spinulose; setae like those of the surface of the stipe; stipe surface of a discontinuous layer consisting of numerous setae which are actually a special kind of broom cells but have the setulae so strongly pigmented that they appear quite well differentiated, usually simple and consisting of a basal cylindric to inflated portion and a setulose upper portion; lower portion hyaline or more often brown, mostly thick-walled, 9-59 X 4-11 ,; setulose upper portion (rarely two to three to one basal portion) (3-)10-45 X 3-5.5 (at base) , simple or more rarely forking, mostly brownish or melleous (at least in lower portion), attenuate to a subacute to needle-sharp acute tip; wall of lower portion up to 1.3 u thick; there all transitions from setae to broom cells in some collections and in the broom cell-like bodies the main body generally strongly reduced.

On wood of dicotyledonous and monocotyledonous (palms) trees, in rain forest and tropical montane forest.

MATERIAL STUDIED. BRAZIL. Guapore, Guajaramirim, 20 Mar 1956, Singer B 1806 (LIL), paratype. BOLIVIA. La Paz: Nor-Yungas, Rio Yariza, 16 Feb 1956, Singer B 1206 (LIL) type.

This is extremely close to M. fulvovelutinus Beeli from Africa (Zaire) which differs in somewhat larger pileus, cream white to pale buff lamellae and broader cystidia.


193. Marasmius cohaerens (Albertini & Schweinitz ex Fries) Cooke & Qualet, Clavis Synopticus Hymenomycetum Europaeorum, p 135. 1878.

Agaricus cohaerens Albertini & Schweinitz ex Fries, Systema Mycologicum 1: 253. 1821. Agaricus ceratopus Persoon, Mycologia Europaea 3: 214. 1828. Marasmius ceratopus (Persoon) Quelet, Flore Mycologique p 319. 1888. Mycena cohaerens (Albertini & Schweinitz) Gillet, Champignons de France, Hymenomyc[tes

p 275. 1874. Agaricus spinuliferus Peck, Ann. Rep. New York State Museum 24: 62. 1872. Marasmius setulosus Murrill, Bull. Torrey Club 67: 150. 1940.

193a. Marasmius cohaerens var cohaerens The type variety has not been collected in the neotropics. The type from Germany

is probably not conserved.

193b.Marasmius cohaerens var americanus Singer, var nov TYPE. Singer N 700, from U.S.A. (Michigan). Differt lamellis minus distantibus, latioribus. Typus in F conservatur. Pileus fulvous to cocoa-brown, the margin paler (pale reddish brown), in button

stage "Mirador, Argentina" (M&P), soon becoming almost "Alamo" to "tarragona" or "Tuscan tan sauterne" (M&P) with "kis kilim," "burnt umber" or "cocoa br" (M&P) smooth or rugulose, very finely hispid, campanulate, the campanulate-umbonate, eventually still more repand, 10-26-(34) mm broad. Lamellae not pure white, e g "Almond biscuit" or "Longbeach" (M&P) under a lens brick-red to brown spotted from the setoid cystidia, with concolorous edge, rather narrow, later medium or moderately broad, (2-3.5 mm broad), intermixed, sometimes ventricose, intervenose, close, later subclose to subdistant, not or scarcely intervenose, adnate, rounded-adnexed, sometimes subfree. Stipe white, early becoming fulvous below, later bay-brown to reddish chestnut with




pallid apex, often canaliculate, finely hispid or pubescent, hollow, subequal, (5-)30-80 (-150) X 1.5-3(-6) mm; basal mycelium abundant, often extensive. Context rather fleshy in the pileus, toughish and elastic in the stipe, white in the pileus, unchanging, inodorous with a slight odor of Marasmius oreades.

Spores 7-8.5 X 4-4.8 p, ellipsoid, with suprahilar depression, smooth, hyaline, inamyloid. Hymenium: basidia 4-spored, occasionally some bisporous; basidioles fusoid; cystidia setoid, 48-98 X 8-13 u, fusoid, thick-walled (wall 1.3-2.2 t thick), acute, rarely forked, base hyaline and mostly remaining so, upper portion more rarely almost entirely rusty tawny or brownish ochraceous in the great majority, few if any entirely hyaline. Hyphae of the pileus trama hyaline, pseudoamyloid, with clamp connections; those of the stipe thick-walled and parallel in adult material. Cortical layers: epicutis of the pileus hymeniform, consisting of broom cells and setoid dermatocystidia, the former with a hyaline to stramineous thin-walled main body and 6-7 p long acute, subhyaline to brown setulae; the setoid dermatocystidia pigmented and acute like the setoid cystidia of the lamellae but reaching only 90 in length; covering of the stipe consisting of similar setoid dermatocystidia.

On leaves of frondose and coniferous trees, rarely on very rotten wood buried in the leafmold, often cespitose to fascicular, in North America, reaching southern Florida.

MATERIAL STUDIED. U.S.A. Vermont: Middlebury, det. Peck, Herbarium Burt (FH); Michigan: Cheboygan County near Douglas Lake, Gorge, 21 Jul 1953, Singer N 700 (F) type; Florida: Arredonda, 29 Jul 1938, West & Murrill F 18267 (FLAS).

This is a marginal form as far as the neotropics are concerned but the southern limit of this form is not known. The American collections seem to belong mostly to this variety which differs only slightly from the European type which has more distant

(16 to 19 through-lamellae) and broader (3.5-5 mm) lamellae and perhaps slightly larger spores and grows more often on coniferous wood or needles than on Dicotyledones. A somewhat intermediate form has been observed by me in New York, Huntington Forest. Here the lamellae were broad, distant, gray and brown-margined, on Fagus leaves (Singer N 171, FH). This shows that var americanus is neither a geographic race nor an independent species.

194. Marasmius flammans Berkeley, Hook. Jour. Bot 8: 136. 1856. TYPE. Spruce 97, from Brazil. Pileus orange fulvous, dried "Burnt Sienna" [Ridgway], according to Dennis, nearer

Mars Orange when soaked up with paler radiating lines, 22 mm broad. Lamellae ochraceous (now yellowish) narrow, of two to three lengths although not quite regularly tri- or didymous, subdistant, attingent-adnate. Stipe said to be fuscous, now cinnamon rufous in the herbarium, glabrous, smooth, equal, 15 X 1 mm.

Spores unknown (none in type specimen). Hymenium: basidia 4-spored; cystidia numerous, setoid, inamyloid, subhyaline to stramineous, 30-48(-55) X 9-9.8(-11) A, mostly acute and simple, rarely with a rounded apex (and then ampullaceous-subcapitate or

clavate) and rarely with lateral spinulose excrescency (and then like most cystidia fusoid and acute), many very deep rooting, with very thick (3-3.7 , diameter!) wall; cheilocystidia subulate, acute, hyaline, thin-walled. Hyphae pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells and setoid dermatocystidia; broom cells with main body about 8 p long and setulae about 6-13 uplong, all apical and erect; setoid dermatocystidia much like cystidia of the hymenium, fusoid, acute, very thick- walled, but + pseudoamyloid.



Marasmius 249

On fallen leaves in tropical rain forest. MATERIAL STUDIED. BRAZIL. Amazonas, "Panure" (Ipanure = Sao Jeronimo) Spruce 97

(K) type. Only the type is known. This was restudied by both Dennis and this author. The

above is a redescription based on the data given by Berkeley, Dennis and this author. ILLUSTRATION: Singer (1958a) fig 19.

195. Marasmius echinatulus Singer, Sydowia 12: 98. 1958. Marasmius echinatus Theissen ex Singer, Lilloa 25: 212. 1952, non Romell (1926). Marasmius cohaerens var brasiliensis Theissen, Broteria 8: 62. 1909. TYPE. Rick, from Brazil. Pileus orange to deep rufous ferruginous (11 C 9, 11 D 10, "mosque," "terra cotta,"

"burnt sienna," "Ginger, Kaiser brown," "talavera," "Eldorado," "ferruginous," even deeper and richer than "Chrysanthemum" sometimes reaching "feuille morte," "tarragona"), pallescent, in age becoming pallid with or without ferruginous spots, at first finely pubescent-subpruinose with hyaline pubescence, at first with a strongly incurved margin, later tending to have uplifted margin, in age somewhat hygrophanous (dry cinnamon white) but young not hygrophanous and little changed on drying and dried, when hygrophanous long-transparently striate, at maturity with sulcate margin, otherwise smooth or rugulose to reticulate-venose, hemispheric, then campanulate to convex and often umbonate, even-

tually subapplanate, 11-25 mm broad. Lamellae white, pallid, cream white, sometimes slightly crenulate, narrow to (more rarely) broad, mostly medium broad, close to subclose, at first (in buttons) with an adnate collarium, i e subcollariate but very early be-

coming rounded-free, later adnexed to even adnate and then eventually separating from

apex of stipe, generally not intervenose but in some large mature caps very weakly sub- intervenose. Spore print white. Stipe whitish, very early becoming subconcolorous from base upwards, later apex white to dull cream, otherwise "Inca gold" to "doubloon" then chestnut or ochraceous brown to cocoa brown ("cocoa" to "Sudan brown", "Saratoga") rarely even mature specimens with almost entirely whitish stipe, slightly pubescent- pru- inate, sometimes glabrescent, equal or slightly tapering upwards or downwards, central, rarely somewhat eccentric, at times with a slight bulb at base up to 0.5 mm broader than base diameter), terete or somewhat compressed, 18-36 X 1-5 mm; basal mycelium at first a mycelial pad (white), later sparsely to distinctly tomentose to strigose, pallid whitish to sordid whitish or pure white. Context white, inodorous or with an odor of "sauerkraut" or Collybia dryophila, or of Marasmius oreades.

Spores 6.5-9.7 X 2.3-4.5 p,, oblong to oblong-subfusoid, thin-walled, hyaline, smooth, inamyloid. Hymenium: basidia 16-28 X 4-6.7 p, clavate, 4-spored; basidioles clavate to fusoid; cystidia none but cystidioles at times few, scattered, subampullaceous e g 29 X 7.5 ,; opaque, oleiferous hyphae occasionally entering the hymenium but never reaching the surface also present; cheilocystidia of two types: (1) of the metuloid type, more numerous in the mature specimens, in buttons scarcely developed, later 18-35(-100) X 3.5-11 ,u, generally deep rooting, versiform but often cylindric to fusoid, more rarely clavate or am-

pullaceous, smooth or granularly incrusted, never projecting strongly, hyaline, generally with rounded tips, very slightly bluing in cresyl blue mounts, thick-walled, (2) of the broom cell type, more numerous in the young specimens and near the margin of the pileus but sometimes persisting in mature specimens, always scattered although sometimes numerous, mostly in form of transitions between normal broom cells and setae, hyaline, with strongly reduced main body and elongated spinulose setulae which are few, mostly once forked, pseudoamyloid, main body up to 22 X 7 ,, setulae 6.8-9(-15), long, strictly




acute; typical setae as found in epicutis of pileus only in young specimens, rarely found on edge, none on sides of lamellae; in some collections the marginal broom cells also occurring (but very sparsely) on the sides of the lamellae in the neighborhood of the edge. Hyphae hyaline, thin-walled in trama of pileus and hymenophore, strongly pseudoamyloid, with clamp connections; nongelatinous. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type and setae and also some transitional formations (between broom cells and setae); broom cells hyaline to pale stramineous, few brownish- ocher at times, about 14.5-16 X 5.3-6 ,, main body mostly hyaline or subhyaline, thin-walled or more rarely somewhat thick-walled, setulae collectively sometimes spreading to as broad as 8 s, setulae 4-8 , long, strictly apical; setae scattered to numerous among broom cells, 20-108 X 4-10.5 (at base) ,, brown to hyaline with golden melleous base, which is bulbous-ventricose, diameter of thin portion in the middle of seta 2.5-4.5 M thick, wall 0.7-1.2 , thick, smooth, weakly to distinctly pseudoamyloid, in older specimens often scarcely pseudoamyloid, tip acute to needle-sharp; transitional formations characterized by deep forking pseudoamyloid setulae, the swollen base assuming the role of the main body, the setulae shorter than the setae; surface of stipe beset with setae of the same type as encountered in the epicutis of the pileus, numerous.

On Pteridophyta and leaves of Monocotyledones (Bromeliaceae), Dicotyledones (e g Inga, mostly mixed leaf mold) perhaps also on conifers (Podocarpus), solitary or more often in groups, not cespitose.

MATERIAL STUDIED. COLOMBIA. Valle: Municipalidad Cali; Saladito, 15 Apr 1968. Siiiger B 6154 (F). BRAZIL. Rio Grande do Sul, Rick (as M. echinatus , ty pe of M. echinatiilus and M. echinatus Theissen ex Singer Theissen), (FH), type. BOLIVIA. La Paz: Nor-Yungas, Coroico, 26 Jan 1956 Singer B 531 (LIL), B 532 (LIL), 11 Feb 1956, B 1079 (LIL). ARGENTINA. Tucuman: Chicligasta, Rio de los Sosas, 27 Jan 1951 Singer T 1130 (LIL), paratype, 12 Mar 1950 Singer T 1694 (LIL), paratype; Las Lenguas, 27 Feb 1957, Singer T 3026 (LIL); Tafi, Ciudad Universitaria, 15 Jan 1955, Singer T 2029 (LIL), 30 Dec 1951.


196. Marasmius venezuelanus Dennis, Kew Bull 15: 97. 1961. TYPE. Dennis 1052, from Venezuela. Pileus cream buff or pale grayish brownish, dried reaching "Peruvian br" (M&P)

in the center, glabrous, finely striate on the margin or entirely smooth, convex, then flattened, 20-30 mm broad. Lamellae whitish or concolorous with pileus, beset with brown setae clearly visible under a hand lens, crowded, narrow, of several lengths, free. Stipe deep chestnut below, paler above, the extreme apex white, covered with minute, brown, downy, setulose hairs, hollow, subequal or tapering upwards, 48-60 X 1.5-3 mm, basal mycelium strigose, sometimes scanty, sometimes abundant, white.

Spores 6-9 X 3-3.5 A, tear-shaped, smooth, hyaline, inamyloid. Hymenium: basidia 17-20 X 5-7.5 ui, spored; cystidia brown, numerous, 60-90 X 11-17 #1, fusoid to ventricose with 1.5-2 M thick walls, simple or forked, the tips acute; cheilocystidia scattered to dense, some like the setoid cystidia, others like the epicuticular broom cells. Hyphae of the pileus hyaline, with clamp connections, strongly pseudoamyloid. Cortical layers: epicutis of the pileus hymeniform, consisting of firm- or thin-walled hyaline or pale stramineous cells, 10-20 X 6-20 u, with interspersed bright brown, rarely hyaline, thick-walled broom cells with setulae 2-5(-26) X 0.6-2.5(-3)u1, also brown, rarely hyaline; occasional short, simple or forked brown setae which are like those of the hymenium; somewhat shorter ones also present; covering of stipe as in Marasmius jalapensis.

On rotten wood and other debris in cloud forest, solitary. MATERIAL STUDIED. VENEZUELA. Dto. Federal, El Avila, 13 Jun 1958, 2000 m alt, Dennis



Marasmius 251

1052 (K) type. COLOMBIA. Valle, N of Dagua road km 25, 1950 m alt, 28 Apr 1968, Singer B 6455 (F).

ILLUSTRATION: Singer (1965), pl 21, fig 114 DV; pl 22 fig 115 DV; Dennis (1961) fig 50.

197. Marasmiusjalapensis Murrill, North American Flora 9: 264. 1915. TYPE. W. A. & E. L. Murrill 84, from Mexico. Pileus cream buff to cream gray (between "sweet meat" and "tansan"), hygrophanous,

dry fading to pale cinnamon or cinnamon white, smooth, later sulcate over two thirds of the radius, drying from the center outwards, glabrous, convex, then subapplanate, subumbonate, but eventually with a depression in the center, 25-67 mm broad. Lamellae whitish to concolorous with pileus, beset with brown setae clearly visible under a hand lens, narrow, intermixed, not intervenose, close to subclose, free to adnate. Stipe red brown ("Mohawk") more fulvous above and gradually paler towards a whitish (or concolorous with the lamellae) apex, brown-pruinate from setae, smooth, equal, 40-180 X 1-8 mm; basal mycelium strigose to tomentose and often connected with a widespread mycelial mat, white or whitish, slightly fulvescent at places when dried. Context white, inodorous.

Spores 7-9 X 2.5-3.5 ,, oblong to ellipsoid-cylindric, hyaline, smooth. Hymenium: basidia small; cystidia in the form of setae numerous on sides and edges of lamellae, setae 44-110 X 7-17.5 A, acicular-fusoid, needle-sharp acute, thick-walled (wall 2.7-3.7 lthick), brownish-fulvous to ochraceous brown in KOH, smooth, partially somewhat pseudoamyloid; cheilocystidia not differentiated or not found. Hyphae of pileus trama and hymenophoral trama hyaline in KOH, strongly pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type and setae, also of some transitional formations (between broom cells and setae); broom cells hyaline to golden melleous in main body which is vesiculose, 7-12.5 A broad, mostly thin-walled and with golden melleous, more rarely hyaline setulae, some of the broom cells without any setulae and some with strongly enlarged few deep brown setulae (transition to setae); setae relatively few, simple, subulate, more rarely fusoid, smaller than those of the hymenium. Stipe beset with setae similar to those found in the hymenium but more subulate.

On leafmold and rotten wood in tropical and tropical-montane (cloud) forest. MATERIAL STUDIED. MEXICO. Veracruz: Jalapa, 12-20 Dec 1909, W. A. & Edna L. Murrill

84 (NY) type. BOLIVIA. Beni: Vaca Diez, Ivon 3 Apr 1956, Singer B 2470 (LIL). The slightly deeper colored (isabelline to fulvous according to Murrill) pilei of the

Mexican type may be due to this having been a younger stage of the carpophore, as in M. echinatulus. The Bolivian material is paler.

ILLUSTRATION: Singer (1965) pl 21, fig 114 (not 114 DV);pl 22, fig 115, (not 115 DV).

Marasmius sect Fusicystides Singer in Singer & Digilio, Lilloa 25: 287. 1952. TYPE SPECIES. Marasmius fusicystis Singer in Singer & Digilio. Pileus pigmented, with an epicutis of the Ramealis-type or irregularly arranged

broom cells; spores large; lamellae not collariate; stipe oblique and eccentric, lateral or rudimentary (thus habit of carpophores pleurotoid) with somewhat mycelioid base, black rhizomorphs absent; partially pseudoamyloid, even in the pileus, with clamp connections; on dead wood.

This is the only section which combines pleurotoid habit (and is macroscopically very similar to the species of the following section) with the absence of a hymeniform layer of the surface of the pileus. It is sharply separated from Marasmiellus sect




Marasmiellus by the distinctly pseudoamyloid hyphae. It comes closest to sect Neo- sessiles, especially Marasmius paulensis.

The only species is M. isabellinus Pat.

198. Marasmius isabellinus Patouillard in Patouillard & Lagerheim, Bull. Soc. Mycol. France 9: 126. 1893.

iMarasmius fusicystis Singer in Singer & Digilio, Lilloa 25: 187. 1951. Pileus yellowish to golden isabelline or with salmon tinge or dull ochraceous

orange (9 B 6, 10 H 7, 12 L 9, 10 B 6, M&P) eventually fading to orange-yellowish or

yellowish (10 F/H - 4/6, M&P) or assuming an "India spice" (brownish) tinge in some

collections, glabrous, smooth, becoming sulcate or radially rugose, elliptical-conchate or

heart-shaped in outline, sinuate behind, convex, but more applanate behind in most mature specimens, sometimes even umbilicate, 5-28 mm broad. Lamellae yellow ("capucine y," "sunburn, French beige") in age or a dull orange (11 J/K 10, M&P), finely orange brown or orange punctate when seen under a lens at least when fully mature, distant or subdistant (5-13 through-lamellae), with a few lamellulae or forked lamellae intermixed, narrow to rather broad, mostly at first narrow and later medium broad (1-4 mm broad), becoming distinctly intervenose in almost all fully mature

specimens, attentuate-attingent, attenuate-subfree, or more or less rounded-adnexed.

Stipe concolorous with the pileus or somewhat paler, often more sordid, at first mostly white, often with a deep orange yellow ring around the apex of the stipe just where the lamellae begin, eventually becoming dirty brownish, subglabrous, pruinate under a lens, often curved, equal or attenuate toward the apex or the base, not insititious or subinsititious, eccentric, mostly strongly eccentric or even laterally attached, sometimes rudi-

mentary, 1-3.5 X 0.1-1 mm; basal mycelium white, sparse or visible only under a lens, forming a small patch at the point where the stipe rises from the substratum. Con- text white, but mostly concolorous underneath the cuticle, thin, not fragile; odor none or weak like fried Boletus edulis.

Spores 11.7-16.5 X 4-5.8 p, fusoid or oblong, often curved and Fusarium-like, some almost clavate, thin-walled, the walls swelling slightly in KOH, with suprahilar depression or applanation, smooth, hyaline, inamyloid. Hymenium: basidia 28-36 X 7.5-9 1, 4-spored; cystidia on both sides and edges numerous, 20-80 X 4-10 p, fusoid, mostly entire, the "neck" sometimes finger-like elongated and about 2 , broad, with obtusate tip, with thin (0.2-0.5 ,) wall but this often appearing thick (up to 1.5 p) because of a hyaline to yellow succineous resinous granular or sheet-like appressed incrustation, the wall proper hyaline but often colored lemon yellow by the pigment of the incrustation in the preparations, deeper rooting than the basidia and projecting 12 p and more beyond the basidia, those of the edge of the lamellae near the margin of the pileus often forked or distorted. Hyphae of the pileus-trama in the lower stratum

hyaline, thin-walled, few firm-walled, 3-5 p broad, with clamp connections, weakly pseudoamyloid to inamyloid; those of the infra-hypodermial layer palest orangy stramineous in the wall and often slightly roughened by granular material, most with firm, few with thin wall, 3.5-7 p broad, rarely broader, pseudoamyloid, at least the firm-walled ones; those of the stipe subparallel, thin- to thick-walled, strongly pseudoamyloid, especially the thick-walled ones; subhymenium inamyloid; hymenophoral trama regular. Cortical layers: epicutis of the pileus of hyphal ends which are partly repent, partly obliquely ascendant or erect and the erect ones in places almost subhymeniform but not forming a continuous hymeniform layer, in places forming a trichodermial layer and thus differentiated from the radially arranged hyphae of the upper



Marasmius 253

layer of the trama of the pileus, the terminal cells mostly oblique or erect, firm- but not thick-walled, some at the apex somewhat thinly incrusted and golden, 17-20 X 7-10 , some elongated to 31 long, mostly clavate or cylindrical, many entire, but some slightly to distinctly diverticulate, the diverticula 2-7 X 0.5-2.5 A, finger-like and mostly erect and apical, some more subulate but the tips of the terminal cells and the diverticula mostly obtuse; dermatocystidia also mostly present but rather rare, similar to the hymenial cystidia but more irregular and with less incrustations, e g 30 X 6 u; hypodermium of filamentous hyphae which ascend towards the terminal hyphae; cov-

ering of the stipe consisting of terminal members of the cortical hyphae and a few

dermatocystidia, these ampullaceous or fusoid but smaller than the epicuticular and

hymenial ones, all hyaline and inamyloid. On wood or bark of dead shrubs and trees. Known hosts: Phoebe porphyria,

Duranta serratifolia, Eugenia pungens, Coffea arabica, Allophylus edulis, Myrtus sp, Piptadenia sp.

MATERIAL STUDIED: BRAZIL. Rio Grande do Sul, Pareci. 2 Nov 1951, Singer B 54 (LIL). ECUADOR. Pichincha, Quito, Rio Machangaro at 2600 m alt, 20 May 1973, Singer B 7541 (F), topotype, Feb 1892. Lagerhei,m (FH), type; Cotocallao (FH), paratype. BOLIVIA. La Paz, Nor-Yungas, Coroico, 26 Jan 1956, Singer B 549 (LIL). ARGENTINA. Tucuman: Anta Muerta, 2 Jan 1950, Singer T 802 (LIL, MICH), type of M. fusicystis, Quebrada de Lules towards Villa Nougues, 8 Feb 1961, Singer T 1641 (LIL); - Rio Cochuna, 11 Feb 1952, Singer T 1837 (LIL); Ciudad Universitaria, 25 Dec 1959, Singer T 3422 (LIL).

In this section we have also a palaeotropical species which is represented by Marasmlius campanella Holtermann, (1898) as interpreted by Petch (1947, p 35; specimen: Peradeniya, rufous form, 14 Oct 1914, Petch 4171 (FH), studied by me) which differs from A. isabellinus in narrower spores (14-16 X 4.3-4.5 u). Petch thinks that Marasnmius rufescens Berkeley & Broome is the same species.

ILLUSTRATION: Singer (1958a) fig 1; (1965), pl 7, fig 10.

Marasmius sect Neosessiles Singer, Mycologia 50: 104. 1958. TYPE SPECIES. Marasmius neosessilis Singer Pileus pigmented or not, with the epicuticular broom cells hymeniformly or sub-

hymeniformly arranged; spores mostly medium-sided to large; lamellae not or indis-

tinctly collariate; stipe oblique and eccentric later, or rudimentary (thus habit pleurotoid) with insititious, subinsititious to mycelioid base (unless rudimentary), hyphae pseudoamyloid or inamyloid with or without clamp connections; on dead wood and leaves.

This is the only section that combines pleurotoid habit with the presence of a

hymeniform or subhymeniform arrangement of broom cells on the surface of the

pileus.

Key to the Species of Section Neosessiles

1. Hyphae with clamp connections. 2. Cystidia thick-walled, non-incrusted; cheilocystidia with setulae; stipe at first central

later strongly eccentric, not insititious; lamellae neither forked nor intervenose, subclose to subdistant; strongly pseudoamyloid, thick-walled elements in the trama of the stipe present. 199. M. polycystis.

2. Cystidia either thin-walled or with resinous incrustation or absent; not combining the above characters. 3. Cystidia or cheilocvstidia with succineous incrustations present; spores 11-15 X

4-6.24 . 4. Stipe never absent, never insititious; pleurocystidia few, hyphae or pileus indis-

tinctly or not pseudoamyloid; on dicot branches. 200. M. paulensis.




4. Stipe insititious or absent; pleurocystidia numerous; hyphae of pileus trama slowly but distinctly pseudoamyloid; on leaves and petioles. 201. M. cecropiae.

3. Cystidia small and basidiomorphous or else absent on the sides of the lamellae. 5. Spores 11.5-14.5 X 2.2-4 p; hyphae of the pileus distinctly pseudoamyloid and

stipe base subinsititious or not insititious. 202. M. griseoroseus. 5. Spores smaller or hyphae of the pileus inamyloid (or both); stipe usually

insititious or subinsititious. 6. Pileus deep fuscous; stipe fuscous; spores 9.5-11 X 5.3-5.5. 203. M1. lstilago. 6. Colors and/or spores different. Pileus pale gray, pinkish cinnamon or white

or dirty ochraceous, dirty fuscous, etc., apricot pink to salmon; spores (7-)8-10 X 4-4.8 . different only if hyphae of pileus all inamyloid. 7. Hyphae of the pileus-trama inamyloid but stipe trama may or may not

be pseudoamyloid; spores up to 10 p long or longer. 8. Spores 7.8-9.7 X 4.7-5.5, ; pileus white to fuscous or white to gray;

stipe dark brown or dark fuscous. 204. Al. sessiliaffinis. 8. Spores either larger or narrower or else color of the pileus different.

9. Pileus 7-!4 mm broad, rusty brown or light grayish fuscous (fresh); spores 8-10.5 X 3.8-4.7 u. 205. AM. teinuissilmus.

9. Pileus smaller and differently colored (often fresh wvhite or salmon); spores 7-10 X 3-5(-6)p or 9-12.3 X 4.3-6 , but within these limits very variable in shape. 10. Spores 9-12.3 X 4.3-6.5u; pileus 4-7 mm broad, white at

first, soon becoming partly or entirely light argillaceous. pale fuscous or dirty brown; lamellac distant; cystidia none. 206. M. spaniophylllus.

10. Spores in an average shorter: 7-10 X 3-5(-6)p,; pileus smaller than 4 mm, salmon pinkish to pinkish ochraceous; lamellae very distant to subclose (2-10 through-lamellae at maturity); cystidioles present, numerous. 207. M. sej/unctus.

7. Hyphae of the pileustrama pseudoamyloid; spores 208. MI. neosessilis. up to 10 p long, not longer.

1. Hyphae without clamp connections. 209. M. pleurotelloides.

199. Marasmius polycystis Singer, sp nov Fig 84. TYPE: Singer B 1499, from Bolivia. Pileo salmoneo-aurantiaco, 6-8 mm lato; lamellis pileo multum pallidoribus; stipite

demum fortiter excentrico. Sporis 9 X 4.2 p; cystidiis crassitunicatis, haud incrustatis; hyphis fibuligeris. Ad corticem ramorum delapsorum dicotyledoneorum in Bolivia. Typus in F conservatus est.

Pileus salmon orange (9 F 6 then 10 C 5/6, M&P) glabrous, sulcate, circular, then elliptical in outline, 6-8mm broad. Lamellae whitish with an orange shade (like the pileus but much paler), numerous but only subclose to subdistant (six or more through- lamellae) intermixed with lamellulae, not forked, rather narrow but never venose when mature, not intervenose, attenuate-subfree or adnexed. Stipe concolorous with the lamellae or the pileus, curved or appressed to the substratum, at first central but later strongly eccentric, not insititious, subequal, mature 1-2 X 0.7 mm; basal mycelium fibrillose, well developed, white, no rhizomorphs. Context inodorous.

Spores (few seen) 9 X 4.2 ,, fusoid, smooth, hyaline, inamyloid. Hymenium: basidia 15 X 5 p, 4-spored; cystidia numerous on both edges and sides of the lamellae, 24-43 X 6-9.8 u, characteristically thick-walled (wall about 1-1.5 p thick), opalescent and wall without clear inner outline, not incrusted, versiform, often subcylindric, clavate, ventricose, hyaline. Cheilocystidia shorter than pleurocystidia, with apical and often somewhat divergent orange hyaline to brown setulae, these about 4-5 p long, knotty or bifurcate in many cells. Hyphae in hymenophoral and pileus-trama hyaline, not or very weakly pseudoamyloid, with clamp connections. Hyphae of the stipe filamentous, also with clamp connections, many walls strongly red-pseudoamyloid, thick



Marasmius 255

cortical layers: epicutis of the pileus hymeniform, consisting of broom cells, main body 11.5-17 X 7-10 , hyaline or brownish, ventricose, setulae either as in the cheilocystidia or orange brownish in their majority (in mature caps) and often simple.

On bark of dead branches of Dicotyledones. Yungas. MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, 26 Feb 1956, Singer B 1499 (F),

type.

200. Marasmius paulensis Singer, sp nov Fig 85, 86. TYPE. Singer B 4099, from Brazil. Pileo aurantiaco vel aurantiaco-pallido, rarius albo, ? 12 mm lato; lamellis pileo

paullum pallidoribus, intervenosis, badio-punctatis; stipite exiguo, demum evanescente.

Sporis 12.5-15 X 4.5-6.2 ,; cystidiis paucis; hyphis pilei haud fortiter pseudoamyloideis vel inamyloideis, fibulatis. Ad ramos dicotyledoneos in Brasilia. Typus in BAFC conservatus est.

Pileus orange to orange-pallid, in some white, glabrous, deeply sulcate, about 12 mm broad. Lamellae concolorous with the pileus but somewhat paler, narrow, distant, strongly anastomosing-connected, under a lens with red brown punctations along the edge. Stipe concolorous with the pileus, small and eventually disappearing, not insititious, up to 3 X 0.3 mm; basal mycelium consisting of a small white tomentum applicate to substratum. Context with a slight odor of sauerkraut.

Spores 12.5-15 X 4.5-6.21, fusoid, with suprahilar depression, thin-walled, smooth, hyaline, inamyloid. Hymenium: basidia 25-37 X 7.5-10 p, clavate, 4-spored; cystidia none or a few like those at the edge but without incrustation; cheilocystidia 35-53 X 4-9 ,, filamentous-cylindrical and wavy or constricted, some narrowly ventricose, rarely clavate, thin-walled, hyaline, rather numerous and some densely congre- gated in bunches, some crowned with a few appendages, most with a thick resinous red-brown or succineous incrustation. Hyphae of the pileus-trama hyaline, with firm to thick (0.3-1 ,) wall which is in most parts inamyloid, with clamp connections, not

gelatinized. Hyphae of the stipe subparallel to parallel with each other, hyaline, some, especially those of the rind zone, slowly but distinctly pseudoamyloid. Cortical layer: epicutis of the pileus rather subhymeniform than hymeniform, consisting of broom cells, main body 9.5-29 X 3.5-11 ,, some cells hyaline, some brown, thin- to thick- walled (wall up to 2 ,), erect but some laterally or apically branched with rounded- obtuse branches, setulae or appendages apical or lateral or on the upper side of the horizontal branches, rarely on the sides of the horizontal branches, many of them sec-

ondarily knotty to diverticulate or forked, 7-19 p long, mostly obtuse, concolorous with the main body or its upper portion.

On branches of dicotyledonous trees in Southern Brazil. MATERIAL STUDIED. BRAZIL. Sio Paulo, Reserva Florestal, 10 km N of Sao Paulo,

17 Dec 1964, Singer B 4099 (BAFC), type. This species may be confused with Marasmius isabellinus which, however, has

numerous, fusoid incrusted pleurocystidia and the epicutis is devoid of broom cells with thick colored walls and colored setulae.

201. Marasmius cecropiae Dennis, Kew Bull. 15: 92. 1961. TYPE. Dennis 1427, from Venezuela. Pileus light ochraceous (10 G 6), by dehydration in situ or in the herbarium be-

coming gradually light fulvous orange to orange-buff, glabrous and smooth, reniform in outline, convex, then applanate, 2-10 mm broad. Lamellae cream to "Spanish




yellow," very distant, 4-7 through lamellae, one or two forked or anastomosing by lower lamellae or veins, rather narrow,adnate or concurrent. Stipe absent or sublateral and inconspicuous, later lateral, inserted in the sinus in rear portion of pileus, rudimentary, short, curved, hyaline, pruinate, insititious, 1.5-0.7 mm, or less. Context very thin, inodorous.

Spores 11-14 X 4-4.8 , clavate-oblong, smooth, hyaline, inamyloid. Hymenium: basidia 20 X 6.3-7 ,, 4-spored; cystidia on sides and edges of lamellae, about 35 X 8 p, conspicuous and projecting, thin-walled or almost so, subfusoid, broader in lower half, hyaline but often crowned by a resinous succineous incrustation; cheilocystidia similar. Hyphae partly pseudoamyloid, with clamp connections. Cortical layers: epicutis of pileus consisting of broom cells which are vesiculose but sometimes shallowly bi-lobed or tri-lobed and the setulae somewhat divergent in places while in the entire cells the setulae are generally apical and erect, in the majority of the cases all erect (Siccus-type), cells alternating succineous melleous and hyaline (scalp view: mottled) 8-10 in diameter, setulae up to 4 long.

On fallen dicotyledonous leaves in tropical forest, perhaps always on Cecropia. MATERIAL STUDIED. VENEZUELA. Miranda: Guatopo, 25 Jun 1958, Dennis 1427 (K)

type. BOLIVIA. La Paz: Nor-Yungas, Carmen Pampa, 2000 m alt, Singer B 1496 (LIL). ILLUSTRATION. Singer (1965) p 11 1, fig 60; Dennis (1961) fig 42.

202. Marasmius griseoroseus (Montagne) Dennis, Kew Bull. 1952: 35. 1952. Agaricus griseoroseus Montagne, Ann. Sci. Nat. IV. 1: 98. 1 54. Pleurotus griseoroseus (Montagne) Saccardo, Sylloge Fungorum 5: 386. 1887. TYPE. Leprieur 1018 (PC), from French Guyana. Pileus white, drying dirty-buff, or somewhat orange-tinted, drying grayish-orange,

smooth, then slightly sulcate, glabrous or subglabrous, conchate to subreniform, sessile or with a stipe rudiment, up to 5 mm broad. Lamellae white, dried ochraceous, few (mostly up to four), narrow to medium broad and some incomplete or connecting ones venose, sometimes some forked, distant (1-5 through-lamellae), concurrent. Hy- menium of youngest stages smooth. Stipe none or rudimentary, minute and lateral; basal mycelium, if a stipe is present, visible under a good lens, radiating-fibrillose, white, or else stipe rising from a byssus-like appressed (to the substratum) mycelium. Context extremely thin, rather soft, white, inodorous.

Spores (9.5-)11.5-14.5 X 22.4p, oblong or subfusoid, smooth, hyaline, inamyloid. Hymenium: basidia 15-21 X 5.3-8.5,, clavate, hyaline, 4-spored; cystidia none; cheilocystidia present, like the epicuticular broom cells. Hyphae hyaline, not gelatinized, with clamp connections, pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of cells of the Rotalis-type with setulae much like those of M. spaniophyllus i e often irregular and forked, hyaline (in cheilocystidia always so) or melleous, main body vesiculose, 9-22 X 7.2-13k, setulae 1-3 X 0.2-0.4(-1)p.

On rotting branches and twigs in tropical rain forest. MATERIAL STUDIED: COLOMBIA. Valle, Finca La Araucana (Corr. Vergel, Municipalidad

Dagua) 1600 m alt, 14 Apr 1968, Singer B 6104A (F). BOLIVIA. Pando: Manuripi, Conquista, 25 Mar 1956, Singer B 2228 (LIL), B 2238 (LIL).


203. Marasmius ustilago Singer, Sydowia 18: 344. 1965. TYPE. Singer B 1870, from Bolivia. Pileus deep fuscous ("teakwood"), dried brown ("toast" to "English oak"), sul-

cate all over, elliptical in outline, convex, then more or less applanate, 5-13 mm



Marasmius 257

broad. Lamellae whitish ochraceous, dried ochraceous, narrow to medium broad, free, adnexed or adnate to a disc and often separating from it and becoming indistinctly collariate, intervenose at least in age, distant, 6-9, often intermixed with 1-2 lamellulae or one or two forked lamellae. Stipe fuscous, pruinose-tomentose, with white apex when fresh, equal, oblique or curved and often partially adnate to lower part of one side of pileus, dried 2-3 X 0.2-0.4 mm. Context white, very tough, inodorous.

Spores 9.5-11 X 5.3-5.5pj, ellipsoid, hyaline, smooth, inamyloid. Hymenium: basidia rather long; cystidia none; cheilocystidia hyaline, few with somewhat darkened setulae or fuscous-hyaline, otherwise like the epicuticular broom cells. Hyphae hyaline, some very thick-walled and elongated, reminding one of the hairs of Crinipellis and these distinctly pseudoamyloid at least in inner layer of wall, others thinner-walled and not or not consistently pseudoamyloid. All hyphae non-gelatinized, with clamp connections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells which are at first hyaline but become pigmented in both the main body and the setulae, reaching bister or fuliginous, thick-walled almost from the beginning, more or less pedicellate, vesiculose, 13-25 X 6.8-11 , some occasional ones reaching 33 X 22, setulae 1-3 X 0.6-2.5,, coarse and broad on the pigmented cells, obtuse, covering divergently the upper half of the cells, the latter becoming angular when seen from above (scalp view) and easily losing the setulae, reminding one somewhat of the spores of some Ustilaginales.

On dead dicot branches fallen to the ground in virgin tropical rain forest, very gregarious.

MATERIAL STUDIED: BOLIVIA. Beni, Vaca Diez, 3 km below Guayaramerin in virgin forest, 12 Mar 1956, Singer B 1870 (LIL) type.

ILLUSTRATION. Singer (1965) p 1 1, fig 59.

204. Marasmius sessiliaffinis Singer, Sydowia 12: 82. 1959. Maraamius spanioplhyllus var sessiliaffinis (Singer) Singer, Sydowia 18: 211. 1965. TYPE. Martin & Weldon 7606, from Panama. Pileus at first white or whitish, soon whitish gray or (because of minute fuscous

punctation) light brown to pale yellowish fuscous, eventually almost uniformly ferruginous ("Ginger, Kaiser br.") smooth, or sulcate to deeply sulcate, old ones at the same time often rugulose, glabrous, elliptical to circular with a sinus on one side, or reniform in outline, convex, later more or less applanate, 2-14 mm broad. Lamellae white or whitish and remaining so unless dried, irregular, 4-7 through-lamellae and 1-2, rarely more lamellulae, with smooth, interlamellar spaces or more rarely with few incomplete venose anastomoses, sometimes some old hymenophores with anastomoses of equal height or intervenose-rugose, at first with obtuse edges, later normally wedge-shaped and somewhat white-fringed but less so than in the preceding variety, moderately broad to rather narrow, free, more rarely adnexed to adnate and even then not collariate-separating, sometimes appearing concurrent because of the inconspicuous stipe. Spore print pure white. Stipe brownish to deep brown or brown black, subpruinate, somewhat glabrescent, in immature specimens whitish (concolorous with pileus), short and strongly curved or appressed to margin of pileus where it is sinuate, insititious, sublateral or lateral, inconspicuous. Context extremely thin, white, inodorous.

Spores (0.8-)7.8-9.7 X (4-)4.7-5.5u, ellipsoid, smooth, hyaline, inamyloid. Hy- menium: basidia 25-29 X 6.8-8.6 A, 4-spored; cystidia none; cheilocystidia on the venose and immature lamellae often not demonstrable but on mature ones present and




making the edge incompletely to completely heteromorphous, hyaline, otherwise like the epicuticular broom cells. Hyphae hyaline, inamyloid, with clamp connections, ir-

regularly shaped to filamentous, not gelatinized. Cortical layers: epicutis of pileus hymeniform or subhymeniform consisting of broom cells which are clavate to vesiculose and then often broader than high, sometimes sinuate-tripartite or shallowly forked above, 12-28 X 7-17g, main body either all hyaline or in some specimens at times few, at times many cells melleous to brown, thin-walled to slightly thick-walled (walls up to 1 , thick, setulae subdivergent or basically erect and apical, verruciform and then 1-2s, high, or else elongate, simple or forked, conic to (more often) rod-shaped and then 1-6 X 1-2 p, hyaline, golden melleous or brown.

On bark and on small sticks and even dead fallen leaves of dicotyledonous trees, gregarious.

MATERIAL STUDIED PANAMA. C. Z.: Barro Colorado Island, 11 Jul 1952, laartini & IVelden 7606 (IA), type, (F) ISOTYPE, 1,5 Aug 1952, 8406 (IA, F), PARATYPE. BOLIVIA. Beni: Vaca Diez; Guayaramerin, 7 Mar 1956, Singer B 1666 (LIL).

ILLUSTRATION. Singer (1965) p 1 1, fig 57.

205. Marasmius tenuissimus (Junghuhn) Singer, comb nov Fig 87. Agaricus tenuissimus Junghuhn, Verhandlungen, Bataviasche Genotenschaft Kunsten

Wetenschaften, Verh. 17 (84). 1839, non Schweinitz, nec Weinmann. TYPE. Junghuhn, from Indonesia. Pileus dirty ochraceous to cinnamon brownish ("Arab" M&P) when dried, when

fresh varying between rusty brown ("Ginger, Kaiser br" or paler) and light grayish fuscous, with minute brown punctations under a strong lens, rugulose uneven almost all over or at least slightly rugulose at margin when dry, smooth otherwise or becoming sulcate on margin, convex, conchate, often sinuate on inner side 7-14 mm broad. Lamellae white to cream pallid, fresh often becoming concolorous with the pileus in the herbarium, distant, (3-6 through-lamellae, sometimes depending on the age of the

carpophores) with entire, not discolorous edges, often some forked, not or only distantly intervenose when mature, narrow, some iamellulae intermixed or not, laterally concurrent or attenuate-subadnate. Stipe white to deep brown, glabrous or subglabrous or very finely whitish flocculose but glabrescent, curved and often appressed to the lamellae, lateral, sublateral or strongly eccentric, sometimes practically disappearing in age, insititious, 1-2 X 0.1-0.5 mm. Context in pileus white, thin, pallid in interior of

stipe; odor none. Spores (7-)8-10.5 X 3.8-4.8 ,, to ellipsoid-oblong, fewer ellipsoid, smooth, hyaline,

inamyloid. Hymenium: basidia 17-25 X 5.5-7.5ps, 4-spored, basidioles fusoid; cystidia none; cheilocystidia present at least near the margin of the pileus, like the epicuticular broom cells. Hyphae of the pileus-trama hyaline, filamentous to irregularly shaped, the majority filamentous, not gelatinized, with clamp connections, thin-walled, inamyloid. Hyphae of the stipe mostly thick-walled but also many thin-walled ones

present, the latter inamyloid, the former weakly but undoubtedly pseudoamyloid, the thick-walled ones most numerous in the rind of the stipe and there tending to become melleous-ochraceous (intraparietal pigment). Cortical layer: epicutis of the pileus hymeniform, consisting of broom cells which are very irregular in shape and size, mostly with a main body 7-21(-35) X 5-16.5,which is hyaline or light golden ochraceous or entirely ochraceous, with thin to slightly thickened wall (up to 1 U), pedicellate, often ventricose or vesiculose, but also often broadened and bi- to tri-lobed, etc; setulae 1-7 X 0.8-3p, hyaline, light golden melleous or brown, mostly golden melleous,



Marasmius 259

some simple, some forked-crested, apical but often slightly divergent. On dead and living twigs and bark of dicotyledonous trees but also passing over

onto dicotyledonous leaves. MATERIAL STUDIED. BRAZIL. Guanabara, Jardim Botanico, Rio de Janeiro, 28 Jan

1961, Singer C 3137 (BAFC). BOLIVIA. Beni, Vaca Diez, Guyaramermn, 14 Mar 1956, Singer B 1975 (LIL). INDONESIA. near Kebokoening, May (L), type.

This species may be confused with Marasmius neosessilis if the pseudoamyloidity of the pileus-hyphae is too slow to be discovered immediately.

206. Marasmius spaniophyllus Berkeley, Lond. Jour. Bot. 2: 631. 1843. Marasmius iguazuensis Singer in Singer & Digilio, Lilloa 25: 189. 1952 (a variety).

206a. Marasmius spaniophyllus var spaniophyllus TYPE. Gardner, from Brazil. Pileus becoming brown, glabrous, sulcate, reniform, convex, 5-7 mm. Lamellae

yellowish pallid or becoming so, 3-6, with smooth interlamellar spaces, ventricose.

Stipe pulverulent, concolorous (beneath covering ?), short and lateral.

Spores 10.5-12.3 X 5.3-6u, smooth, hyaline, inamyloid. Hymenium: basidioles fusoid; cystidia none; cheilocystidia like epicuticular broom cells but mostly hyaline. Hyphae with clamp connections, inamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells which are globose or subglobose, clavate-vesiculose, varying from 14.5 X 141to 17.5-20 X 9.8-13.8u, the upper portion covered with somewhat diverging but often only apical and subapical short verrucose (projecting 1-1.5u) to elongate diverticula or setulae, these subhyaline to deep melleous, often secondarily diverticulate or forked, wall of main body hyaline to deep melleous and thin to up to 1.5/l thick.

On dead woody sticks, erumpent from bark, gregarious. MATERIAL STUDIED. BRAZIL. Goyaz: Natividade, Nov 1839, Gardner (K) type. ILLUSTRATION. Singer (1958a) fig 10.

206b. Marasmius spaniophyllus var iguazuensis (Singer in Singer & Digilio) Singer, Sydowia 18: 210. 1965.

Marasmiius iguazuensis Singer in Singer & Digilio, Lilloa 25: 189, 1952. TYPE. Singer & Digilio M84, from Argentina. Pileus young white, then becoming very pale fuscous (sayal br. M&P) to grayish

fuscous ("cracker" to "adobe"), dried reaching "Bombay," glabrous, subsulcate to-

deeply sulcate, reniform, convex, eventually applanate, often umbilicate, 4-7 mm broad. Lamellae concolorous with pileus or remaining white a little longer than the

pileus, distant, 5-9, mostly about 6, intermixed with 0-4 lamellulae or forked ones, with the interlamellar spaces often rugulose, edges slightly white-fringed, adnate but in

age separating and forming a very incomplete collarium, rarely not separating, often almost concurrent (stipe so reduced!), rather broad (1 mm). Stipe concolorous with the pileus, pruinose, becoming more umber brown under the pruina when handled and dried especially at base, curved to appressed, rarely straight and merely oblique, equal or tapering downwards, eccentric, then strongly eccentric to lateral, insititious, 0.5-3.5 X 0.2-0.6 mm. Context very thin, inodorous.

Spores 9-11 X 4.3-6.5u, ellipsoid to ellipsoid-oblong, smooth, hyaline, inamyloid. Hymenium 22-39 X 7-8.3g, 4-spored; cystidia none; cheilocystidia 13-28 X 7-21u, thin-walled, hyaline to melleous, rarely thick-walled, vesiculose-pedicellate, excrescen-




cies setulose-diverticulate, often forked or nodulose to subramose, 2.5-4.7iLlong. Hyphae hyaline, inamyloid, in stipe partly pseudomyloid with clamp connections. Cortical layers: epicutis of pileus hymeniform but sometimes fragmentary and then leaving an uppermost layer of irregular hypodermial filamentous, sometimes rough- walled hyphae, hymeniform covering consisting of broom cells, hyaline, later melleous with a hyaline minority interspersed, thin-walled, later many slightly thick-walled, vesiculose to globose-pedicellate, beset with small verruculose setulae or excrescencies as in the cheilocystidia, main body 9-20u1diameter, setulae reaching 5p long. Covering of stipe with similabroom cells.

On dead stems of various herbs and small vines and twigs of trees (dicotyledonous species) also on bark of dicotyledonous trees, always gregarious.

MATERIAL STUDIED. VENEZUELA. Sucre, NW of Irapa, 8 Jul 1972, Diimont et al VE 4259 (NY). ARGENTINA. Misiones: Iguazu; Cataratas, 26 Nov 1949, Singer & Digilio M 84 (LIL), type, 22 Nov 1949, M 23 (LIL); Refugio "Yaguarete" 15 Apr 1957, Singer M 1012 (LIL).

ILLUSTRATION: Singer (1965), pl 11, fig 56.

207. Marasmius sejunctus Singer, sp nov Fig 88. TYPE: Singer B 980, from Bolivia. Pileo pallide salmoneo vel roseolo-ochraceo, 2-3.5 mm lato; lamellis albis vel

cremeo-albis, subconfertis in maturis, haud fortiter intervenosis; stipite albo vel salmoneo vel umbrino-fusco, admodum excentrico vel sublaterali, exiguo, insititio.

Sporis 7-10 X 3-6s,, magnitudine formaque variabilibus; cystidiolis ad lateral lamellarum praesentibus. Ad ramulos ligneos et ramos delapsos arborum dicotyledonearum. Typus in F conservatus est.

Pileus pale salmon to pinkish ochraceous (9 C 5 to 9 H 6, M&P), often becoming reddish brown ("tortoise" M&P) in the herbarium, smooth or with very indistinctly sulcate margin, macroscopically glabrous but under a lens appearing subtomentose or beset with tiny reddish brown punctations, convex, later more repand, reniform (the stipe passing through the sinus), with a shallow umbilicus or depression above the insertion of the stipe when mature, eventually slightly lobed or wavy, 2-3.5 mm broad. Lamellae white to cream white, with the edges concolorous with the sides, at first only 3-4 through-lamellae, later 8-10, irregularly or didymously inserted with lamellulae, subclose when mature, with occasional and few anastomoses or intervenations in some of the mature carpophores but not consistently nor strongly intervenose, not collariate, narrow to rather broad, adnexed to adnate. Stipe white or concolorous with the pileus, or umber-fuscous, curved and/or recurved, very eccentric to sublateral, often appressed to the underside of the pileus, very finely flocculose to pruinose but glabrescent, often somewhat compressed, hollow, insititious, 0.5-1 X 0.1-0.2 mm. Context very thin, white, or whitish when fresh, inodorous.

Spores 7-10 X 3-6u,, mostly about 8-9.5 X 3.5-4.8A, variable in size and shape from as elongated as 10 X 3, to as broad as 8.5 X 4.5p, i e subfusoid-oblong to ellipsoid, with very thin eventually firm wall, smooth, hyaline, inamyloid. Hymenium: basidia (13-)16-24 X 6.5-8,u, 4-spored; basidioles fusoid, fusoid-obtuse or sometimes the majority clavate-subcylindrical i e equally variable as the spores; cystidia present as cystidioles, these neither deep-rooting nor strongly projecting, numerous, especially near the margin of the pileus, 15-18 X 5-7u, clavate to ventricose, obtuse, with firm to slightly thickened wall (wall 0.5-0.6p thick), entire, stramineous to stramineous-hyaline, or hyaline; cheilocystidia like the epicuticular broom cells but mostly hyaline or else



Marasmius 261

pale golden melleous here and there, never with long flagella-like appendages, diverticula either indistinct or setulose and then only 1-3p high. Hyphae of the pileus-trama and the regular to irregular hymenophoral trama hyaline, thin-walled, interwoven, not gel- atininzed, dense, with clamp connections but some major septa secondary and without clamps, inamyloid. Hyphae of the stipe subparallel, more or less distinctly dimorphic, with the thinner-walled ones (wall 0.3-0.6,6) not or extremely slowly and weakly pseudoamyloid, whereas the thick-walled ones (wall 0.7-1.5u thick) subhyaline to

yellowish in KOH and weakly but distinctly pseudoamyloid, more strongly so when seen in thick layers, all hyphae of variable diameter (2-122) but elongated. Cortical layers: epicutis of pileus hymeniform or subhymeniform, consisting of broom cells and similar cells which are devoid of setulae but often with a finger-like outgrowth, the latter always in the minority, e g 22 X 6g, with firm to slightly thickened wall, with acute to rounded tip, fusoid, cylindric-subclavate, more rarely subisodiametric, mostly pale golden melleous to melleous; the broom cells with a main body, 11-20 X 5-7(-17) -, subglobose, clavate or fusoid to ventricose, hyaline to pale stramineous or pale golden melleous, mostly thin-walled but some with slightly thickened wall, not in-

crusted, setulae extremely variable between short and broad and needle- to flagella-like and thin and long (3-8 X 0.2-0.4 ), the short ones coarsely verrucose to conical, obtuse or acute, covering the apical region (sometimes down to the middle) of the cell but more or less diverging, 1-3 X 0.4-1.5 ,), forked or simple, the thin ones mostly hyaline, the short ones often light golden melleous or goldbrown, with all transitions between the Siccus- and the Rotalis-type. Covering of the stipe: the outermost hyphae of the

stipe-rind on their outer side beset with setula-like bodies (like the setulae of the broom cells but with strongly reduced main body), these setulae often in bunches, mostly acute, 2-3.5p long.

On woody twigs and branches fallen off dicotyledonous trees in wet forests, gregarious.

MATERIAL STUDIES. VENEZUELA. Dto Federal, Mount La Naiguata, 25 Jun 1971, Dumont et al VE 725 (NY). BOLIVIA. La Paz, Nor-Yungas, gorge near San Jeronimo, 1900 m alt, 7 Feb 1956, Singer B 980 (F), type.

208. Marasmius neosessilis Singer, Mycologia 50: 103. 1958

208a. Marasmius neosessilis var neosessilis TYPE. Singer & Digilio M 57, from Argentina. Pileus at first almost white but soon "Apricot" to salmon flesh-color (9 F 7),

pinkish cinnamon (11 B 8) or sordid salmon (12 B 10), glabrous cross-veined and

radially deeply sulcate-grooved when mature, reniform in outline, 1-19 mm broad. Lamellae white, distant (3-4, in larger caps ? 6), with numerous veins on intralamellar surfaces but veins rarely reaching neighboring lamellae, intermixed with about 1-5 lamellulae, not intervenose and not intermixed in small carpophores, narrow, adnate or in smaller caps concurrent. Stipe absent in small carpophores even at maturity, present although rudimentary and appressed to both substratum and carpophore, apex whitish, below pallid to pale fuscous, later fuliginous, slightly subtomentose, insititious, 0.5-2 X 0.1-1 mm. Context thin. white, inodorous.

Spores 7-10 X 3.3-4.8,, ellipsoid to oblong-subfusoid, smooth, hyaline, inamyloid. Hymenium: basidioles fusiform; cystidia none, cheilocystidia like epicuticular broom cells but all hyaline or merely with pale golden yellow setulae, absent over part of the edges. Hyphae with clamp connections, in the pileus thin-walled, later becom-




ing mixed thin-walled and moderately thick-walled, some pseudoamyloid, some inamyloid (trama heteromerous as far as cell wall structure is concerned, not morphologically); those of stipe thin- to thick-walled and pseudoamyloid, reminding one of the hairs of Crinipellis intermixed only with a few thin-walled connective hyphae, all hyphae with clamp connections. Cortical layers: epicutis of pileus basically cellular and hymeniform, cells granular to diverticulate in upper part, some cells brown because of the ornamentations being so pigmented, in young material generally with a very slight and often indistinct yellowish pigment incrustation, main body hyaline, in some cells

golden-melleous, mostly vesiculose, 9-25 X 6.5-18,u; setulae mostly golden-yellow to

golden-melleous, rodshaped or spinulose, some forked, 1.2-9 X 0.5-1.5l,; covering of the stipe consisting of similar broom cells.

On various dead twigs and vines, etc, rotting in the subtropical and tropical forest.

MATERIAL STUDIED. ECUADOR. Napo, Shushufindi, 13 May 1973, Singer B 7415 (F). BOLIVIA. Beni: Vaca Diez; Guayaramerin, 16 Mar 1956, Singer B 2047 (LIL). ARGENTINA. Misiones: Iguaziu, Cataratas, Singer & Digilio M 57 (LIL), type.

The type is very small, and sessile to subsessile whereas the Bolivian material is 12-19 mm broad and constantly stipitate. The type has only 3-4 through-lamellae whereas the Bolivian material has about six. The Ecuadorian material is intermediate

(pileus 4-10 mm; 2-6 through-lamellae).

208b. Marasmius neosessilis var montepiensis Singer, var nov Fig 89. TYPE. Singer M 8757, from Mexico. Differt pileo griseolo, 9-17 mm lato, lamellis admodum distantibus. Typus in F. This differs from the type variety in the colors of the pileus and in the fact that

the pileus is large but the lamellae are fewer i e relatively more distant; the stipe is

relatively more developed than in the type variety. Pileus pale dirty gray to gray when fresh and wet, usually paler at margin, be-

coming "yellow-beige" on drying or remaining gray ("seaside" M&P), folded-sulcate, often scrobiculate, glabrous, eventually lobed, 9-17 mm broad. Lamellae white, be-

coming cream or pale ochraceous on drying, very distant (4-7 through-lamellae), at first not, later weakly intervenose, very narrow but not veniform, adnexed to adnate.

Stipe fuscous, glabrous, equal, curved, insititious or subinsititious, 0.5-4 X 0.5-0.8 mm; basal mycelium, if present, hyaline and very scanty. Context very thin, inodorous.

Spores 7.5-9 X 4.5-3,u, ellipsoid, smooth, hyaline, inayloid. Hymenium: Basidia 20-27 X 6-8.5y, 4-spored, some thin-walled and alternating with somewhat thick-walled ones which are easily mistaken for cystidia before sterigmata are formed; cystidia none; cheilocystidia not making the entire edge heteromorphous, much like the epicuticular broom cells. Hyphae of the pileus-trama slowly distinctly pseudoamyloid (but some

inamyloid) all with clamp connections, nowhere gelatinized. Cortical layers: epicutis of the pileus hymeniform, consisting of broom cells, these at least at first subhyaline with the main body 13-19 X 3.8-9.5,, ventricose to clavate, setulae 3.4-6E1long, apical but slightly divergent, some simple, some forked; covering of the stipe scattered, con-

sisting of broom cells like those of the epicutis of the pileus and terminal cells of the

hyphae of the rind of the stipe which are ascendant and with or without a septum. On wood of dicotyledonous trees and shrubs in rain forest, gregarious. Mexico to

Ecuador. MATERIAL STUDIED. MEXICO. Veracruz, 3 km from Montepio, 30 Jul 1969, Singer M

8757 (F), type. ECUADOR. Napo, Shushufindi, 13 May 1973, Singer B 7382 (F).



Marasmius 263

This species can easily be confused with M. tenuissimus (no. 205) but differs in the colors, the strongly sulcate pileus and by the presence of pseudoamyloid hyphae in the trama of the pileus.

209. Marasmius pleurotelloides Singer, sp nov Fig 90. TYPE. Singer B 4240, from Brazil. Pileo albo, 2.5 X 2 mm; lamellis albis, distantibus; stipite atrobrunneo, curvato,

1 X 0.1 mm. Sporis 7-7.5 X 3,; cystidiis nullis, sed cystidiolis interdum praesentibus; hyphis pseudoamyloideis, defibulatis; cheilocystidiis et elementis epicuticularibus typi Marasmii rotalis.-Ad ramulos in Brasilia. Typus in BAFC conservatur.

Pileus white in fresh, whitish in dried material, smooth, glabrous, convex, oval-

subcircular, 2.5 X 2 mm. Lamellae white, turning ocherpallid when dried, distant (six to seven through-lamellae), rather broad to broad, broader and narrower lamellae alter-

nating but not intermixed with lamellulae, rounded-adnexed. Stipe dark brown, curved, 1 X 0.1 mm. Context white, thin, inodorous.

Spores (6-)7-7.5 X (2.5-)3,, fusoid, smooth, hyaline, inamyloid. Hymenium: basidia 14-23.5 X 5.2-6.2M, 4-spored; cystidia none but cystidioles sometimes found which are e g 16 X 5 u, subvesiculose, with flexuous pedicels; cheilocystidia like the

epicuticular broom cells. Hyphae of the pileus-trama filamentous, hyaline, without

clamp connections, pseudoamyloid. Cortical layer: epicutis of the pileus subhymeniform, hyaline, main body 14-40 X 8-171, broadly cylindrical to subvesiculose, of the

Rotalis-type, since the setulae cover one third to one half of the cells and are more or less divergent, of very variable size and shape, the apical ones often elongated and

secondarily diverticulated at right angles (almost asterostromelloid), but most simple and only 2-5, long, many broom cells ramose or appendiculate.

On small stems, twigs, etc., in Brazil. MATERIAL STUDIED. BRAZIL. Parai, Estancia Pirelli, 7 Jun 1966, Singer B 4240 (BAFC),

type. This species may be mistaken for a Marasmiellus but the subhymeniform arrange-

ment of the epicuticular broom cells and the fact that no clampless Marasmielli with

pleurotoid habit are known, show that it belongs in Marasmius.

Marasmius sect Alliacei Kuhner, Le Botaniste 25: 60. 1933 (as Alliaceae). TYPE SPECIES. Marasmius alliaceus (Jacq. ex Fr.) Fr. Pileus pigmented or not, with the epicuticular cells in hymeniform of frag-

mented-subhymeniform arrangement, but never in the form of broom cells, i.e. smooth or with uneven surface but occasionally sometimes not beset with setulae; spores small to medium sized, rarely long; stipe central, with distinct basal mycelium and mostly with pruinate to velutinous covering; hyphae inamyloid, with clamp connections; on dead wood and leaves.

Key to the Species of Section Alliacei

1. Spores acicular-fusiform, 11-17.5 X 3-3.5 u; cystidia none; stipe scurfy-velutinous. 210. M. perlongispermus.

1. Spores shorter; cystidia present or absent. 2. Pileus green. 211. M. smaragdinus. 2. Pileus not green.

3. Cystidia absent. 4. Pileus gray, tomentose under a lens; spores 5.5-8.8 X 2.5-3.8 , on dicotyledon-

ous wood in Brazil. 212. M. batistae.




4. Pileus not gray but ochraceous brown or pale golden cinnamon etc. 5. Spores 5.3-6 X 2.3-3u; epicutis with fragmentary epicutis layer; on dead

rotting leaves; pileus about 6 mm broad. 213. M. oligocysstis. 5. Spores about 5.5 X 4.3,; epicutis rather continuous, subhymeniform to

hymeniform; on humus and sandy soil; pileus larger. 214. M. aimiara. 3. Cystidia on the sides of the lamellae present.

6. Pileus dirty violet in the center, brown towards the margin; diameter 20-60 mm; lamellae distant; cystidia thin-walled. 215. A. ionides.

6. Pileus without violet tinges, often smaller, or else lamellae close or cystidia thick-walled. 7. Spores over 4 broad.

8. Cystidia with firm but not thick-wall; spores up to 5.5 p broad; pileus rufescent, setoid dermatocystidia on the pileus absent; on tree trunks.

216. A. latirsculosperlmus. 8. Hymenialcystidia and dermatocystidia thick-walled; spores still broader;

pileus fuliginous with isabelline disc; setoid dermatocystidia on the pileus present; on clay soil. 217. ,MI. nmutrillianus.

7. Spores less than 4 broad. 218. Al. hinnuii leiformis.

210. Marasmius perlongispermus Singer, sp nov Fig 91. TYPE. Singer B 6489, from Colombia. Pileo atrobrunneo, margine pallidiore, hygrophano, 30-31 mm lato; lamellis brun-

neis, angustis, subconfertis vel subdistantibus; stipite atrobrunneo, apice pallidiore, frequenter subexcentrico; sapore fortiter alliaceo. Sporis 11-17.5 X 3-3.5y; cystidiis nullis; hyphis inamyloideis. Ad lignum in silvis montanis columbianis. Typus in F conservatus est.

Pileus deep brown with paler margin, dried "auburn" (M & P), hygrophanous, fades to buff when fresh, sulculate in the marginal portion, glabrous, convex soon with

depressed center, eventually subapplanate, earlier flattened on the larger side if eccen-

tric, umbilicate or subumbilicate, 30-31 mm broad. Lamellae brown, narrow, subclose to subdistant, not intervenose, tridymous, adnexed to subdecurrent. Stipe deep brown with paler apex, entirely scurfy to pubescent-velutinous or pruinate, slightly tapering towards the base, often more or less eccentric, 20-21 X 1-1.2 mm; basal

mycelium scanty, short-strigose, often visible only with a hand-lens. Context thin; odor weak, but taste strongly alliaceous.

Spores 11-17.5 X 3-3.5 , acicular-fusiform, smooth, hyaline, inamyloid, with

many small internal oil droplets. Hymenium: basidia 24 X 7p1, clavate, 4-spored; cystidia none; cheilocystidia like the cells of the epicutis of the pileus but smaller:

17-18(-40) X 7-9u, hyaline or subhyaline, hyaline or subhyaline, smooth, thin-walled.

Hyphae of the pileus trama rather loosely arranged, but not or scarcely gelatinized, subhyaline, radially arranged, subparallel, with firm walls, 2-7 mm broad, with clamp connections and inamyloid; hymenophoral trama regular but its hyphae somewhat inter-

woven, hyaline to subhyaline, not or scarcely gelatinized, elongated, with firm walls

up to 0.6u in diameter, 3-9ubroad. Cortical layers: epicutis of the pileus subhymeniform, consisting of smooth cells, 16-31 X 10-21u, some with wavy-sinuate outline, or

obliquely elongated, with or without pedicel, fuscous; covering of the stipe, a dense

layer of erect hairs with firm walls up to 0.5 u thick, these 50-160 X 6-9(-10)A, hya-

line, inamyloid, irregularly cylindrical with occasional slight swellings, apex often dif- form or crooked.

On wood in tropical montane cloud forest, subparamo. Colombia. MATERIAL STUDIED. COLOMBIA. Cauca: road to sulphur mine at Purace at 2700 m

alt, 30 Apr 1968,SingerB 6489 (F), type.



Marasmius 265

211. Marasmius smaragdinus (Berkeley) Singer, Sydowia 12: 75. 1959. Agaricus smaragdinus Berkeley, Hook. Jour. Bot. 8: 131. 1856. Omphalia smnaragdina (Berkeley) Saccardo, Sylloge Fungorum 5: 322. 1887. TYPE. Spruce, from Brazil. Pileus green or deep olive, almost black at the margin and along the striae ac-

cording to Dennis, glabrous and smooth except for the striation, convex, shallowly or deeply umbilicate, 10-14 mm. broad. Lamellae cream colored or green, tridymous, subdistant or distant, rather narrow or rather broad, somewhat intervenose when mature and dried, decurrent. Spore print white. Stipe green or avellaneous with a white bloom, equal, hollow, 25 X 0.5-2 mm. Context cream under the cuticule of the

pileus, avellaneous in the stipe. Spores 4-4x5 X 3,s, elliptical in outline, inamyloid. Hymenium: basidia 25 X

4-5p, 4-spored, cystidia, none seen. Hyphae hyaline, non-gelantinized, inamyloid, with clamp connections those of the stipe parallel, slightly thick-walled and submel- leous. Cortical layers: epicutis of pileus hymeniform, consisting of broadly clavate smooth elements' hypodermium seemingly pseudoamyloid but probably not (a combination effect of pigment color and staining); pigment dissolved in epicutis, incrusting in hypodermium (at least in the dried material studied).

On woody humus and rotten twigs and branches in tropical forest. MATERIAL STUDIED. TRINIDAD. Siparia, 3 Nov 1949, Dennis 274 (K). BRAZIL. Ama-

zonas: Panure (i e Ipanure = Sao Jeronimo), Spruce (K) type. Dennis considers Wright 41 from Cuba (Marasmius viridifuscus Berkeley & Cur-

tis) to be the same as the species described above. The type material deposited in the Curtis Herbarium (FH) did not yield enough data to be certain about this synonymy.

ILLUSTRATIONS. Dennis (1952) fig 24; (1970) p1 7 fig 24

212. Marasmius batistae Singer, Sydowia 18: 352. 1965. Fig 92. TYPE. Singer B 3436, from Brazil. Pileus gray, glabrous, under a lens subtomentose, sulcate, convex-subapplanate,

with slightly depressed center, 18 mm broad. Lamellae whitish, distant, broad, adnate. Stipe white, finely subtomentose-pubescent, smooth, equal or slightly tapering towards the apex, 17 X 1.5-2 mm; basal mycelium substrigose, white. Context white, unchanging, inodorous.

Spores 5.5-8.8 X 2.5-3.8iu,oblong to fusoid, thin-walled, smooth, hyaline, in-

amyloid, with a globose central oil drop. Hymenium: basidia 18-25 X 4.3-5,u; basidioles fusoid, acute, some succineous-brown incrusted (incrustation sometimes dissolved in Melzer's reagent to form brown droplets); cystidia and cheilocystidia none. Hyphae of the irregular hymenophoral trama running in denser strands of parallel hyphae, but between these strands seemingly somewhat gelatinized, variable in size and shape, some thin-walled, others slightly thick-walled and appearing glassy in alkali mounts; subhymenium of more axially arranged filamentous hyphae which are also denser; all hyphae inamyloid, with clamp connections. Cortical layers: epicutis of the pileus consisting of a single layer of almost to quite hymeniformly arranged swollen hyphal ends, these broad-cylindric-subvesiculose or vesiculose with or without a mucro or ventricose- mucronate, or more cystidiform, and then ampullaceous, 23-48 X 16-20/, all hyaline in alkali, inamyloid; there are also, but scattered, some larger dermatocystidia on the

pileus, much like those encountered on the stipe. Covering of the stipe consisting of hairlike dermatocystidia 88-100 X 6-12.5, cylindric or cylindric-subampullaceous, hyaline.




On dicotyledonous dead wood in tropical rain forest. MATERIAL STUDIED. BRAZIL. Pernambuco, Mata de Sao Joao, 16 Jul 1960, Singer B

3436 (BAFC), type. The color and anatomy of this fungus makes one think of Hydropus or Cypto-

trama but the entirety of characters place it in this section of Marasmius.

213. Marasmius oligocystis Singer, Sydowia 12: 75. 1959. TYPE. Singer B 403, from Brazil. Pileus ochraceous brown with whitish central disc, not viscid but possibly hygro-

phanous, glabrous but very slightly pruinate, under a lens smooth, convex, 6 mm broad. Lamellae whitish, becoming chestnut when dried, narrow, close to subclose, subfree. Stipe cinnamon, ochraceous brown to fuscous ochraceous at the apex, dis-

tinctly pruinate all over, not institious, filiform, equal, 17 X 0.8 mm.; basal mycelium white. Context almost none, inodorous.

Spores 5.3-6 X 2.3-3 j, hyaline, smooth, inamyloid, guttiform or cylindrical- elongated rarely reaching 6.5 , in length. Hymenium: basidia 15.5-18(-20) X 4.3A, clavate and relatively short, 4-spored; basidioles fusoid; cystidia on edges of lamellae very conspicuous but scattered, 20-30 X 9-15.7 p saccate to vesiculose-clavate, hyaline or subhyaline, smooth. Hyphae: hymenophoral trama regular, hyaline or subhyaline: hyphae of stipe parallel; all hyphae with clamp connections, inamyloid. Cortical layers: epicutis of pileus consisting of smooth cells 15-26 X 7-9 u, repent, ascendant or erect, ventricose to clavate or globose-pedicellate, cystid;oid, forming a fragmentary and interrupted layer, neither clearly hymeniform nor in palisade in adult material, covering the hypodermium; hypodermium, a cutis consisting of repent hyphae which are cinnamon to pale chestnut, with very fine and not very conspicuous pigment incrustations; pruina of the stipe made up of dermatocystidia which are most frequently subcapitate as in M. chordalis and M. elongatipes but also sometimes cylindric, ventricose, or ampullaceous, 20-30 X 5-8 p.

On decaying leaves in tropical coastal forest. MATERIAL STUDIED. BRAZIL. Parana, Near Paranagua, Sep 1952, Singer B 403 (F),

type. ILLUSTRATION. Singer (1965) fig 3.

214. Marasmius aimara Singer, Sydowia 18: 333. 1965. TYPE. Singer B 1134, from Bolivia. Pileus pale golden cinnamon ("golden wheat"), not shining, irregularly convex to

campanulate-subumbonate but obtuse, 19-20 mm broad. Lamellae cream gray("tan- aura"), subclose, broad, adnate. Stipe concolorous with the pileus, equal solid, slightly minutely fibrillose becoming subglabrous, 24-35 X 2-2.5 mm. Context rather thick, pale ochraceous cream, inodorous.

Spores 5.4-5.6 X 4.3 , without a suprahilar depression, smooth and hyaline, inamyloid. Hymenium: basidia 16-21 X 5-6u, 4-spored; cystidia, none. Hyphae with

clamp connections, inamyloid. Covering layers: epicutis of pileus subhymeniform, consisting of subisodiametrical cells which vary from vesiculose to globose, with some few cylindrical ones mixed in, all or most strongly incrusted, yellowish, some with one or two diverticula.

On sandy soil mixed with humus, margin of mango plantation. Gregarious, not cespitose.

MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Charobamba, 13 Feb 1956, Singer B 1134 (LIL) type.



Marasmius 267

215. Marasmius ionides Patouillard in Duss, Enumeration Champignons de Guadeloupe, p 45, 1903.

TYPE. Duss 539, from Guadeloupe. Pileus dirty violet in the center, brown towards the margin, opaque, glabrous,

under a lens pulverulent, convex then applanate, 20-60 mm broad. Lamellae yellowish white, distant, narrow, intermixed, adnate. Stipe rufescent, pilose, central, equal, 20-40 X 1-3 mm. Context thin.

Spores 5.5-7.2 X 3.2-3.8 , (10 X 3 j, Patouillard), ovoid, with or without suprahilar depression, smooth, hyaline, inamyloid, Hymenium: basidia about 30 ,llong, 4-spored; cystidia 30-50 X 12-16 ,, subvesiculose-elongated, hyaline, thin-walled. Hyphae with inamyloid wall. Cortical layers: epicutis of pileus hymeniform, with vesiculose- elongated cells filled with a violet or brown intracellular pigment, these cells smooth, thin-walled, 20-50 X 20-30,'; covering of stipe consisting of hair-like dermatocystidia which are 600-700 X 10-19 A, apparently thick-walled, awl-shaped, hyaline.

Said to grow on soil. MATERIAL STUDIED. GUADELOUPE. Bois des trois Rivieres, Mar. 1902, Duss 539 (FH),

type.

The description is partly borrowed from the description and drawings by Patouillard, but checked on the type specimen.

216. Marasmius latiusculospermus Singer, sp nov Fig 93. TYPE. Guzmrn 1582, from Mexico. Pileo rufescente, siccando coriicolore, 10-15 mm lato; lamellis albis vel albidis,

angustis, confertis; stipite atrobrunneo, pubescente, cavo, 17-40 X 1-1.5 mm. Sporis 8.5-8.8 X 4.5-5.5 ,; cystidiis sparsis; hyphis inamyloideis; stipitis tegumento pilis crasse tunicatis longis consistente. Ad truncos arborum in Mexico. Typus in Herbario Instituti Polytechnici Mexicani conservatus est.

Pileus rufescent (dried "tan"), then "gypsy," "Agate" (M & P), glabrous, slightly to strongly sulcate on the margin, strongly campanulate at first, then convex and soon with depressed center, 10-15 mm broad. Lamellae white or whitish, narrow, close, somewhat intervenouse when quite mature, adnate to subfree from the widened

apex of the stipe. Stipe deep brown all over, excepting the apex of the young car-

pophores which is pallid, distinctly pubescent, hollow, subequal, not instititious, 17-40 X 1-1.5 mm. Context white excepting the dark brown rind of the stipe.

Spores (7.5-)8.5-8.8 X 4.5-5.5 ,, ellipsoid, smooth, hyaline, often at first granular inside, sometimes taking up internally the dissolved pigment of the pileus, inamyloid. Hymenium: basidia 29-36 X 6-7.7, , 4-spored; cystidia scattered, 42-55 X 4.5-8 ,, ventricose below, tapering to a constriction below a more or less subcapitate apex, constriction 2.5-3.5 , across, capitulum 4.5-5 , across, with firm but not thick wall, with occasional oil droplets in the interior and sometimes becoming colored like the spores. Hyphae thin- to firm-walled in the pileus-trama, thick-walled in the stipe, hyaline ex-

cepting the rind of the stipe where they are chestnut brown, all inamyloid, with clamp connections, nowhere gelatinized. Cortical layers: epicutis of the pileus hymeniform consisting of smooth elements 15-24 X 7.5-14.5(-16)., golden, firm-walled (wall to 0.5 , thick), clavate. Covering of the stipe consisting of long hairs of the Crinipellis- type, but inamyloid, all entire, cylindric-filamentous or slightly and gradually tapering to an obtuse tip, sometimes with a socle-like broadened base up to 12 3, wide, with 1-3 , thick wall, 45-105 X 4-7 ,.

On tree trunks in woods with Liquidambar styraciflua. Mexico.




MATERIAL STUDIED. MEXICO. Queretaro, 5 km W of Lobo, 26 Jun 1969, alt 1500 m, Guzmant 1582 (ENCB), type.

This species was found barely outside the neotropical zone proper but its southern limits of distribution are unknown.

217. Marasmius murrillianus Singer, Lilloa 25: 488. 1952 ex Singer, nov nom Gymnopus setulosus Murrill, North American Flora 9: 373. 1916. Collybia setulosa (Murrill) Murrill, Mycologia 8: 219. 1916. Marasmius setulosus (Murrill) Singer, Lilloa 22: 326. 1951, non Murrill (1940). TYPE. W. A. & E. L. Murrill 632, from Jamaica. Characteribus sectionis; cystidiis setiformibus. Typus in NY conservatur. Pileus fuliginous with isabelline disc, dark purplish brown in the herbarium

(A. H. Smith 1938), beset with bristles, uneven, rugose, dry, with undulate slightly incurved concolorous margin, irregularly conical to convex, truncate, 16 mm broad. Lamellae pure white, rather broad, distant, adnate to a collarium (?). Stipe isabelline with a dense coat of brown bristles, cinnamon brown in the herbarium (Smith 1938), hollow, enlarged and radicant at the base as in Oudemansiella radicata, 50 X 2-3 mm. Context tough in the stipe.

Spores 8-10 X 7-9,, globose or subglobose, inamyloid. Hymenium: basidia 4- spored; cystidia 60-80 X 12-20(-28)g, smooth, hyaline with blunt apex and somewhat inflated mid-portion, at the edges of the lamellae not differentiated. Hyphae inamyloid. Cortical layers: epicutis of the pileus, a hymenial palisade consisting of versiform elements (11-)28-36(-45) X 8-12(-15) ,, filled with brown pigment, subglobose to fusoid- ventricose or with acute apex; also setoid dermatocystidia, 50-180 X 8-12(-28) A, gradually tapering to a point, light yellowish brown; covering of the stipe consisting of similar setae as those of the pileus, according to my measurements reaching 500, in length.

On a moist,mossy bank on clay soil in the West Indies. MATERIAL STUDIED. JAMAICA. Cinchona, about 1500 feet alt, between 25 Dec 1908

and 8 Jan 1909, W. A. & E. L. Murrill 632 (NY), type. The description given above is adapted from Murrill's original account for the

macroscopical characters and on A. H. Smith's (1938) account for most of the microscopical details. For further information on the type specimen I am indebted to Joanne Williams, Seattle.

ILLUSTRATION. Smith (1938), fig 3a, b, c, g.

218. Marasmius hinnuleiformis Murrill, North American Flora 9: 263. 1915. Fig 94. TYPE. Murrill 163, from Mexico. Pileus "beige" with "oakwood" (M&P) center according to sketch, but apparently

often partly whitish at first, smooth or becoming plicate on drying, convex becoming flattened with convex center, thus appearing obtusely umbonate, 5-20 mm broad. Lamellae white, under lens dotted by the cystidia, narrow, close to subclose, not intervenose, adnexed to adnate, sometimes collariately separating from the apex of the stipe. Stipe yellowish brown when fresh, minutely pruinate from the dermatocystidia, equal or subequal, 30-50 X 1-2 mm; basal mycelium abundant, whitish. Context thin.

Spores 6-7.5 X 2.5-3.3,, oblong, subfusoid, smooth, hyaline, inamyloid. Hy- menium: basidia 20 X 4.3 1, 4-spored; cystidia 36-49 X 9.3-16.3 A, fusoid-ampullaceous, thick-walled (wall 1.2-2.2 , thick), mostly entire but sometimes with 1-3 inconspicuous, small, obtuse diverticula at the apex, opalescent, hyaline. Hyphae inamyloid, not gel-



Marasmius 269

atinized. Cortical layers: epicutis of the pileus hymeniform, consisting of smooth, clavate or ventricose to subvesiculose elements which are often apically mucronate, with 0.4-1 , thick wall, interspersed with dermatocystidia 33-100 X 7.7-8.7 M, ampullaceous, the apex of the "neck" often subcapitulate but otherwise smooth and obtuse and 2.2-4.5 , in diameter, subhyaline or hyaline, inamyloid; covering of the stipe consisting of similar dermatocystidia.

On the ground (?) in virgin forest, Mexico. MATERIAL STUDIED. MEXICO. Veracruz, Jalapa, 12-20 Dec 1909, W. A. & Edna L.

Murrill 163 (NY), type. Marasmius subagricola (Murrill) Singer, comb nov (Gymnopus subagricola Murrill,

Mycologia 33: 439. 1941) is very similar but has very scanty basal mycelium, thinner- walled cystidia, subdistant lamellae. It was described from Gainesville, Florida, U.S.A. but does not seem to occur in the neotropics.

Marasmius sect Globulares Kihner, Le Botaniste 25: 60. 1933 (as Globularinae). TYPE SPECIES. Marasmius wynnei Berkeley & Broome. Pileus pigmented or not, with the epicuticular cells as in the preceding section

from which this section differs mainly in the strongly pseudoamyloid trama.

Key to the Species of Section Globulares

1. Spores large: 10.3-16 6 long, at least most spores longer than 1A. 2. Pileus white (cf also "16" below). M. plunieri (see p 283). 2. Pileus distinctly pigmented.

3. Spores about 16 p pileus cinnamon tawny growing from white rhizomorphs in Brazil. 219. M. viegasii.

3. Spores (6-)10.5-11.5 X 3-4 u; pileus mouse gray. Colombia. 220. M. myocephalus. 1. Spores smaller, up to 10.5 plong.

4. Pileus wax yellow or even brighter yellow; lamellae also yellow, intervenose. 221. M. rhyssophyllus.

4. Pileus not so colored or lamellae not intervenose. 5. Pileus deep spadiceous-otter brown or deep purple; lamellae intervenose.

6. Spores 7-9 X 4.5-4.8,. 222. M. ditopotrama. 6. Spores 5.7-6.2 X 3.5-3.8 u. 223. M. poromnycenoides.

5. Pileus some other color or else lamellae not intervenose. 7. Cystidia none on the sides of the lamellae.

8. Growing in open places, directly on the earth or on rotten herbaceous plant material, rarely on dung or in water puddles, sometimes in parks and plantations; lamellae moderately distant to rather distant, rather broad to very broad, often somewhat anastomosing but not regularly or strongly intervenose-poroid, margin of pileus subsmooth at maturity. 9. Stipe stuffed; not tending to gray.

10. Introduced species; spores (7-)8-10 X 4-5.8 ,; odor of HCN.. 224. M. oreades.

10. Native neotropical species; spores 5.5-7.2 X 2.8-4.3p; (see also "I1" below). 225. M. cibarius.

9. Stipe very early tubulose and/or tending to grayish. 11. Pileus and stipe tending to yellowish (see "12" below). 11. Pileus and stipe tending to grayish. 226. M. albogriseus.

8. Growing on forest and plantation litter, mostly dead foliage, buried and rotten wood etc, in dense subtropical and tropical marginal forest, shady plantations, etc; not combining the characters indicated above. 12. Lamellae not intervenose; growing on foliage under trees in forests,

plantations, etc. 13. Pileus white with whitish to pale cinnamon or stramineous center

and whitish margin. 14. South American species with spores up to 3.5, broad and

merely sulculate margin and umbilicate center. 227. M. niveus.




14. North American species with spores >3.5, broad and mostly merely corrugated, ?umbonate center. 228. M. strictipes.

13. Pileus not so colored. 229. M. heliomyces. (compare also M. tortipes, p 284).

12. Lamellae weakly to strongly intervenose in mature specimens or fungus not growing on foliage but among grasses and herbs in semi- open places.

15. Lamellae weakly intervenose, broad; growing among grasses and herbs in semi-open places on earth; pileus transparently striate when wet, smooth when dry. 230. M. pseudocollinus.

15. Lamellae strongly intervenose when quite mature, narrow to broad; growing on wood or leaves in the forest, semi-arid woods and plantations; pileus mostly sulcate or pustulate-scrobiculate when dry.

231. M. cohortalis. 7. Cystidia present on the sides of the lamellae.

16. Spores 8.1-20.2 long; pileus colored like M. oreades: lamellae distant; in marginal subtropical forest in Argentina. 232. M. riparius.

16. Spores up to 8.5 p long and/or with different color. 17. Spores 14 u (see "2" above). 17. Spores 8.5 M.

18. Pileus white to lead color; stipe subdiaphanous (see "12" above). 18. Pileus deep cinnamon to ochraceous brownish in the center and on

the striae, paler between the striae, dried more uniformly brown in center, paler on margin, stipe not diaphanous. 233. A'. silvicola.

219. Marasmius viegasii Singer in Viegas, Bol. Superintendencia Servicio do Cafe 32(369): 14. 1957.

TYPE. Viegas 7800, from Brazil. Pileus cinnamon tawny (13 C 11), glabrous, furrowed-sulcate almost to the cen-

ter, at first globose, then becoming convex, gradually flattening eventually more or less depressed or concave, sometimes, especially in age, umbonate, 60-70 mm broad. Lamellae concolorous with pileus, broad, (5 mm), very distant, little or not anastomosing, adnexed or adnate. Spore print white. Stipe deeper colored than the pileus, hollow, finrillose-tomentose below in continuation of the basal tomentum, longer than the diameter of the pileus, slightly tapering upwards, also sometimes tapering downwards into the base from a point below the middle, sometimes spirally twisted, at apex glabrous to subglabrous, about 5 mm broad; basal tomentum white arising from white rhizomorphs which are smooth and become brown, eventually blackish.

Spores about 16 X 4,u when fully mature, cylindrical to subfusiform but the inner side subapplanate to slightly concave in profile view, developing (when overmature) a cross-septum and easily germinating on paper, thin-walled, smooth, hyaline, inamyloid. Hymenium: basidia 20-28 X 4-5 ,, 4-spored; cystidia none. Hyphae pseudoamyloid with clamp connections. Cortical layers: epicutis of pileus hymeniform, its elements more or less clavate, pedicellate and often strongly shortened and then subisodiametrical, smooth, hyaline, arising from thinner hyphae which form a trichodermial palisade of which the clavate cells are the terminal elements; hypodermium, beneath this palisade, forming a cutis. Setae none.

On roots of Coffea arabica. MATERIAL STUDIED. BRAZIL. Sgo Paulo: Campinas, from laboratory culture made on

basis of rhizomorphs of a dead coffee tree of Fazenda Baixadao, Ribeirao Preto, 5 Aug 1948, A. P. Viegas 7800 (IACM), type.

Apparently the rhizomorphs are common in Sao Paulo, Parana and Minas Gerais. The carpophores, on the other hand, are obviously not common or else the species had not been described as new at such a late date.

This is the organism which causes the root rot of coffee in Brazil ("Pfahlwurzel-



Marasmius 271

f'iule," "podridao das raizes") or contributes to it being found regularly in form of rhizomorphs on the roots of affected trees. The phenomenon was formerly attributed to nematodes (Ihering, Noack) and was fully clarified later (Viegas, 1957). About the history, importance, symptomatology, etiology, pathogenicity, life history of the fungus and control of the "disease," see Viegas.


220. Marasmius myocephalus Singer, sp nov Fig 95. TYPE. Singer B 6152, from Colombia. Pileo murino, + 54 mm lato; lamellis pallidis, confertis, haud intervenosis, adnatis; stip-

ite castaneo; odore nullo. Sporis 10.5-11.5 X 3-4 1; hyphis pseudoamyloideis; dermatocystidiis stipitis 8-60 X 2-10 ,. Ad folia et ligna putrescentia in Columbia. Typus in F conservatus est.

Pileus mouse gray, center becoming brownish ochraceous when dried, rugose, not viscid, with sulcate margin, convex with soon applanate or slightly depressed center, about 54 mm broad. Lamellae pallid, with entire, pallid edge, narrow to medium broad, close, not intervenose, adnate. Stipe chestnut, subglabrous to pruinate, hollow, equal, about 95 X 5 mm; basal mycelium dirty pallid. Context inodorous.

Spores (few seen) (6-)10.5-11.5 X 3-4 1, cylindric-fusoid, smooth, hyaline, inamyloid. Hymenium: cheilocystidia like epicuticular cells, often more irregular or narrower. Hyphae strongly pseudoamyloid, in pileus hyaline, hymenophoral trama regular, in stipe parallel with each other, yellow in KOH. Cortical layers: epicutis of the pileus hymeniform, consisting of a layer of smooth, vesiculose, often pedicellate cells 15-25 X 10-24 , with 0.2-0.4 thick wall; covering layer of stipe consisting of dermatocystidia 8-60 X 2-10 A, ascendant to erect, versiform, hyphous to ampullaceous, ventricose, clavate or irregularly cylindrical, often with constrictions, many forked or even branched, all hyaline with firm wall.

Attached to leaves and rotting wood in the litter of the tropical montane forest. Co- lombia.

MATERIAL STUDIED. COLOMBIA. Valle, Municipalidad Cali, Saladito, 15 Apr 1968, Singer B 6152 (F), type.

221. Marasmius rhyssophyllus Montagne in Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 294. 1869 (sensu Patouillard).

Dictyoploca ryssophylla (Montagne in Berkeley & Curtis) Baker & Dale, Mycol. Pap. Commonwealth Mcyol. Inst. 33: 86. 1951, pl, III, d, dl.

TYPE. Wright, from Cuba (not seen); Pileus fresh wax yellow, 25 mm broad. Lamellae yellow, medium broad or rather nar-

row, very strongly intervenose to merely distinctly anastomosing, distant. Stipe concolorous or white, partly chestnut-black when mature, 25 X 2 mm, base strigose.

Spores 7-8.3 X 3.3-3.8 , smooth, hyaline, inamyloid. Hymenium: cystidia none seen, apparently absent. Hyphae pseudoamyloid because of strongly pseudoamyloid walls of numerous thick-walled hyphae in both pileus and stipe, aside from them also inamyloid to strongly pseudoamyloid thin-walled hyphae present in the trama of the pileus and the hymenophore. Cortical layer: epicutis of pileus cellular.

On rotting and dead branches in forest. MATERIAL STUDIED. GUADELOUPE. Duss 1769 (FH), compared with type by Patouillard. This is indicated also from the Guyanas (Saccardo), andTrinidad (Dennis). Dennis

(195 la) gives a complete description of the material from Trinidad but according to him




the pileus surface is "formed of radiating hyphae 5-6 ,wide, their terminal cells often swollen to 30 X 12-13 ,. Trama nonamyloid."

In Dennis's (195 la) interpretation, the species may not belong in Marasinius but probably in Collybia. In Patouillard's interpretation it is a Marasmius of this group. I have not been able to find the type in Montagne's collection (PC).

222. Marasmius ditopotrama Singer, Sydowia 18: 335. 1965. TYPE: Singer B 2068, from Bolivia. Pileus deep spadiceous, otter brown ("piccaninny") in depressions of sulci and on re-

ticulations between sulci, otherwise light brownish gray ("Manila"), not viscid, glabrous, sul-

cate, convex-gibbous with a tendency to become convex around the center because of the uplifting of the margin, about 78 mm broad. Lamellae light brownish gray ("Manila") moder-

ately broad (5 mm), subdistant, intervenose, adnate. Stipe gray, pruinose, somewhat ventri-

cose, hollow when dried, subglabrescent, 80 X 6 mm, only about 4 mm broad at apex and base; basal tomentum short-strigose, gilvous in dried material. Context whitish, inodorous, somewhat toughish, not at all fragile fleshy as in Poromzycena or Hydropus.

Spores 7-9 X 4.5-4.8 , ellipsoid, thin-walled, hyaline, smooth, inamyloid. Hymenium: basidia 31-55 X 5.5-7.5 u, varying from normally long and clavate to strikingly elongate as in Hygrophoraceae, 4-spored; cystidia none; cheilocystidia not confined to extreme edge but also occurring near edge, not striking as they do not project noticeably but distinct and

large, 35-55 X 16-21 ~, "empty," thin-walled, hyaline, yellowish hyaline in Melzer's reagent, smooth, vesiculose to vesiculose-clavate, broadly rounded above, scattered. Hyphae of the

subhymenium small, intermixed, versiform, hyaline, inamyloid. Hymenopodium extraord-

inarily well developed, consisting of thin-filamentose, inamyloid, parallel hyphae. Hymeno- phoral trama proper hyaline, consisting of opaque filamentous hyphae which are axially arranged, subparallel with each other but somewhat away from the edge of the lamellae

they become more and more intricately interwoven-interlaced and quite irregularly arranged, some very broad with thick walls, hyaline, pseudoamyloid (orange brown in Melzer's rea-

gent), easily separating as a layer from the hymenium (when dry sections are made under the binocular one can observe how the trama separates from the subhymenium and hymeniun with the white trama showing; these preparations curl up into ringlets when moistened). Trama of stipe consisting of parallel hyphae which often have thick walls (1.5 M in diameter), strongly reddish pseudoamyloid; all hyphae of trama proper hyaline in NH40H and KOH, partly pseudoamyloid, not gelatinized, with clamp connections. Long oleiferous hyphae present in trama of lamellae. Cortical layers: epicutis of pileus not a continuous hymeniform layer but consisting of isolated nests or fascicles of a somewhat irregular palisade of swollen elements which are either erect or ascending or repent, mostly vesiculose, some cylindrical, 8-22 s, broad, most of these elements filled with a fuscous intracellular pigment but walls hyaline and varying between quite thin and about 1.5 juthick, smooth. Hypoder- mium consisting of repent hyphae which are filamentous, many with a deep sepia-fuscous internal pigment, others partly hyaline, some relatively broad, most narrow.

On rotten leaves and small sticks on humus in tropical rain forest. MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, Guayaramerin, 17 Mar 1956, Singer B 2068

(LIL), TYPUS. ILLUSTRATION. Singer (1965) p 12, fig 61.

223. Marasmius poromycenoides Singer, Sydowia 18: 342. 1965. TYPE. Lowy 365, from Peru.



Marasmius 273

Pileus deep purple ("ambulance", 8 L 6), glabrous, sulcate, reticulate, convex, umbilicate, 11-16 mm broad, Argentine material up to 50 mm broad. Lamellae whitish or

purplish pallid (when fresh ?) whitish to buff-discolored, distant but with numerous lamellae and crowded anastomoses, many of which reach about half the breadth of the through- lamellae and occasionally even more, the lamellulae generally narrower than the through- lamellae which are medium-broad, adnate. Stipe concolorous with or darker than the pileus, smooth and glabrous, hollow when fresh, equal, 35-50 X 1-1.5(-11) mm; basal mycelium strigose, white when dried.

Spores 5.7-6.2 X 3.5-3.8 ,, ellipsoid, smooth, inamyloid, hyaline. Hymenium: basidia 18.7-22 X 5.5-6.3 ,, 4-spored; cystidia none; cheilocystidia numerous and making the edges of the lamellae heteromerous, 22-28 X 5-12 p, hyaline, thinwalled, inamyloid, extremely versiform, some vesiculose, others fusoid-ventricose, others subcylindrical or clavate.

Hyphae of the pileus hyaline, moderately strongly pseudoamyloid, hymenophoral trama

regular, hyaline, also moderately pseudoamyloid; hyphae of stipe parallel with each other, some rather long and relatively wide, strongly (brown-red-) pseudoamyloid. All hyphae with clamp connections. Cortical layers: epicutis of pileus hymeniform, consisting of smooth, vesiculose, subisodiametric cells which are inamyloid, e g 20 X 12.5 , hyaline; hypodermium hyphous, not distinctly pigmented in KOH.

On almost humified mixture of rotten wood and foliage in tropical rain forest, gregarious to cespitose, apparently also on rotten wood in subtropical forest.

MATERIAL STUDIED. PERU. Loreto: Rio Nanya, 31 Oct 1958, B. Lowy 365 (LIL) type. ARGENTINA. Misiones: Iguazu: Refugio "Yaguarete," 29 Feb 1960, R. T. Guerrero 132 (LIL).

By its aspect this would be identified as "Poromycena" i e a Mycena sect Purae with

subporoid hymenophore. However, an analysis of the specimens shows that they belong to Marasmius sect Globulares where they stand out because of their color. The Argentine material is much larger than the Peruvian one and sterile. It is included here with some doubt.


224. Marasmius oreades (Bolton ex Fries) Fries, Epicrisis Systematis Mycologici, p 375. 1838.

Agaricus oreades Bolton ex Fries, Systema Mycologicum 1: 127. 1821. Marasmius caryophylleus (Schaeffer) ex Schroter in Cohn, Kryptogamenflora Schlesiens 3a: 561.

1889. TYPE. Bolton, England (apparently not conserved). Pileus fulvous to testaceous-tan to pale cinnamon buff, the paler forms often develop-

ing in age at dry weather conditions, not strongly hygrophanous ("Yucatan" to "raw sienna", then often 11 G/H 7, 12 G 8), not viscid, glabrous, in age with striate margin or short- furrowed-crenate, not long-deep-sulcate, mostly smooth over most of the surface, campanulate-convex, then convex-subumbonate or somewhat gibbous, eventually tending to be

applanate, obtuse to more often obtusely umbonate or gibbous with uplifted margin, 20-50 mm broad. Lamellae white, then ochraceous-cream, rather distant to distant, eventually sometimes slightly intervenose by low veins, rounded-subfree. Spore print white. Stipe white to concolorous (with pileus) but paler or pallid with merely a flush of the pileus color, covered with a fine tomentum or pubescence, equal or subequal, solid to stuffed, hollow only when overmature and sometimes on drying, 35-75 X 3-5 mm; pseudorhiza slightly developed or none; basal mycelium not conspicuous, white woolly, delicate, some occasional strands of white rhizomorphs occurring. Context white, rather thick in center of pileus, flesh-tough, tough in stipe, odor of cyanic acid, but weak.

Spores (7-)8-10.5 X 5-5.8 p, ellipsoid-subfusoid, smooth, inamyloid, hyaline. Hymen-




ium: basidia 25-30 X 6 ,, 4-spored; cystidia none; cheilocystidia inconspicuous. Hyphae strongly pseudoamyloid, with clamp connections, hymenophoral trama of strongly interwoven hyphae, subirregular near context. Cortical layers: epicutis of pileus hymeniform, cellular.

On earth in open places, pastures, meadows, in South America only adventive on flower beds and lawns, often forming fairy rings.

MATERIAL STUDIED. Numerous European, Asiatic and North American collections. In South America: ARGENTINA. Buenos Aires: Gonnet, 26 May 1940, Singer S 26 (LIL); Finca Pascuala, 16 Nov 1962, Singer 5363 (BAFC); Sierra de la Ventana, 14 Nov 1962, Singer S 229 (BAFC); Pampa: Trelen, 24 Apr 1966, Singer (BAFC).

This species is not properly South American but has been introduced in regions which have subtropical to temperate vegetation. It is therefore here admitted. The South American material has probably been introduced with grass seeds from North America or Europe. MAar- asmius oreades has also been observed in Florida, U.S.A., probably likewise adventitious.

The collection from the province of Pampa was consistently whitish. It is possible that a whitish variety, also observed in Europe, can be separated from the type variety.

ILLUSTRATION. Singer (1961) pi 1. Horak, (1968) fig a,b,c,d,e,f.

225. Marasmius cibarius Singer, sp nov. TYPE. Vargas 10050,, from Peru. Pileo brunneo vel fuligineo-brunneo in siccis, + 20 mm lato; lamellis pallidis siccando

grisascentibus, sat latis, ventricosis, sat distantibus, adnexis; stipite pallido vel pileo pallidiore, farcto, haud insititio. Sporis 5.5-6.5 X 2.8-3.8 ,; cystidiis nullis; cheilocystidiis versiformibus; hyphis pseudoamyloideis. Ad terram extra silvas in zona alta andina, Peru. Ty- pus in LIL conservatus est.

Pileus dried "burnt umber" on the margin, "cocoa" to "seal" (M&P) further inwards and often reaching a deep blackish brown, opaque, smooth, nonviscid, non-hygrophanous, convex, then more repand, obtuse, about 20 mm broad. Lamellae pallid, tending to dirty grayish when dried, rather broad, ventricose, rather distant, adnexed. Stipe pallid or lighter colored than the pileus, subglabrous to slightly fibrillose, stuffed, not tubular-hollow, longer than the diameter of the pileus, about 3 mm broad, not insititious. Context white, unchanging on drying.

Spores 5.5-6.5 X 2.8-3.8 , ellipsoid to oblong with flatter inner side, often as narrow as 6.5 X 3 u or as broad as 6 X 3.5 C,, smooth, hyaline, inamyloid. Hymenium: basidia 27- 28 X 4.8-6.3 u, 4-spored, clavate, hyaline; cystidia none; cheilocystidia versiform, mostly inconspicuous and basidiomorphous. Hyphae strongly pseudoamyloid, with clamp connections. Cortical layer: epicutis of the pileus hymeniform or subhymeniform, consisting of clavate to vesiculose smooth elements which are mostly melleous to light fuscous by an intracellular pigment.

On earth, outside the forest near water; alpine zone of the Andes. MATERIAL STUDIED. PERU. Cuzco, Canal, El Descanso, 4200 m alt, 5 Feb 1951, Vargas

10050 (LIL), type. This species is edible and used for food in the Cuzco region like the following species.

226. Marasmius albogriseus (Peck) Singer, Ann. Mycol. 41: 130. 1943. Collybia albogrisea Peck, Bull. Torrey Club 22: 199. 1895. Gymnopus albogriseus (Peck) Murrill, North American Flora 9: 368. 1916. Collybia fimicola Earle, Bull New York Bot. Gard. 3: 298. 1904. TYPE. McClatchie, from California (type not seen but possibly available at the Pas-

adena Museum).



Marasmius 275

Pileus fresh "yellow beige" to pale fuscous (13 G 7), central portion very frequently deeper fuscous in youth ("winter leaf," "English oak," at times reaching "Congo") but bleaching to paler and dirtier than "ivory" and often found in intermediate colors, dried assuming a color near "olive wood," "teak wood," "mummy," "cocoa," "seal" in the center, the margin remaining pallid or drying to "cookie," "burnt umber" or "roe," i e generally reddish brown to grayish fuscous with paler margin and the more rapidly dried and the less pressed the less reddish in tone, subhygrophanous or almost non-hygrophanous, short transparently striate tending to become short coarsely sulcate when dried, opaque to weakly opimous, not viscid, glabrous, naked, smooth in fresh condition, with sometimes somewhat irregular margin, not shining or very little so when dried, convex, then irregularly flattened, subumbonate, more rarely quite obtuse or distinctly umbonate, 18-45 mm broad. Lamellae sordid beige ("Malacca" to "Arizona"), bleaching in age to paler and dirtier than "ivory," dried whitish to subconcolorous with margin of pileus, usually whitish, subdistant to distant, broad to very broad, ventricose, adnexed to rounded-adnexed, not intervenose except in large, old specimens occasionally very minutely intervenose. Spore print white. Stipe whitish to grayish pallid or pale cream gray (usually about "seaside, tea time" to "cracker" below and paler to pallid above), finely pruinate to pruinate fibrillose especially above, but glabrescent, hollow, sometimes becoming canaliculate, equal, subequal, or with slightly enlarged base, 20-50 X 2-9 mm; basal mycelium typically tomentose near base and at the base unless scanty when base immersed in strongly adhering earth particles, sordid white to grayish or dull cream. Context white, unchanging but under the cuticle concolorous with surface, in stipe concolorous with surface; odor weak, like that of M. oreades; taste agreeable, mild, not farinaceous.

Spores 5.5-7 X 2.8-4.3 A, mostly 6-7 X 3-4 p, hyaline, smooth, ellipsoid to oblong with flatter inner side, thin-walled, inamyloid. Hymenium: basidia 27-29 X 4.8-6.3 p, 4- spored; cystidia none; cheilocystidia very inconspicuous, filamentous, clavate or clavate- forked, mostly basidiomorphic, sometimes septate, hyaline. Hyphae: Hymenophoral trama regular, consisting of hyaline, thin-walled, subparallel hyphae; all hyphae with clamp connections and strongly pseudoamyloid. Cortical layers: epicutis of pileus subhymeniform, consisting of thin-walled hyaline or (by a dissolved intracellular pigment) mielleous to light fuscous cells which are clavate to balloon-shaped.

On earth among leaves or grasses in open places, in North America in low altitudes, in South America only in the drier portions of the alpine and puna zones but preferring the margin of wet places, water puddles and generally wet soil, gregarious to generally cespitose or subcespitose. Fruiting in South America from January until March.

MATERIAL STUDIED. U.S.A. California: San Mateo County, Stanford University, C. F. Baker (NY) type of M. fimicola. PERU. Cuzco, Urubamba, Chic6n, 3400 m alt, 6 Feb 1963, Zamaolla 202, 203 (BAFC). BOLIVIA. Cochabamba, Colomi, 3400 m alt, Jan 1949, Ctrdenas 243 (LIL). ARGEN- TINA. Tucuman: Cumbres Calchaquies, Laguna Seca, 4150 m alt, 14 Feb 1952, B. Sparre 9587 (LIL); Infiernillo, 3000-3200 m alt, 23 Jan 1960, Singer T 3550 (LIL); Catamarca: between Minas Capillitas and Cerro Negro at 3700 m alt, 3 Mar 1952, B. Sparre, comm. Singer T 1915 (LIL).

In the Andes of Peru, the Quichua population uses this species as food. There the species has a certain economic importance since in those altitudes few other edible species can be gathered in quantity.

The South American high altitude populations are morphologically inseparable from the California type. The whitish gray pileus turns reddish brown in both the North Amer- ican and the South American forms. Some differences in spore measurements are due to individual variability of the average spore sizes in different populations but do not coincide with area or altitude differences. A certain weak intervenosity of the lamellae can likewise be observed here and there. The larger spores of the type of C. fimicola are explained by




the presence of 2-spored (larger) basidia, this again being, perhaps, a consequence of the fimicolous habitat which is obviously a coincidence rather than an ecological specialization.

ILLUSTRATION. Horak (1968), fig a-e.

227. Marasmius niveus Montagne, Ann. Sci. Nat. IV. 1: 117. 1854. Collybia nivea (Montagne) Dennis, Trans. Brit. Mycol. Soc. 34: 434. 1951.

TYPE. Leprieur 1038, from French Guyana. Pileus white or cream white, center sometimes light cinnamon, strongly sulculate

to center, glabrous, with a shallow central umbilicus, otherwise convex, soon convex-

applanate and sometimes with wavy margin, 30-50 mm broad. Lamellae white or whit-

ish, subclose to distant, mostly subdistant, narrow, adnate, not inter venose but occasion-

ally some forked. Stipe fulvous to chestnut with white or cream colored apex, glabrous or subglabrous with pruinose but glabrescent apex, equal or slightly tapering downwards, hollow-tubular, up to 105 mm long, 2.5-3 mm broad, basal mycelium tomentose, white or whitish. Context white, somewhat tough, unchanging, with odor of crab or fresh fish.

Spores 5.3-9 X 2.7-3.5 y,, according to Dennis up to 11 X 3 u, ellipsoid-oblong to

fusoid, hyaline, smooth, inamyloid, with a tendency to form, when overmature, two or one internal septum with a small apical cell or appendage and sometimes a basal cell, much like Crinipellis mirabilis or Marasmius arborescens. Hymenium: basidia clavate,

hyaline, 4-spored; cystidia none; cheilocystidia vesiculose, small, 12.3-13.7 X 5.5-8.3 ,

hyaline, smooth. Hyphae: hymenophoral trama regular, hyaline, in trama of pileus hyaline to subhyaline, of very different sizes and shapes although always elongated; all

hyphae strongly or at least distinctly pseudoamyloid, many thick-walled, with clamp connections. Cortical layers: epicutis of the pileus hymeniform, hyaline, consisting of cellular elements which are often clavate-pedicellate e g 29 I long and 11-19 9 broad.

On dead twigs and leaves of monocotyledonous and dicotyledonous trees in tropical and tropical-montane forest, gregarious.

MATERIAL STUDIED. VENEZUELA. Caracas: Rio Chacaito, 18 Nov 1949, Dennis 265 B

(K). FRENCH GUYANA. Leprieur 1038 (PC), type. BRAZIL. Amazonas: Remate dos Males, 1927, P. Occhioni 11 (RB 5.2.04) (LIL). BOLIVIA. Pando: Manuripi, Conquista, 24 Mar 1956, Singer B 2179 (LIL); 25 Mar 1956, B 2252 (LIL). PERU. Cuzco: Convenci6n; Sahuayaco, 1200 m alt, 17 Jan 1947, Vargas 6315 (CUZ, LIL).

This species is also reported to occur in Trinidad (Dennis, 1970). ILLUSTRATION. Singer (1965) pl 12, fig 65; Dennis (1970), pl 3, fig 1.

228. Marasmius strictipes (Peck) Singer, Lilloa 22: 326. 1951. Fig 96. Collybia strictipes Peck, Ann. Rep. New York State Museum 41: 62. 1888.

Gymnopilus strictipes (Peck) Murrill, North American Flora 9: 357. 1916.

TYPE. Peck, from New York, U.S.A. Pileus white with stramineous center or entirely whitish, not distinctly hygrophan-

ous, but later developing in many cases pale fulvous to rusty fulvous spots or areas, not

viscid, glabrous, mostly with corrugated-rugose or reticulate-rugose center, more rarely entirely corrugated, with entire or lobed margin which is incurved at first, convex, later more applanate and even depressed around a low and obtuse umbo or subumbonate, 26- 60 mm broad. Lamellae white to whitish, not or weakly intervenose or more or less

anastomosing, narrow to very broad, crowded, close or subdistant, rounded-adnexed, ad-

nexed or subfree to free. Stipe white, often tending to orange cinnamon below, white fibrillose to subglabrous but with mealy apex, equal or subequal or with slightly broader

apex or base, hollow, usually straight, 47-88 X 1.5-5.5(-8) mm; basal mycelium abund-



Marasmius 277

ant, white, often connection with the base of neighboring specimens. Context white or whitish, unchanging, odor of crabs or crayfish mixed with the odor of Marasmius oreades.

Spores 7.5-8.5 X 3.8-5 ,, mostly about 7.7-8.3 X 4-4.8p , ellipsoid to oblong or narrowly ovoid, smooth, hyaline, inamyloid. Hymenium: basidia 23-28 X 5.5-8 5, 4-

spored; cystidia none; cheilocystidia 26-28 X 2-5 p, irregular in shape and size, often filamentous to cylindric-flexuous or cylindric-subclavate, stramineous to subhyaline. Hyphae hyaline, with clamp connections, with thin walls, some with thick walls (these having the aspect of hairs of Crinipellis and are particularly strongly pseudoamyloid), all pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform, consisting of erect hyaline or subhyaline cells which are subglobose, clavate vesiculose or piriform, entire, thin-walled, not intermixed with dermatocystidia.

On rotting leaves and rotten pieces of wood of dicotyledonous trees growing gregariously or mostly cespitosely, fruiting in summer and fall. Known hosts: Quercus, Carpinus, Inga, Coffea. Restricted to the temperate to tropical-montane zones of North America.

MATERIAL STUDIED. U.S.A. New York: Catskill Mountains (NYS), type. MEXICO. Michoacan, San Miguel del Monte, R. Palacios, 18 Nov 1967, 2200 m alt, (ENCB); Oaxaca: Huautla de Jimenez towards Rancho del Cura, 1500 to 1560 m alt, 10 Jul 1957, Singer M 1510 (LIL); Guz- tlibn 948 (ENCB), 7 Jul 1969, Singer M 8331 (F); near Chiltepec, 30 Oct 1960, Guzman 2786 (ENCB).

ILLUSTRATION: Kauffman (1918) pl 161. Smith (1938) fig 1 b, d.

229. Marasmius heliomyces Murrill, Bull. Torrey Club 67: 149. 1940. TYPE. West, Arnold & Murrill, from Florida (U.S.A.). Pileus young "Mummy Brown" (Ridgway) to fuscous black, soon gradually whit-

ish towards the margin, or inversely, "Mummy Brown" all over excepting the disc where the color is sharply changed to much paler, eventually often brownish gray or mouse gray, excepting a characteristic but not fully constant pallid zone between disc and margin leaving the center zone star-shaped, glabrous, dry or subhygrophanous, striate, then

deeply long-furrowed-sulcate, sometimes even rimose, usually ? rugulose, semiglobate to convex, then convex-applanate or subapplanate, usually umbilicate, circular and entire, eventually often lobed, 13-40 mm broad. Lamellae varying from white (especially in

youth) to "cartridge buff' (Ridgway), especially in age and when dry, at first ascendant, soon horizontal, mostly subventricose, broad (4-8 mm), distant to very distant, deeply emarginate-adnexed to subfree. Stipe hyaline, then white with chestnut colored base, later "burnt sienna" to "Mahogany" (Ridgway) from the base upwards, sometimes leaving an intermediate citrinous zone between the apex and the rest of the stipe, on

drying the chestnut color often replaced by a cinereous to pale umber color, macroscopically glabrous, shining, smooth, equal or more rarely tapering downwards, tubulose- hollow, 40-80 X 2-3(-5) mm; basal mycelium abundant, white. Context thin; odor none or weakly of Marasmius oreades but sometimes when fresh either rancid or of honey, but these odors soon disappearing.

Spores (5.2-)6-8.8 X 2.8-3.7 ,j, smooth, hyaline, inamyloid. Hymenium: cystidia none; cheilocystidia 9.8-18 , broad, vesiculose and mostly broadly rounded above, some mucronate, some broadly cylindric, some more clavate, without conspicuous appendages or setulae, hyaline. Hyphae of the context of the pileus hyaline, strongly pseudoamyloid. Cortical layers: epicutis of the pileus hymeniform, much like the cheilocystidia but not all hyaline, fuscidulous in the dark brown zones.

On the base of stumps and on rotten trunks, also on dead leaves, solitary or gregarious, in Florida often on Quercus virginiana




MATERIAL STUDIED. U.S.A. Florida: Highlands County, Highlands Hammock State Park, Aug to Sep 1942, Singer F 73, F 416, F 513 (FH); Alachua County. Gainesville, 21 Jul 1938, West, Arnold & Murrill F 18277 (FLAS), type; Dade County: Matheson Hammock, 20 Oct 1942. Singer F 1152 (FH).

The form occurring in southern Florida has less chestnut brown stipe, more hygrophanous pileus, and an odor of honey; it may represent a variety and, again, be the same as Marasmius tortipes, see p 284.

230. Marasmius pseudocollinus (Singer & Digilio in Singer) Singer, Sydowia 18: 229. 1965.

Mlarasmius cohortalis var pseudocollinus Singer & Digilio in Singer, Sydowria 12: 91. 1959.

TYPE. Singer T 1329, from Argentina. Pileus between light golden ochraceous and light buff in the center ("Inca gold."

"Pablo"), gradually whiter outwards with white or whitish marginal zone, transparently striate when wet, subhygrophanous, dried light buffish ochraceous (11 G 6) in center and white on margin, smooth when dry, glabrous, convex, sometimes subumbonate, in age applanate, 8-31 mm broad. Lamellae white, eventually often sordid pallid, adnexed to adnate, subclose to subdistant, at first rather narrow but soon moderately broad to

very broad (more often very broad in fully mature material) not or slightly intervenose, more generally moderately intervenose when quite mature, at times some forked. Spore print white. Stipe white, in age light cinnamon with white apex, not deep colored ever in lower portion, finely pruinate to slightly tomentose, equal or tapering downwards, narrowly tubular, 20-38 X 1.5-2.5(-4) mm; basal fibrillosity scanty, white. Context white, odor none or of Marasmius oreades (HCN), or agreeable (as in some Suilli).

Spores 6.5-7.5 X 3.3-3.5 ,, ellipsoid to ellipsoid-oblong, rarely slightly sausage- formed or tilda-shaped, eventually sometimes bicellular (in overmature spores) as in Crinlipellis mirabilis or Marasmius arborescens, hyaline, smooth, inamyloid. Hymenium: basidia 29.2 X 5-5.3 p,, 4-spored; cystidia none; cheilocystidia none, or very inconspicuous. Hyphae of pileus trama very slightly to distinctly pseudoamyloid, those of stipe strongly pseudoamyloid, all with clamp connections. Cortical layer: epicutis of pileus hymeniform, consisting of erect elements, the terminal ones subisodiametric or short- clavate e g 20-26 X 10-12.3 #, below often shorter or longer cells rising vertically from the cutis formed by the hypodermium, sometimes with crystals adhering to the apex or with very scattered epimembranal pigment bodies.

On deep earth, often among herbs or grasses, open places of the plains and up to the montane zone (up to 1200 m altitude) also under scattered trees on earth among Gramineae, gregarious.

MATERIAL STUDIED. ARGENTINA. Tucuman: Aguadita, 450 m alt. 25 Feb 1951, Singer T 1329 (LIL) paratype; Capital, Villa Lujan 450 m alt, 22 Jan 1955, Singer T 2070 (LIL) paratype; Taft: Anta Muerta, 1200 m alt, 26 Dec 1951, Singer T 1667 (LIL) type.

ILLUSTRATION. Singer (1965) pl 12, fig 63

231. Marasmius cohortalis Berkeley, Videnskaps Meddelingen Naturhistoriska Fereniging, Kjobenhavn, p 34. 1879.

Collybia nivea var cohortalis (Berkeley) Dennis, Trans. Brit. Mycol. Soc. 34: 435. 1951. Agaricus hymeniicephalus Spegazzini, Anal. Soc. Ci. Argentina 16: 244. 1883. Collybia hymeniicephala (Spegazzini) Saccardo, Sylloge Fungorum 5: 242. 1887. Heliomyces hymeniicephalus (Spegazzini) Spegazzini, Bol. Acad. Nac. Ci. Cordoba 23: 386.

1919. Marasmius hymieniicephalus (Spegazzini) Singer, Lilloa 23: 197. 1951. Gymnopilus ludovicianus Murrill, North American Flora 9: 355. 1916. Marasmius ludovicianus (Murrill) Singer, Lilloa 22: 326. 1951, non Planchon. Marasmius alachuanus Murrill, Lloydia 5: 140. 1942.



Marasmius 279

231a. Marasmius cohortalis var cohortalis TYPE. Glaziou 9171, from Brazil. Pileus "mellow glow" to "Inca gold" (M&P) in center, watery white on margin,

hygrophanous, pure white on the margin and whitish to yellowish cinnamon white on the disc when dry, sulcate or rugose in a zone about 2-3 mm broad from the margin inwards, glabrous, not viscid, convex to convex-umbonate in age, convex with slightly depressed to umbilicate center, eventually sometimes applanate and umbilicate, sometimes somewhat eccentric, 14-23 mm broad. Lamellae pallid to pale ochre cream (9 D 3), rather narrow to moderately broad (2-2.8 mm broad), moderately close to distant, strongly intervenose in the larger mature carpophores, less or not so in the young and small carpophores, adnate. Stipe chestnut color below with gradual transition towards the white apex, entirely light castaneous-cinnamoneous when dried, finely pruinate but glabrescent, hollow, equal, somewhat eccentric, 36-42 X 1.2-2 mm, basal tomentum whitish, becoming sordid when dried. Context whitish, watery when wet, partly concolorous with the surface, especially in the lower portion of the stipe, almost inodorous.

Spores 7-8.3 X 3 ,, oblong-ellipsoid, smooth, hyaline, inamyloid. Basidia and cheilocystidia as in var hymenicephalus; Hymenium: basidioles fus-

oid: basidiole-like cystidioles or pseudoparaphyses present but scattered, "empty," some with slightly forked-sinuate apex, not projecting and not deeper-rooting than basidioles, not opaque or opalescent, not conspicuous; true cystidia none. Hyphae: hymenophoral trama consisting of hyphae running not all in the same direction, intermixed with a few very broad and short cells, some hyphae with moderately thick wall, most thin-walled, all hyaline; all hyphae with clamp connections, pseudoamyloid. Cortical layer: epicutis of pileus hymeniform, consisting of vesiculose elements which are smooth but often forked into two rounded lobules or protracted and nodulose and ramose or with 1-2 appendages or a mucro, some broader than high, some somewhat elongated, hyaline.

On very rotten wood, on humus and on leafmold in tropical, subtropical and sub-

tropical-montane forest, gregarious. MATERIAL STUDIED. BRAZIL. Glaziou 9171 (K), type. ARGENTINA. Tucuman: Taft:

Vieja carretera al paso San Javier, 24 Apr 1951, Sinlger T 1527 (LIL); Rio de los Sosas, 4 Feb 1955, Singer T 2119 (LIL).

ILLUSTRATION. Singer (1958) fig 14.

231b.Marasmius cohortalis var hymeniicephalus (Spegazzini) Singer, Sydowia 12: 89. 1958. Agaricus hymeniicephalus Spegazzini, Anal. Soc. Ci. Argentina 16: 244. 1883. TYPE. Balansa 3380, from Paraguay. Pileus hyaline-white, hygrophanous, dry-faded white or whitish, sulcate and/or ret-

iculate to pustulate, center smooth to rugulose-venose, sometimes from margin inwards

pustulate, further inwards sulculate, further smooth and in the very center venose, glabrous, not viscid, convex, soon becoming applanate, with a sometimes rather shallow central umbilicus, 10-37 mm broad. Lamellae white, sordid white or cream, medium close to distant, narrow to moderately broad (up to 3.5 mm broad), generally linear, often forked, subarcuate, intervenose to anastomosing, i e connecting lamellae venose to high and lamellar (reaching up to three quarters of the height of the through-lamellae, more rarely as high as the latter) adnexed to adnate or even adnate-subdecurrent. Sport print white. Stipe chestnut brown or ochraceous fulvous below, gradually or abruptly hyaline or white at apex, at first fresh finely pruinate all over or at least at apex, sometimes




finely fibrillose all over, but some glabrescent, often eccentric, equal or sometimes slightly attenuate in lower half, smooth, soon tubular, terete or canaliculate, otherwise smooth, 30-100 X 1.2-3 mm; basal mycelium tomentose, white, in dried condition often sordid white, often continuing onto surface of lower part of stipe as a fine pubescence, often

covering substratum with white arachnoid filaments, forming thin membranous films, but sometimes scanty. Context white, inodorous or with slight crustaceous (shrimp) or marine algae odor.

Spores 4.8-8.2 X 2.74.8 u, mostly 5.5-7 X 3-3.5 p, ellipsoid, oblong, or more rarely subcylindrical, smooth, hyaline, inamyloid. Hymenium: basidia (15-)22-25.5(-33) X 5-7 ,p

4-spored; cystidia none; cheilocystidia small to medium sized cells like those of the epicutis of the pileus but rather inconspicuous. Hyphae of pileus trama and hymenophore varying from slightly and incompletely pseudoamyloid to distinctly and strongly pseudoamyloid, all hyaline and with clamp connections. Cortical layer: epicutis of pileus con-

sisting of subglobose, pedicellate or clavate-vesiculose cells which are hymeniformly arranged and either completely smooth and entire, or in some sections frequently nodulose or hook-bearing or beset which a few prongs at the apex where they might be somewhat

gelatinized outside since the spores often adhere to the excrescences of the epicuticular cells, the latter 17.8-23 X 7-22 ,. hyaline.

On woody humus and on leaves and branches rotting on the ground in subtropical, tropical-montane, tropical and subtropical-montane forests, gregarious to fasciculate.

MATERIAL STUDIED. PARAGUAY. Guarapf, January 1880, Balansa 3380 (LPS, K) type. BRAZIL. Sao Paulo: Apiaf (Apiahy) Jun 1890, Puiggari (LPS): Rio Grande do Sul: Sao Leopoldo, J. Rick (as M. svnodicus) (FH). BOLIVIA. Pando: Manuripf, Conquista, 23 Mar 1956, Singer B 2178 (LIL); La Paz: Nor-Yungas: Rio Llolosa, 31 Jan 1956, Singer B 782 (LIL); Coroico, 30 Jan 1956, B 714 (LIL). ARGENTINA. Tucuman: Quebrada de Lules, 18 Feb 1951, Singer T 1246 (LIL); Rio de los Sosas, 1 Jan 1951, T 1084 (LIL); Vieja carretera al San Javier, 13 Feb 1951 T 1200 (LIL); Misiones, Arroyo Piray Guazui, 24 Feb 1960, R. T. Gurenero 71 (LIL); Buenos Aires: Santa Catalina. Llavallol, 19 Mar 1966, Singer S 632 (BAFC).


231c.Marasmius cohortalis var arenicolor Singer, Sydowia 12: 92. 1958. TYPE. Singer T 1445, from Argentina. Pileus sand color to gray ("cracker" to 13 F 7, "lariat" or 12 F 6 to 11 G 5),

hygrophanous, dry-faded white to whitish, dried mostly "clay" or grayish, rarely white, fresh, dry and dried often with paler margin, smooth, but in age becoming more or less sulcate and + sulcate on the margin when dry, and dried, transparently striate when wet, glabrous, not viscid, convex, soon becoming umbilicate or applanate on disc, eventually more applanate-repand and often with uplifted margin, 11-38 mm broad. Lamellae hyaline white to pale creamy gray ("Longbeach," "Malacca," "old Ivory"), mostly strongly intervenose except for immature carpophores, subdistant to distant, rather narrow to broad (to 4 mm broad), somewhat ventricose at times, adnexed to adnate, eventually often subcollariate. Stipe white, in lower portion soon becoming yellow to brown ("clay," "spruce y," "cognac") with gradual transition, extremely finely pruinate-fibrillose, glabrescent, hollow, sometimes canaliculate, often spirally twisted when dried, otherwise smooth, equal or sometimes with slightly enlarged base, or base and apex 17-80 X 1.8- 3.8 mm; basal mycelium whitish, rarely forming small white rhizomorphic filaments. Context white, either inodorous or (mostly) with a slight and pleasant fruity odor (as in some Suilli).

Spores (5.5-)6.5-8.2(-8.8) X 2.5-4(-5.2) s, ellipsoid, thin-walled or almost so, rarely with suprahilar depression, hyaline, smooth, inamyloid. Hymenium: basidia 31.5 X 6.8-



Marasmius 281

7.5 ,, 4-spored; cystidia none; cheilocystidia versiform. Hyphae distinctly pseudoamyloid, especially in the stipe. Cortical layer: epicutis of pileus hymeniform, consisting of clavate- vesiculose to subglobose cells whose upper portion is very rarely and rather indistinctly nodulose, hyaline, inamyloid, 23-29 X 10-24 p, dermatocystidia of the surface of the stipe erect or appressed to the uppermost parallel hyphae of the stipe, thin-walled, polymorphous but often cylindric, inamyloid, 9-41 X 3.5-4.8 u.

On woody humus and leafmold in tropical-montane, subtropical, subxerophytic and xerophytic forest and in parks. Southern Hemisphere.

MATERIAL. STUDIED. BOLIVIA. La Paz: Nor-Yungas, Charobamba, 13 Feb 1956, Singer B 1126 (F). ARGENTINA: Tucuman: Capital, Parque Avellaneda, 28 Mar 1951,Singer T 1445 (LIL) type; Trancas: between Salta Road and La Higuera, 25 Jan 1955, Singer T 2103 (LIL) paratype; Dry hills above San Pedro de Colalao, 23 Jan 1955, Singer T 2059 (LIL) paratype; Chicligasta: Los Sosas, 4 Feb 1955, Singer T 2270 (LIL).

231d.Marasmius cohortalis var alachuanus (Murrill) Singer, comb nov Miarasmius alachuanus Murrill. Lloydia 5: 140. 1942. TYPE. Murrill, from Florida, U.S.A. Pileus "Cream Buff" to "Cartridge Buff" (Ridgway), hygrophanous, becoming rather

uniformly milk-white dried or faded, transparently reticulate when moist, scrobiculate- rugose when dried, besides coarsely sulcate over about three quarters of the radius at least when fresh and faded and over one side of the pileus, more rarely slightly sulcate to subsmooth on the margin, glabrous, not viscid, convex, soon umbilicate or with depressed center, eventually often applanate or irregular, 8-33 mm broad. Lamellae white to buffish white but sometimes eventually cream-grayish, intervenose, even anastomosing in some old specimens, horizontal, variable in number, varying from subclose to subdistant (distant to very distant if only the through-lamellae are taken into consideration), 1-4 mm broad, not ventricose, adnexed, often adnexed to a widened apex of the stipe and the eventually either appearing decurrent or separating from the apex but not truly collariate. Stipe white becoming tawny, russet and Mars brown (in this order) when mature or when old, remaining pallid at apex, smooth but sometimes canaliculate, macroscopically glabrous, + subpruinate under a lens, hollow, equal or tapering downwards, 30-48 X 1.5-4 mm; basal mycelium abundant, sometimes extensive and membranous, white or cinnamon-pallid. Context very thin, white, unchanging, with an odor of crabs when quite fresh (of "anise" according to Murrill); taste mild.

Spores (4.5-)6-8(-8.5) X (2.3-(3-4(-4.5) u, fusoid or ovoid-oblong to almost ellipsoid, in profile mostly somewhat applanate on inner side but generally without a distinct suprahilar depression, thin-walled, smooth, hyaline, inamyloid. Hymenium: basidia relatively long, 4-spored; cystidia few like the cheilocystidia and near the edges or none whatever: cheilocystidia not very numerous in most collections and not making the edge of the lamellae heteromorphous, 24-29 X 5.5-10.8 u, versiform, often clavate to vesiculose, hyaline. Hyphae of the regular hymenophoral trama hyaline, interwoven, thin- to slightly thick-walled, not gelatinized, with clamp connections, strongly pseudoamyloid; in pileus trama similar; in stipe trama parallel. Cortical layer: epicutis of the pileus hymeniform, consisting of erect smooth cells, 13-25 X 7.5-20 u, mostly vesiculose to piriform, obtuse or a few with appendage, without setulae, hyaline or subhyaline in KOH.

On leaf mold in humid forest, low, mesophytic and high hammock type but also in tropical hammock, on dicotyledonous leaves, often of Quercus but also other trees, in the Gulf region including all of Florida, Louisiana, Puerto Rico, fruiting from June until October.




MATERIAL STUDIED. U.S.A. Florida: Alachua County, Gainesville, 16 Aug 1939, Murrill F 19885 (FLAS), type, 2 Jul 1943, Singer F 2511 (F); Dade County, Miami, Simpson Park, 14 Sep 1942, Singer F 688 (F). Louisiana: New Orleans, City Park, 3 Sep 1908, Earle 36 (NY), type of Gymnopus ludovicianus. PUERTO RICO. El Yunque, Jun 1968, P/z. Clark(F).

ILLUSTRATION. Smith (1938), fig 1, c, g.

232. Marasmius riparius Singer in Singer & Digilio, Lilloa 25: 198. 1952. TYPE. Singer T 1244, from Argentina. Pileus colored like A. oreades, hygrophanous, cinnamon-white when faded, semi-

globose, then covex, soon umbilicate, not viscid, glabrous, deeply and coarsely sulcate over half or three quarters of the radius, 25-45 mm broad. Lamellae concolorous with pileus, regularly intermixed by lamellulae which are rounded at inner end, ventricose, up to 10 mm broad, i.e. broad, distant, not intervenose, adnexed. Stipe white, then cinnamon with cinnamon-white apex, rarely becoming uniformly cinnamon, tubular, velutinous, at the apex somewhat ribbed, often spirally twisted, 25-60 X 1-3 mm. Context white in pileus, cinnamon brown in stipe; odor weak, of Micromphlale .tetidum.

Spores 8.2-10 X 4.8-5.5 p, ellipsoid-fusoid, smooth, thin-walled, often with one oil droplet or with very small droplets, hyaline, inamyloid. Hymenium: basidia 4-spored: cystidia 40-44 X 8-9 ,, more rarely narrower, opaque, hyaline-yellowish in Melzer's reagent, clavate but slightly short-attenuate at the extreme apex, very numerous on sides and edges of lamellae. Hyphae all with clamp connections, pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of obpiriforn pedicellate cells which are 13.7-18 X 11-13.7 ,, not incrusted, hyaline or with a very dilute umber colored intracellular pigment.

On forest detritus in marginal (gallery) woods of the subtropical zone. MATERIAL STUDIED. ARGENTINA. Tucuman: Quebrada de Lules. 18 Feb 1951, Singer

T 1244 (LIL) type. ILLUSTRATION. Singer (1958a) fig 15.

233. Marasmius silvicola Singer in Singer & Digilio, Lilloa 25: 199. 1952. TYPE. Singer T 1479, from Argentina. Pileus deep cinnamon to ochraceous-brownish ("Java," "Maracaibo") in the center

and on the striae, paler between the striae, also often "sayal br.," "amber br.," "Martin- ique" in the center, otherwise 9 F 6 to "walnut taffy," hygrophanous, fading to avellaneous, dried colored like M. pampicola or somewhat paler, glabrous, smooth or shallowly to coarsely sulcate on margin, especially when dry, transparently striate over 1/7 to 4/5 of radius when wet, convex, eventually applanate or with slightly depressed center, often umbonate, 17-95 mm broad. Lamellae pale and sordid grayish cinnamon ("almond") but in some collections more yellowish ("golden corn" to "saratoga") or more gray but in many cases on sides and lamellae-ground in the part near the margin of the pileus concolorous with the latter, regularly intermixed with lamellulae, the lamellulae often crisp or venose, ventricose or subventricose, rather broad to broad (2-4 mm, mostly 4 mm broad), with entire edge, rounded-adnate to rounded subfree or narrowly adnexed, rarely broadly adnexed to adnate, not intervenose, subclose to distant. Stipe pallid to cream-grayish (concolorous with the edges of the lamellae), fresh subglabrous to white or whitish, thin-tomentose of fuscous ground, deeper colored below (concolorous with pileus or somewhat paler) at least in most adult specimens, often canaliculate, otherwise smooth, tubular-hollow at least in upper portion, eventually always entirely hollow, equal or towards the base slightly to distinctly subbulbous or attenuate, 20-118 X 1.5-7



Marasmius 283

mm, if widened at base, reaching up to 9 mm in diameter there; basal mycelium tomentose or woolly white to sordid whitish, sometimes very abundant. Context whitish to fuscous, mostly concolorous with surfaces in cortical zones, especially in cortex of stipe, inodorous or with very weak odor.

Spores 4.5-8.3 X 2.5-4.2 s,, mostly 5.5-6 X 3-3.5 ,, ellipsoid or tear-shaped, smooth, hyaline, the hilar portion often somewhat incurved, thin-walled, inside granular or with one round or more rarely elongated oil droplet, inamyloid. Hymenium: basidia 26-36 X 5.5-7.5 ,, 4-spored, rarely a few 2-spored ones present; cystidia 33-100 X 5.8-16 u, ventricose (broader below or in middle or above), sometimes ventricose-cylindric or clavate to ampullaceous or fusoid, usually with curved pedicel or with very short pedicel, sometimes capitate with mucro, always broadly rounded at tip even if this is attenuated towards a narrow apex, remarkably opaque in ammonia, refringent, eventually less opaque hyaline to yellowish hyaline in iodine, deeper rooting than basidia, mostly distinctly projecting beyond basidia, very numerous or rather numerous on sides and edges of lamellae, hyaline, sometimes some pale melleous (yellow in Melzer), smooth and entire; cheilocystidia differentiated, differing from the pleurobystidia in not being opaque, otherwise either similar to these (in one Amazonian collection) and 58-62 X 6.8-12.3 1, or else more inconspicuous, vesiculose hyaline, smooth, 11-28 X 6-10g. Hyphae: hymenophoral trama regular although some of the hyphae are interwoven, hyaline in ammonia, all hyphae with clamp connections and pseudoamyloid. Cortical layer: epicutis of pileus hymeniform or subhymeniform consisting of cells 10-48 X 4-39 ,, these cells isodiametric or subisodiametric, globose, vesiculose-clavate, laterally elongate, cordiform, erect ellipsoid, in the middle sometimes constricted or at apex mucronate, sessile or pedicellate but the pedicel often separated by a septum, hyaline, sometimes some brownish ones intermixed, smooth. Chemical reaction: very weak reaction of HCN (picric acid method).

On forest humus and on dead foliage (e g under Enterolobium, Allophylus) sometimes on very rotten wood, in moderately moist to moist subtropical forest but also in tropical rain forest, gregarious.

MATERIAL STUDIED. BRAZIL. Rio Grande do Sul: Pareci, 2 Nov 1951, Singer B 53 (LIL). ECUADOR. Napo, Sacha 4, 10 May 1973, Singer B 7336 (F). BOLIVIA. Pando: Manuripi, Conquista. 25 Mar 1956, Singer B 2220 (LIL). ARGENTINA. Tucuman: 10 km south of Tapia, 10 Feb 1951, Singer T 1193 (LIL), paratype, T 1191 (LIL), paratype; Sierra de San Javier, 14 Apr 1951, Singer T 1479 (LIL) type, 4/5 Mar 1949, Singer T 184, T 277 (LIL), 9 Jan 1959, Singer T 3210 (F).

The Ecuadorian material was remarkable for having particularly large carpophores and smaller spores (mostly 5-6 X 3 ,), but was not otherwise separable.

ILLUSTRATION. Singer (1965) pl 12, fig 66; pl 13, fig 67; (1958a) fig 16.

Species incompletely known:

Marasmius ardosiacus Montagne, from Bolivia. No material found. Marasmius auriformis Hennings, from Brazil. No material found. Marasmius bruchianus Spegazzini, from Argentina. Material insufficient for study. Marasmius cantharelloides Hennings, from Brazil. Type destroyed. Marasmius clitocybiformis Hennings, from Brazil. Type destroyed. Marasmius cyperinus Spegazzini, from Argentina. Type insufficient for study. Marasmius fulvus Montagne in Saccardo, a nomen confusum, see Dennis (1951b) p 157. Marasmius ochraceopapillatus Hennings, from Brazil. Type destroyed. Marasmius plumieri (LUveill6) Singer = Heliomyces plumieri Ldveille'.

A type specimen of LUveillM has not been discovered by me but what Patoui-




Hard determined so, comparing it with Plumier pl 168 apparently from the West Indies, is material collected by Duss in the West Indies, evidently belonging in Marasmius section Globulares (FH). I have studied this material but too little could be found out about this species to insert it in the sequence of numbers here treated. However, I did establish that the spores are large (14.5-15.5 X 4.4-5 p,), the hyphae pseudoamyloid and the epicuticular cells hymeniformly arranged, vesiculose and about 14.5 p broad. Duss's material is said to have originally white

pileus which is now dirty liver brown in the herbarium; it has the aspect of a Mic-

romphale. These data make it possible to identify the fungus when rediscovered; therefore the species were included in the key, p 269. No. 319 may be too close.

Marasmius symbiotes Theissen. from Brazil. Type not found. Marasmius tortipes Berkeley & Curtis, from Cuba.

The type has been studied by Dennis (1951d) and myself (K, FH). The

very distant lamellae without noticeable intervenation and the habit would suggest that this is the same as Marasmius helionlvces Murrill. But the data obtained from the type are not fully convincing inasmuch as in my preparation the epicutis of the

pileus appeared to be partially destroyed and poorly rehydrating in KOH. Dennis did not find cystidia (which would be in agreement with Al. heliomyces) but I saw some cells which might be cystidia. A further study of the type and topotypical material will show whether this is an independent species or A. heliomiyces which, in this latter case, would become a synonym ofM. tortipes.

Marasmius weddellianus Montagne, from Brazil. Type not found; no material available.

7. Gloiocephala Massee, Grevillea 21: 34. 1892. TYPE SPECIES. Gloiocephala epiphylla Massee.

Carpophores stipitate or sessile, stipe if present, lateral, eccentric or central; hymenophore present or absent, if present vein-like, merulioid, or lamellate; pigment present or absent; epicutis hymeniform, without broom cells; cystidia in form of hymenial or derm-

atocystidia present, metuloid and/or oleo-cystidia, frequently capitate and/or extremely long; trama monomitic, inamyloid or more rarely partly pseudoamyloid, mostly with

clamp connections; basal mycelium present or absent. On both living and dead leaves, sticks, culms, wood of Cormophyta. See figure 118.

Key to the Sections and Subsections of Gloiocephala

1. Pileus covered with setoid hairs or thin-walled hairs up to 450j long; stipe central, on dead leaves and their petioles; spores 6 to 15 long. sect Longifimibriatae p 285.

1. Pileus and stipe covered with dermatocystidia or hairs not longer than 200 ; if any elements of the hymenium or the epicutis are longer than 100u they are capitate (dermato) cystidia, or else spores different. 2. Stipe or pseudostipe present, central to lateral; if spores are longer than 12.5 l, the Q 2; diverticulate elements in epicutis none or rare. On various kinds of sub-

stratum. 3. Spores <6 p long; carpophores spathula- or spoon-shaped; hymenophore mostly

venose, often merulioid, sometimes absent. sect Spathulariae p 300. 3. Spores >6,u long. sect Gloiocephala p 291.

4. Spores 11-18 X 4-7.2 p; on Monocotyledones. subsect Macrosporae p 291. 4. Spores not longer than 12.5 p, mostly smaller. On conifers and Dicotyledones.

5. Pileus pigmented or fungus growing on conifers; base of stipe insititious. subsect Religiosae p 294.

5. Pileus usually white or pallid (sometimes punctate brownish because of the resinous incrustations of some oleo-cystidia). Base of stipe insititious or not. On Dicotyledones. subsect Confusae p 296.



Gloiocephala 285

2. Stipe or pseudostipe none; carpophores sessile; spores (10-)12.5-14.5 X (4.5-)5.5-8, (Q = 2 or smaller); amyloid metuloids and/or diverticulate elements present in the epicutis. On Dicotyledones. sect Sessiles p 302.

Gloiocephala sect Longifimbriatae Singer, sect nov Pilis longissimis praesentibus; sporis 15 p haud longioribus nec 6 brevioribus. TYPE SPECIES. G. longifimbriata Singer.

Key to the Species of Section Longifimbriatae

1. Spores more than 10.2 p; hairs often or always pigmented at least near the base. 2. Spores 12-15 X 3 p; hairs to 450 p long and 5-7 p broad; stipe about three times the

diameter of the pileus. 1. G. occidentalis. 2. Spores 10.5-13.5 X 3.5-4p; hairs 80-270 X 9-15u; stipe more than three times the

diameter of the pileus. 2. G. capillata. 1. Spores up to 12.5 p long, usually only up to 10.2j, long; setoid hairs pigmented or not.

3. Pileus white. 4. Lamellae well formed; pileus 4-5 mm broad; stipe central, 20 or more mm long.

3. G. longicrinita. 4. Lamellae none or replaced by a few vein-like lamellae; stipe, if central, up to 15

mm long. 5. Stipe central, rarely slightly eccentric. 4. G. longifimbriata. 5. Stipe lateral (see sect Gloiocephala).

3. Pileus cinnamon or brown. 5. Spores 8.2-10 X 4-6p; lamellae relatively well formed and rather broad; stipe with

basal mycelium. 5. G. lamellosa. 6. Spores 6.5-9 X 3-5p; lamellae vein-like, very narrow or very broad; stipe in-

sititious. 7. Lamellae vein-like; spores 3-3.8pu broad. 6. G. cinnamomea. 7. Lamellae broader than long; spores 3.8-5 ju, ellipsoid. 7. G. allomorpha.

1. Gloiocephala occidentalis (Dennis) Singer, comb nov Gloiocephala albocapitata var occidentalis (Dennis) Singer, Sydowia 14: 262, 1960. Crinipellis albocapitatus var occidentalis Dennis, Trans. Brit. Mycol. Soc. 34: 72. 1951. TYPE. Dennlis 414, from Trinidad. Pileus white, with scattered stiff brown setae, flat or depressed, 1-2 mm in diameter.

Lamellae white, 3-4, fold-like, adnate. Stipe red-brown, bearing numerous setae, very slender, wiry, cylindrical, about three times as long as the diameter of the pileus.

Spores 12-15 X 3 p, tear-shaped, hyaline. Hymenium: basidia about 20 X 5-6 u;

cystidia 25-30 X 8 , cylindrical, thin-walled, obtuse, often secreting a cap of gummy brown material (oleocystidia). Cortical layers: epicutis of the pileus formed by smooth

thin-walled, cylindrical or pyriform cells, somewhat pointed above, hyaline, 20-25 X 5- 9 p. Setae of pileus and stipe up to 450 p long, 5-7p broad, straight, stiff, tapering upwards, brown, with moderately thick walls.

Scattered on dead leaves in forest litter. MATERIAL STUDIED. None. The description is adapted from Dennis (1951a). Illustration see Dennis, (1970), Fig. 2 G. Marasmius minutissimus Peck with which Dennis compares his fungus is a temper-

ate North American species of Marasmius sect Epiphylli with smaller spores and smaller

cystidia; also lacks brown setae. ILLUSTRATION. Dennis (1951a) fig 9; (1970), fig 2 G.

2. Gloiocephala capillata Singer, sp nov Fig 97. TYPE. Singer M 89 79A, from Mexico. Pileo, lamellis, apice stipitis insititii albis; lamellis rudimentariis vel subnullis. Sporis




10.5-13.5 X 3.5-4 ,; cystidiis subcapitatis; pilis pilei 85-270 X 9-15 p, stipitis 80-230 longis. Ad folia emortua arborum -dicotyledonearum silvae tropicalis in Mexico. Typus in F conservatus est.

Pileus white, pilose, smooth and glabrous otherwise, not glutinous, applanate, about 1.5 mm broad. Lamellae scarcely developed or few and very narrow, hymenial surface white. Stipe white, below deep chestnut, pilose, tapering upwards, insititious, about 24 X 0.1 mm. Context very thin, white.

Spores 10.5-13.5 X 3.5-4 p, fusoid, smooth, hyaline, inamyloid. Hymenium: basidia about 19 X 7 u,, 4-spored; cystidia 30-40 X 7-8 p, ventricose-ampullaceous with rather narrow ventricose portion a subcapitate-swollen at the apex, constriction below apex 1-4. 5 p wide, capitulum about 4 p wide, wall hyaline, thin or firm, inamyloid. Hyphae hyaline, in lower portion of stipe tawny and thick-walled, inamyloid, with clamp connections. Cortical layers: epicutis of pileus consisting of hymeniformly arranged cells, + 23 X 12-20 p, some more elongated and reaching 30 X 20 p, thin-walled to rarely moderately thick- walled, vesiculose or cylindrical; among these some similar elements 20-26 X 8.5-11 p with the wall 2-2.5, at the apex, tawny, below hyaline and rather thin-walled: dermatocystidia 35-44 X 9.5-1 1 p, ventricose and capitate, constriction 4-6 p across, capitulum 7-9 , in diameter, often incrusted by hyaline to brownish resinous crusts; hairs 85-270 X 9-15 , below swollen and sometimes fulvous, thick-walled otherwise gradually thinned to 4-6 1 and at apex about 1.5 p, the larger upper portion with gradually thinner wall, not acute, inamyloid; covering of the stipe with the same elements, i e numerous dermatocystidia but these sinaller and more like the short epicuticular elements forming all transitions to dermatocystidia 7-16 X 3-7.5 p, erect, clavate, cylindric, subcapitate or simply obtuse, with rather thin, firm wall, hyaline to pale straminous, inamyloid. Hairs like those of the pileus, but with often shortened base and often shorter effilate portion (80-230 p long).

On dead fallen leaves of dictyledonous trees in tropical forest. Mexico. MATERIAL STUDIED. MEXICO. Chiapas, between Finca Sospiro and El Pozo. 4 Aug 1969.

Singer M 8979A (F), type. This species is closely related to the preceding one, but differs in somewhat smaller

spores, hyaline, slightly shorter hairs and longer stipe.

3. Gloiocephala longicrinita Singer, sp nov Fig 98. 99. TYPE. Singer B 6282, from Colombia. Pileo albo; lamellis bene evolutis; stipite albo, atro ad basin, 20 mm longiore, in-

sititio. Sporis 6.5-8 X 3-4 p; cystidiis nullis; cheilocystidiis haud manifeste capitatis sed resinaceo-incrustatis incrustatione fulva, crassiuscula, fugaci, interdum in multis cellulis vix visiblili; pilis pilei usque ad 130 p longis et usque ad 18 p latis; eis stipitis usque ad 300,u longis. Ad petiolos foliorum in Columbia. Typus in F conservatus est.

Pileus sordid white, dried whitish with brown dots (under a lens), pilose, convex, about 4-5 mm broad. Lamellae white, about 8 through-lamellae and 3 lamellulae, not intervenose, narrow to medium broad, distant, adnate. Stipe white with blackish base, pilose, the hairs projecting about as much as the diameter of the stipe, equal, insititious, somewhat over 20 mm long and about 0.2 mm thick.

Spores (6-)6.5-8 X (2.8-)3-4 p, ellipsoid, smooth, hyaline, inamyloid, eventually at times developing a cross septum. Hymenium: basidia 15-20 X 5.5-6.2 p, 4-spored; cystidia none; cheilocystidia making the edge partly heteromorphous, 35-45 X 5.5-10.5 ,L, characteristically narrowly ventricose or narrowly club-shaped or subcylindrical, with hyaline thin wall but many with a fugacious thick resinous incrustation at the apex which



Gloiocephala 287

is fulvous. Hyphae mainly thin-walled, loosely arranged and gelatinized, filamentous, (1-3 p broad), mainly radially arranged, in the stipe parallel hyaline in the apex, brownish melleous to brown below, medium thick-walled and non-gelatinized; all hyphae with clamp connections, inamyloid. Trama monomitic. Cortical layers: epicutis of the pileus consisting of a hymeniform layer of clavate to vesiculose obtuse elements (e g 24-26 X 10-18 u) which are hyaline with thin, rarely somewhat thickened (to 1 u) wall; these interspersed with dermatocystidia 14.5-24 X 6.5-7.5 p which differ in being thick-walled (wall fulvous to fulvous-brown, 1.2-2 p thick) with intraparietal pigment, the wall thicker at the apex, mostly clavate-ventricose, not projecting, even partly imbedded in the hypodermial layer which is non-gelatinous; hairs 50-130 X 13.5-18 u, with broadened base and effilate long apex (2-6 p across), at the base often thick-walled (wall 1, rarely 1.5 , thick) but the larger effilate portion thin to firm walled, base subhyaline to melleous, apical portion hyaline or subhyaline; surface elements of the stipe not hymeniformly organized but the same type of elements found here as on the pileus, the thick-walled ones 6-30 X 3-11 p,, versiform, melleous, sometimes ampullaceous, mucronate or capitate; hairs here like those of the pileus but reaching 300 u in length with the base broadened to only about 9 u.

On leaf petioles. Solitary. Western Colombia. MATERIAL STUDIED. COLOMBIA: Valle, Buenaventura, San Joaquin, 22 Apr 1968, Singer

B 6282 (F), type.

4. Gloiocephala longifimbriata Singer, Sydowia 14: 266. 1960. TYPE. Singer T 3520, from Argentina. Pileus white, macroscopically appearing glabrous but conspicuously long-pilose

from numerous but scattered hyaline hairs at least near the margin where they are at times rather dense and make the margin appear fimbriate, smooth, rarely indistinctly sulcate in age, convex, broader than high, 1-4.5 mm broad. Lamellae developing late, at first lower side of pileus with smooth hymenial surface, later lamellae almost veniform and eventually mostly at least some of them distinctly lamellar (about 0-7 lamellae in mature specimens) but some or all of them remaining venose or not reaching the margin of the pileus or the stipe, mostly simple, white, sometimes drying cream, up to 0.4 mm broad, about 80-90 u thick, attingent to subdecurrent, not collariate. Stipe umber, fuliginous, or black, at first with white apex, macroscopically glabrous but under a lens pilose, in age sometimes glabrescent, filiform, insititious, 4-15 X 0.1-0.3 mm, at times accompanied by dark rhizomorphs. Contex white, very thin, inodorous.

Spores (6-)7.5-10.5 X (2-)3.3-4.8(-5.2) Q = 2-2.2 or slightly larger, more rarely slightly smaller), ellipsoid to fusoid, with or without suprahilar depression, smooth, hyaline, inamyloid. Hymenium: basidia 22-31 X (3.5-)6-9.5 ,, clavate, 4-spored, sometimes some 2-spored ones intermixed; basidioles fusoid, then clavate; cystidia 21-50 X 6-11.7 p, ventricose below, with a constriction (2.2-5 p across) underneath a capitulum (2.5-7.5 , in diameter), some merely capitate or ampullaceous, with firm to thick (0.3-2 u, mostly 0.3-0.5 ,), hyaline wall and occasional resinous incrustation, especially on the capitulum (incrustation pale succineous to subhyaline where present), varying from extremely scarce to numerous; besides there may or may not be one or another long thick-walled hair (like on the epicutis and the stipe) in the hymenium, and/or some thin-walled paraphysis- like hyphal ends. Hyphae of the pileus-trama and the hymenophoral trama moderately dense but only partly very weakly gelatinized, thin-walled, hyaline, inamyloid, with clamp connections, but many secondary (clamp-less septa observed), filamentous, radially ar-




ranged in the pileus, regularly arranged in the hymenophoral trama, Subhymenium in

places rather broad, of irregularly interwoven, small, short-filamentous elements, not gelatinized, hyaline. Hyphae of the stipe mostly thick-walled, inamyloid to weakly pseudoamyloid, strictly parallel, in lower part of the stipe mostly pigmented. Cortical layers: epicutis of the pileus hymeniform or subhymeniform, consisting of vesiculose elements which are hyaline, in places sometimes melleous-hyaline, with thin to firm, exceptionally up to 1 p thick wall, 15-37 X 7.5-25 p, interspersed with dermatocystidia which are ventricose below and often capitate, subcapitate, ampullaceous or mucronate, 10-50 X 6.5- 12 ,, constriction or neck 2.5-5.5 ,, capitulum where present 4.5-9.5 in diameter, these often granularly or resinously hyaline to succineous incrusted: aside from these, also long Crinipellis-type hairs present, scattered to rather dense and along the margin of the pileus usually rather numerous, these 33-240(-500) X 3-8.5 A or 57-365(-500) X (2.5-)3-9.5 , many with swollen base (7-18 wide) and slightly narrowed to long-effilate apex, the latter entirely or in the lower portion thick-walled (wall 0.5-2 p1 thick), hyaline, sometimes melleous-hyaline, some with resinous or mucilaginous-granular, amorphous incrustations, inamyloid, in some hairs with an inner wall-layer very slightly pseudoamyloid or an outer layer amyloid; vestiment of the stipe consisting of similar hairs, some of these as long as about 1 mm, the lower portion often pale melleous, the apex often long-effilate and acute, 11-70(-300) X (at base) 3-4 p, wall at base inamyloid or slightly amyloid, wall 0.3-0.7 , thick; near apex hairs with thinner wall and swollen base. 4-25 X 2.5-6 p, cylindric, obtuse-conical or pulvinate, wall hyaline, 0.2-0.4 1thick.

On leaves and petioles as well as herbaceous stems of all kinds of hosts (Pteridophyta to Myrtaceae) in tropical-montane or fog forest. subtropical-montane, and rain forest of the plains, also in inundated forest on earth-covered foliage, solitary or gregarious.

MATERIAL STUDIED. BRAZIL. Serra dos Orgoos. 1500 in alt. 25 Oct 1961, Singer B 4033 (F). ECUADOR. Napo, Lago Agrio, 8 May 1973, Singer B 7274 (F). BOLIVIA. Pando: Man- uripi, Santa Rosa, 22 Mar 1956, Singer B 2146 (LIL): La Paz: Nor-Yungas. Carmen Painpa, 200 m alt, 26 Feb 1956, Singer B 1535 (LIL). ARGENTINA. Tucuman, between Ciudad Universitaria and Estancia San Javier, 900-1000 m alt, 9 Jan 1960, Singer T 3520 (LIL). type.

ILLUSTRATION. Singer (1960), pl. 38, fig. 5; pl. 39, fig. 6. The amyloidity and distribution as well as the size of the metuloid hairs of the

pileus and the stipe is apparently somewhat variable, and it is possible that some varieties may be distinguished. The amyloid reaction of these bodies was observed in the collection B 4033; the hairs were much more evenly distributed up to the apex of the

stipe in coll~ction B 1535 (on Pteridophyta). The size and distribution of the cystidia also appears to be variable.

5. Gloiocephala lamellosa Singer, Sydowia 14: 278. 1960. TYPE. Singer B 15285, from Bolivia. Pileus brown ("Alamo" to "Arab" M&P), smooth, beset with hyaline stiff long

hairs all over (when seen under a lens), unicolorous and not changing when dried, neither papillate nor umbilicate, conic-campanulate, often slightly flattened at apex, with

straight margin, 1.5 X 1.5 mm. Lamellae white with white edges, few (5-6), distant, equal or with one lamellula, moderately broad, not pilose under a lens. Stipe brown, setose, filiform, hirsute from scattered long, hyaline hairs, almost shining, smooth, 18- 23 X 0.1-0.2 mm; basal mycelium varying between weakly developed to well developed in form of radiating white mycelial fibrils around the base of the stipe. Context paler than the surface to pallid, very thin, inodorous.

Spores 8.2-10 X 4-6 p, ellipsoid, smooth, hyaline, inamyloid. Hymenium: basidia 24.5-30.5 X 4.7-8.2 ,L, clavate, hyaline, 4-spored, basal septum clamped; cystidia present



Gloiocephala 289

but very scattered in the hymenium, 31-5-44 X 6.8-7 ,, ventricose with capitate apex, constriction underneath the capitulum about 2 A, across, capitulum about 4.8-5 , diameter, wall thin, hyaline, sometimes found only rarely and on or near edge of lamellae. Hyphae: subhymenium subcellular, consisting of very irregular, small elements, hyaline, not gelatinized. Hymenophoral trama regular, consisting of hyaline to stramineous, thin, filamentous, interwoven hyphae which are not or scarcely pseudoamyloid. Hyphae of the stipe parallel, the thin-walled ones mostly inamyloid or scarcely pseudoamyloid, the thicker-walled ones in superposition weakly pseudoamyloid, with clamp connections, but many clampless secondary septa present. Cortical layers: epicutis of pileus consisting of a hymeniform layer of elements 16-25 X 7.5-9 A,, these fusiform to ventricose or vesiculose-subclavate, smooth, non-incrusted, thin-walled; in distances between 25 and 40 p, there are dermatocystidia interspersed, these 24-29 X 10-14 1, varying from ventricose- subampullaceous and short-pedicellate to vesiculose-subclavate, thick-walled, cinnamon. stramineous to melleous; furthermore, there are also long-setose hairs, reminding one of those of Crinipellis, these (37-)130-255 X 8-16 u, not very numerous but very conspicuous, broadest just above the hyaline to melleous base, equal above and there hyaline, more rarely slightly tapering upwards, with obtusely rounded tip, apex (5-)8-10 , broad, aseptate, wall moderately thick to thick (if thick: 2-6 j in diameter), thickest just above the broadest portion of the hair, not incrusted, not or not distinctly pseudoamyloid but often slightly amyloid; covering of the stipe consisting of the same hairs but these rather scattered, between them surface layer of hyphae not differentiated from the other cortex-hyphae of the stipe.

On dead fallen dicotyledonous leaves in tropical-montane forest, Yungas. MATERIAL STUDIED. BOLIVIA. La Paz, Nor-Yungas, Carmen Pampa, 200 m alt, 26 Feb

1956, Singer B 15285 (LIL), type. ILLUSTRATION. Singer (1960), pl. 38, fig. 4.

6. Gloiocephala cinnamomea Singer, sp nov Fig 100. TYPE. Singer, M 8971, from Mexico. Pileo cinamoneo, radiatim rugoso, piloso e pilis 200-380 X 7-13.5 ,, 6 mm cir-

citer lato; lamellis veniformibus, pallide cremeis: stipite cremeo, ad basin castaneo vel castaneo-fuligineo, subtiliter pubescente sub lente, insititio. Sporis 7-9 X 3-3.8 u; cystidiis frequenter subcapitatis; dermatocystidiis plerumque capitatis. Ad folia mortua arborum dicotyledonearum in silva, Mexico. Typus in F conservatus est.

Pileus cinnamon, radially rugose, not glutinous, pilose with hyaline hairs, especially at the margin, convex, soon flat, about 6 mm broad. Lamellae in form of veins which are

light cream colored, narrow and obtuse, eventually up to seven through-veins and 1-2 shorter ones, the through-veins occasionally forked, but not intervenose, under a lens brown punctulate at the ridges, pilose near the margin of the pileus, collariate or subcollariate. Stipe cream colored, chestnut to fuliginous below, finely pubescent under a lens, subequal and filiform, insititious, 17-18 X 0.05-0.1 mm. Context pale cinnamon in the pileus, dark cinnamon in the cortex of the stipe, inodorous.

Spores 7-8.5(-9) X 3-3.5(3.8) ,, ellipsoid to oblong, more often oblong when mature, thin-walled, hyaline, smooth, inamyloid. Hymenium: basidia about 19 X 5 u, 4-spored; cystidia 27-40 X 6.8-12 ,, either thin-walled or with thick walls and then opaque-glassy with the inner surface of the wall poorly marked in KOH, ammonia, and Melzer reagent, ventricose ltelow, constricted to a broad neck and very frequently subcapitate; cheilocystidia like the pleurocystidia or larger (transition to hairs). Hairs often scattered or restricted to the margin of the pileus, (75-)200-380 X 7-13.5 1, be-




low often with brown and thick-walled, hyaline and thin-walled (0.2-0.5 ju) in the long effilate portion, there often covered by a brown resinous incrustation, base of the ef- fidate portion about 5-6 wide, apex 1.2-2.5(-3) wide with rounded tip, inamyloid. Hyphae with clamp connections, inamyloid, parallel in the stipe, thin-filamentous and not gelatinized in the pileus, hyaline or subhyaline. Cortical layers: epicutis of the pileus hymeniformly arranged consisting of thin- to moderately thick-walled vesiculose elements which may also be ampullaceous or ventricose, 18-27 X 10-13.5 u, hyaline to brownish; among these short thick-walled elements, 12-19 X 6.5-10 ,, distributed in almost irregular pattern among the basic elements, broadly clavate, wall increasingly thicker towards the apex and reaching a thickness of 2 p, also increasingly more pigmented there (deep chestnut to tawny), smooth and broadly rounded above: derm-

atocystidia much like the cystidia of the hymenium, 20-32 X 5.5-10p, often more strongly capitate (constriction 2-3.5 p across, capitulum 4-6.5 l wide), with a wall up to 1 thick. Hairs conspicuous, as described above.

On dead leaves of dicotyledonous trees in tropical forest. Solitary. Mexico. MATERIAL STUDIED. MEXICO. Chiapas, between Finca Sospiro, and El Pozo 4 Aug

1969, Singer M1 8971. (F), type.

7. Gloiocephala allomorpha Singer, sp nov Fig 102. TYPE. Singer B 6451, from Colombia. Pileo ochraceo-brunneo, longe hyalino-piloso e pilis 150-260 X 9-13;:' lamellis

eximie latis (latioribus quam sunt longae!), cremeis, adnatis. stipite pileo concolori vel obscurius brunneo, e pilis hyalinis crinito, 12-23 mm longo, insititio. Sporis 6.5-8 X 3.8-5 p, cystidiis nullis; oleocystidiis acieis capitatis. Ad folia dicotyledonea emortua in Columbia. Typus in F conservatus est.

Pileus ochraceous brown with long hyaline erect hairs, otherwise glabrous, sulcate conic to campanulate and often papillate, 2-4 mm high, 0.8-2.8 mm broad. Lamellae extraordinarily broad (broader than long) for the genus, cream color, adnate, not collariate. Stipe concolorous with the pileus or deeper brown, beset with hyaline, erect hairs, apex concolorous with lamellae, insititious, 12-23 X 0.1-0.2 mm. Associated with black rhizomorphs which are fuscous blackish, pilose but glabrescent. Context extremely thin in the pileus, hyaline or ochraceous, inodorous.

Spores 6.5-8 X 3.8-5 p, ellipsoid, thin-walled; hyaline, smooth, inamyloid. Hymen- ium: basidia 17-18 X 5.5-7 u, 4-spored; basidioles fusoid; cystidia none; cheilocystidia on and near edge 26.5-28 X 5-6.8(10)i1, not making the edge heteromorphous but intermixed with basidia and basidioles, fusoid-ventricose with capitate apex (3 p in diameter), with hyaline rather thin wall but often incrusted by a yellow-brown resinous sheet-like

covering, inamyloid, Hyphae filamentous, thin, thin-walled, not gelatinized, with clamp connections (not many secondary septa observed) hyaline and radially arranged in the pileus; hymenophoral trama regular. Cortical layers: epicutis of the pileus hymeniform consisting of smooth (rarely with knob-like appendage), elements, these short-clavate (20-25 X 0-11 uor more frequently ventricose to ampullaceous (20-27 X 6-8.3 u) or more frequently ventricose to ampullaceous (20-27 X 6-8.3 p), hyaline or in the lower portion or entirely brownish yellow to brass color, with thin to moderately thick (to 1 u) walls, inamyloid, pigment intraparietal. Hairs (18-)150-260 X (5-)9-13 u, long-cylindrical or more often slightly and gradually tapering upwards to an acute to rounded-obtuse tip, thick-walled, the walls hyaline and (l-)4(-7) j in diameter even in the upper portion. Similar hairs also on the surface of the stipe.

On dead (dicotyledonous) leaves in tropical-montane forest, gregarious.



Gloiocephala 291

MATERIAL STUDIED. COLOMBIA. Valle: Corr. Vergel, Mun. Dagua, N of KM 25 of Dagua- road, about 1950 m alt, 28 Apr 1968, Singer B 6451 (F), type.

Gloiocephala sect Gloiocephala Extremely long hairs absent; pileus not sessile; spores larger than 6 u. TYPE SPECIES. Gloiocephala epiphylla Massee

Gloiocephala sect Gloiocephala subsect Macrosporae Singer, subsect nov Pilis eximie longis nullis; sporis sat magnis (11-18 X 4-7.2 p). Ad Monocotyledones. TYPE SPECIES. Gloiocephala inobasis Singer.

Key to the Species of Subsection Macrosporae

1. Pileus white, eventually ochraceous or with brown dots, smooth when dried; hymenial surface smooth or venose: stipe central or eccentric, over 8 mm long.

2. Spores 11.5-18 X 4-4.3(-4.8)p; stipe with basal hyphal fibrils; metuloids strongly incrusted. 9. G. inobasis.

. Spores more elongated, 15-20 X 3-4p; stipe with more numerous metuloid hairs at base but otherwise insititious; metuloids little or not incrusted. 8. G. longisperma.

1. Pileus orange, hymenophore consisting of veins or narrow lamellae; stipe sublateral to lateral, less than 5 mm long. 10. G. palmaruim.

8. Gloiocephala longisperma Singer, sp nov Fig 101. TYPE. Singer B 4103, from Brazil. Pileo albo, demum ochrascente, levi, pilosulo sub lente; lamellis nullis (superficie

hymeniali levi), dein venis perpaucis frequenter obviis; stipite albo, ad basin nigro. Sporis 15-20 X 3-4 ,, fusiformibus; cystidiis sparsis; dermatocystidiis typi oleocystidiorum; pilis 100-200 p longis in pilei stipitisque superficiebus obviis. Ad folia monocotyledonea in Brasilia Australi. Typus in F conservatus est.

Pileus white, eventually ochraceous, smooth, pilose under a lens, 2-3 mm broad. Lamellae practically none, only some few distant veins present in mature carpophores, hymenial surface smooth. Stipe white at apex, blackish below, dried deep blackish chestnut, macroscopically glabrous, under a lens with scattered erect hairs, these more numerous at the otherwise insititious base, not connected or accompanied by rhizomorphs, equal, often eccentric, about 12 X 0.2 mm. Context very thin, inodorous.

Spores 15-20 X 3-4 p, fusoid or fusoid-oblong, in profile the inner side applanate to concave, thin-walled, smooth, hyaline, a few overaged ones brownish, inmayloid. Hymenium: basidia 22-30 X 7.5-8 ,, 4-spored; cystidia scattered, 30-40 X 6-6.5 A, ventricose with tapering apex, or slightly constricted in the middle, or irregularly cylindrical, hyaline. Hyphae hyaline, near surfaces somewhat to strongly pigmented, thin-walled, some slightly thick-walled, radially arranged in the pileus, parallel in the stipe, not gelatinized anywhere, with clamp connections, inamyloid. Cortical layers: epicutis of the

pileus hymeniform, formed by smooth, non-incrusted, hyaline, inamyloid, vesiculose elements 16-30 X 12-20 p; among these some thick-walled hyaline and inamyloid elements, with deep succineous rusty-chestnut intraparietal pigment, 12-23 X 8.5-9.5 j (wall to 3 p thick). Interspersed with these elements hairs 100-200 pu long, base swollen and 8.5-10.2 p broad, here thin-walled or at least the lower portion thin-walled, the effilate apex 4-5.5s broad and thick (1-2 p) walled, hyaline, some pale brownish, tip sometimes again thin-walled, acute or more often obtuse. Surface of the stipe beset with similar hairs but these scattered and only near the base of the stipe crowded and then

pale melleous brown and often lacking the swollen base.




On monocotyledonous (probably bromeliaceous) dead leaves in the forest, gregarious.

MATERIAL STUDIED. BRAZIL. Sao Paulo, Serra da Cantareira, Reserva Florestal, 17 Jul 1964, Singer B 4103 (F), type.

This species should not be confused with G. occidentalis which has longer brown hairs and somewhat smaller spores.

9. Gloiocephala inobasis Singer, Sydowia 14: 277. 1960. TYPE. Singer B 2296, from Bolivia. Pileus pure white but when dried with brown dots from the exudations of the met-

uloids, not orbicular in outline but with a sinus on one side which does not reach the center, smooth, glabrous macroscopically, convex, fresh about 9 mm broad. Lamellae none, veins about four, radial, the whole hymenial surface showing small brown dots from the exudations of the metuloids, otherwise white. Stipe fresh whitish in upper, dark brown in lower portion, distinctly fibrillose under a lens from the dermatocystidia, equal, appearing insititious macroscopically but under a lens rising from a socle on a blackened spot of the substratum and basal mycelium well visible as radiant and somewhat ascendant brown fibrils, about 11 X 0.4 mm, socle up to 2 mm broad, upper portion of stipe straight or oblique, eccentrically attached to the pileus so that carpophores appear spathuloid. Context white, thin, inodorous.

Spores 11.5-18 X 4-4.3(-4.8) p, oblong to subfusoid, with a slight suprahilar de-

pression or somewhat curved in lower portion, thin to firni walled, smooth, hyaline but

browning in contact with the cystidial exudations, inamyloid. Hymenium: basidia 30- 34 X 8-8.5 u, clavate, 4-spored, with clamped base; cystidia metuloid, 40-135 X 16-28 p,

conspicuous but moderately numerous, with often more or less thickened wall (wall 0.3-2 p thick), hyaline and not strongly colorable in cresyl blue mounts but covered with

amorphous hyaline to succineous brown resinous material which turns deep violet to bright blue in cresyl blue mounts, with (in NH40H,) colorless indistinct contents which

may turn partly deep blue in cresyl blue mounts, very weakly pseudoamyloid in the innermost wall or inamyloid, clavate to clavate-subcylindric, often with a slight constriction underneath the apex and thus almost subcapitate, strongly projecting beyond the basidia and deeper rooting than these. Hyphae: subhymenium consisting of ir-

regularly interlaced small hyaline elements; trama of the pileus consisting of irregularly arranged elements which are mostly strongly elongated, not gelatinized, with often

slightly thickened wall, hyaline, inamyloid, with clamp connections: hyphae of the

stipe parallel, inamyloid and not metachromatic in cresyl blue mounts, with clamp connections. Cortical layers: epicutis of the pileus hymeniform, consisting of isodiametric cells with or without a pedicel, some vesiculose with firm wall, smooth, 14-44 X 9-13 p, some slightly pseudoamyloid in the innermost wall layer, but mostly practically inamyloid; dermatocystidia few, intermediate in appearance between the epicuticular vesiculose cells and the metuloids of the hymenium, somewhat larger than the former and also exuding a brown matter. Vestiment of the stipe of numerous, at

places almost crowded dermatocystidia of the metuloid type much like the hymenial metuloids, not forming a continuous palisade.

On monocotyledonous leaves in inundated rain forest fruiting after recession of the water.

MATERIAL STUDIED. BOLIVIA. Pando, Madre de Dios, Santa Teresa, 28 Mar 1956, Singer B 2296 (LIL), type.




Gloiocephala 293

10. Gloiocephala palmarum Singer, Sydowia 14: 277. 1960. TYPE. Singer B 2093, from Bolivia. Pileus pale orange cinnamon, ochraceous orange, sometimes more whitish to och-

raceous, or cinnamon, but usually some pilei in a population showing distinct orange pigmentation, e g 11 G 7 (M&P), dried near 12 F 9 (M&P), finely and minutely pallid pilose under a lens but macroscopically glabrous, smooth but eventually over half the radius sulcate or rugulose all over, with a suborbicular outline but with a deep sinus- like or heart-shaped incision on the side where the stipe is attached, convex, then ir-

regularly repand, becoming flattened, 1-5 mm broad. Lamellae white, very distant, very narrow, often almost vein-like, up to seven lamellae, of these up to four through-lamellae, the others mostly vein-like and sometimes forking with the through-lamellae, with rather obtuse edges, subfree to adnate. Stipe brown, chestnut or black, at first with'a white or orangy apex, often oblique or appressed to the substratum or curved above, attached to the pileus at the edge of the sinus or very near the edge, i e lateral or sublateral, usually forming an angle near the point of attachment, macroscopically glabrous but equally pilose as the pileus under a lens, equal or more rarely slightly tapering upwards, solid, insititious. Context white, very thin, inodorous.

Spores (11-)12.5-13(-16.5) X (4.5-)5(-7.2) +? 5p, in purely 4-spored carpophores, oblong with thin to firm wall, smooth, inamyloid. Hymenium: basidia 27-31 X (7-) 9-1 1.5 ,, 4-spored, more rarely 2-4-spored; cystidia rather scattered, 38-51 X 7.5-12 u, ventricose with subcapitate or capitate apex, constriction 3-5 p across, capitulum 4.5-7 p, diam., wall thin, more rarely somewhat thickened in the lower portion, hyaline, in-

amyloid: cheilocystidia and metuloids not differentiated, of thin, interwoven hyphae, hyaline in ammonia, not gelatinized, inamyloid, those underneath the epicutis often with orangy wall. Hyphae of the stipe cortex chestnut brown, thick- or thin-walled, strictly parallel with each other, filamentous, inamyloid, few extremely weakly pseudoamyloid in some carpophores, those of the more hyaline core inamyloid, all with clamp connections. Cortical layers: epicutis of pileus hymeniform or palisadic consisting of smooth broad elements, these (8-)14-47 X (5-)9-28 p, vesiculose, globose or pear shaped, with hyaline to more often orange thin to somewhat thickened (to 1 sp) inamyloid wall, often mucronate, some with a pedicel up to 6 long, intermixed with two kinds of

dermatocystidia: (a) capitate dermatocystidia 17-55 X 6-14.5 ,, constriction 1.5-3 p,

capitulum 3-4.5 p in diameter, rarely absent and then with a long filamentous mucro instead of the capitulum or with an acute conic apex, with the wall firm or thin, rarely up to 1.3 pin diameters: (b) metuloids 55-135 X (at base:) 8.5-31 p, in the long hair- like apex 4.5-7 , wide, with mostly rounded tip, rarely subcapitate, hyaline to stramineous with a wall 1-4 p thick, inamyloid to weakly pseudoamyloid or amyloid, strongly projecting, numerous to scattered, often more numerous along the edges of the pileus, always with widened base and subcylindric apex, most frequently between 80 and 115 p

long; covering of the stipe consisting of scattered to numerous (but not forming a continuous palisade) metuloid hairs like those of the pileus, usually however somewhat smaller and neither pseudoamyloid nor amyloid.

On monocotyledonous, mainly palm leaves in tropical rain forest. Amazonas region.

MATERIAL STUDIED. ECUADOR. Napo, Francisco de Orellana (Coca), 12 May 1973, Singer B 7377 (F). BOLIVIA. Beni, Vaca Diez, Guayaramerin 17 Mar 1956, Singer B 2093 (LIL), type; 8 Mar 1956, B 1 744 (LIL).

This species has the faint amyloid reaction of some metuloids in common with

Gloiocephala anastomosans which differs however in having metuloids with a more con-




sistently amyloid reaction, and the lack of a stipe as well as the habitat. ILLUSTRATION. Singer (1960), pl 37, fig 2; pl 38, fig 3.

Gloiocephala sect Gloiocephala subsect Religiosae Singer, subsect nov Pileo maturitate pigmentato; sporis eis specierum subsectionis Macrosporarum sim-

ilibus; lamellis in specimininus omnino maturis praesentibus; stipite insititio; generatim ad coniferas.

Pileus when mature pigmented; spores like those of subsection Macrosporae; lamellae in mature carpophores present; stipe insititious; generally on conifers.

TYPE SPECIES. Gloiocephala religiosa Singer.

Key to the Species of Subsection Religiosae

On leaves of Podocarpus; spores 6.5-7.5 X 2.5-3 p; dermatocystidia of the stipe with a long tail-like effilate apex. 11. G. podocarporum. On dead branches of twigs; spores reaching larger size: dermatocystidia of the stipe not long-effilate. 2. Stipe 4-7 mm long; many cystidia and dermatocystidia thick-walled; Mexico. 12. G. religiosa. 2. Stipe 1.5-2 mm long; most or all cystidia and the dcrmatocystidia of the stipe thin-

walled; Chile. 13. G. ciliata.

11. Gloiocephala podocarporum Singer, sp nov Fig 103. TYPE. Singer T5300, from Argentina. Pileo albo dein luteolo vel pallide cinnamomeo, campanulato vel convexo, 0.5-1.5 mm

lato; 1 mellis primum nullis, dein 6-8, angustis, admodum distantibus; stipite albo, basin versus dilute vel atrius castaneo, insititio, usque ad 31 mm longo. Sporis 6.5-7.5 X 2.5-3 p; cystidiis in et prope aciem lamellarum capitatis vel subcapitatis; dermatocystidiis stipitis appen- diculo longo tenuissimo effilatis. Ad folia Podocarpi in Argentina septentrionali. Typus in F conservatus est.

Pileus white, becoming yellowish or pale cinnamon, smooth, later becoming sulcate with as many sulci as there are lamellae, finely pubescent under a lens, campanulate to convex, 0.5-1.5 mm broad. Lamellae at first none, but when fully mature 6-8 narrow lamellae

present, these (and the smooth hymenial surface) concolorous with the pileus, very distant: adnexed to adnate. Stipe white, below light to dark chestnut, finely pilose under a lens, insititious, usually slightly attenuate towards the apex, up to 31 mm long, about 0.1 mm thick. Context very thin, inodorous.

Spores 6.5-7.5 X 2.5-3 , fusoid, smooth, hyaline, inamyloid. Hymenium: basidia about 15 X 5 , 4-spored; cystidia on and near edge of lamellae, 17-25 X 5-8 $A, ventricose- fusoid but with subcapitate to capitate apex (this 2.5-3 pLwide) thin-walled, hyaline. Hyphae filamentous, hyaline, inamyloid, with clamp connections, thin-walled, non-gelatinized. Cor- tical layers: epicutis of the pileus hymeniform, formed by subisodiametric to ellipsoid or fusoid cells which are 9-15 X 4-7 , hyaline or subhyaline in KOH, with thin to slightly thickened wall, inamyloid; among these some thick-walled elements which are more elongated (20-27 X 6.5-8 4), with succineous-tawny wall 0.6-4 ,thick, gradually becoming thinner near the base of the cell, fusoid or ampullaceous, more rarely with a constriction in the middle and almost subcapitate; dermatocystidia numerous, much like those of the hymenium; covering of the stipe consisting of numerous erect hairs which have a thick-walled, subfusoid to subulate base, 9-37 X 3.5-5 ,lapex effilate up to 50 ,long and 1 pwide, long



Gloiocephala 295

tail-like, thin-walled and acute, entire hair hyaline or subhyaline, wall in basal portion 0.6- 1.5 Mthick, rarely effilate portion missing.

On leaves of Podocarpus parlatorei. MATERIAL STUDIED. ARGENTINA. Jujuy, 15 km NE of Yala, 20 Feb 1966, Singer T 5300

(F), typc.

12. Gloiocephala religiosa Singer, sp nov Fig 105. TYPE. Singer MA O1, from Mexico. Pileo aurantiaco-ferrugineo vel brunneolo-ochraceo, marginem versus interdum pall-

idiore vel albido, plus minusve sulcato in vegetis, 1-6 mm lato; lamellis albis, distantibus, primum veniformibus, demum usque ad 8 lamellis percurrentibus praesentibus; stipite insititio, 4-7 mm longo. Sporis 5.5-11 X 3-5 ,u; cystidiis et dermatocystidiis frequenter capitatis vel subcapitatis praesentibus et multis ex eis crassitunicatis. Ad frustula lignea in Abieteto mexicano. Typus in F conservatus est.

Pileus orange ferruginous to brownish ochraceous with more pallid or whitish margin in some, not viscid, more or less sulcate when fresh, sometimes less so when dried, pubescent all over under a lens, especially so towards the margin, convex, 1-6 mm broad. Lam- ellae white, distant, narrow and almost venose when young, later well formed (about up to 8 lamellae and 0-2 lamellulae), not intervenose but sometimes forked, adnexed to adnate.

Stipe whitish to orange above, later dark chestnut to black, eventually all over so, pubescent under a good lens, insititious, equal, central, solid, later narrowly hollow, 4-7 X 0.1-0.5 mm. Context of the pileus white or whitish, inodorous.

Spores 5.5-1 1 X 3-5 A, ellipsoid to oblong, sometimes inequilateral with a strong or weak suprahilar depression, thin-walled, hyaline, smooth, inamyloid. Hymenium: basidia 23-28 X 6-9 ,, 4-spored; basidioles fusoid; cystidia on edge and sides (more numerous on

edges), 27-66 X 6.8-11 p, subulate to fuscoid, or fusoid-ventricose, to ampullaceous, with or much more rarely without a subcapitate to capitate apex 2.5-4 ,across, wall hyaline, inamyloid and thin to thick (0.2-1.5 A,), in the latter case wall thickest in the upper part of the ventricose portion and the lower part of the neck. Hyphae filamentous, not gelatinized, with clamp connections, inamyloid, in the trama of the pileus mostly with slightly thickened wall, about 3 1, broad, in the stipe parallel and brown in the basal portion of the stipe, thick-walled; hymenophoral trama regular, with the hyphae hyaline and strongly interwoven in the portion near the pileus-trama, less interwoven near edge, with thin to moderately thick (0.2-0.5 A) wall. Cortical layers: epicutis of the pileus hymeniform, its elements 10-22 X 8-11 , hyaline, vesiculose to ventricose-subvesiculose or clavate-subvesiculose, wall 0.2-2 Mlthick, smooth; among these some mucronate or even slightly subcapitate elements, with hyaline to ochraceous-brownish wall; dermatocystidia 37-57 X 7-1 1 , much like those of the hymenium, but more frequently thick-walled, more or less ampullaceous and often with subcapitate or capitate apex, wall 1.5-3.3 Mthick, inamyloid, often with knob-like apex; covering of the stipe of the same elements as the epicutis but not hymeniform, the subvesiculose elements hyaline to ochraceous or melleous, e g 12 X 6 M,, the dermatocystidia numerous (causing the pubescence of the stipe), here reaching 20-70 X 7.5-10 l

On fallen twigs and other woody matter on the ground in Abietetum (Abies religiosa) in the fir-belt of the Central Mexican mountains.

MATERIAL STUDIED. MEXICO. Road from San Pedro de Nexapa to the Paso de Col6n, 16 Jul 1969, Singer MA-10 (F), type; Ladera W of Cerro Ocopozco near Topilejo, 3200 m alt, 27 Jul 1969, Singer M 8225 (F).

This species differs from Gloiocephala caucasica (Singer) Singer comb nov




(Marasmius caucasicus Singer Rev. Mycol 2: 230, 1937) which would key out with it although the stipe is much shorter in the Mexican species, by growing on conifers rather than fagaceous branches, the different measurements of the stipe and less grayish colors on the marginal surface of the pileus. An illustration of the Caucasian species given (Fig 104) for comparison.

13. Gloiocephala ciliata (Hennings) Singer, Sydowia 00: 00, 1973. Fig 106. Discocyphella ciliata Hennings, fversigt af Kongligar Vetenskaps- Akademiens Forhandlingar

56: 318. 1900. TYPE. Dusen P320, from Chile. Pileus probably ochraceous when fresh, drying pale brownish, glabrous, smooth con-

vex, 0.7-1.5 mm. Lamellae either none at first or venose, eventually generally (1)3-4(5) through-lamellae present which are entire and simple, with entire edge, moderately broad. Stipe central or subcentral, white, below ochraceous brown, eventually entirely brown, subequal, solid, insititious, 1.5-2 X 0.05-0.1 mm.

Spores 8.5-12.5 X 3-4.3 p, most frequently 10-11 X 3.7 u, fusoid, smooth, hyaline, thin-walled, inamyloid. Hymenium: basidia 25-29 X 6-8 u, 4-spored but some 2-spored ones intermixed; basidioles narrowly clavate; cystidia 29-50 X 4.8-11 , ventricose below and capitate above, with the constriction above 1.84.8 ,wide, the capitulum (3.5-)4.8-8. 9(-9) jin diameter, hyaline, thin-walled, inamyloid; another type of cystidia more scattered, about 30 X 4.5-5 p, ventricose-fusoid or fusoid-subcylindric, hyaline, without constriction or capitulum but at the apex mostly attenuate to about 1.8 p. Hyphae of the regular hymenophoral trama and the trama of the pileus filamentous, hyaline, about 1-3.5 ,broad, not gelatinized, thin-walled, inamyloid; those of the stipe parallel with a wall 1-2.5 jthick, inamyloid to vaguely pseudoamyloid, with clamp connections, broader than hyphae of pileus at least in the core of the stipe. Cortical layers: epicutis of the pileus hymeniform or subhymeniform, consisting of broad subisodiametrical or clavate cells with more or less long pedicel, hyaline, with thin to thick wall, 18-30 X 8.5-20.5 4,; among these dermatocystidia of the two types encountered in the lamellae and these numerous and conspicuous at the edge of the margin of the pileus; covering of the stipe consisting of dermatocystidia of both types, the capitate type more numerous, all thin-walled, sometimes short and rudimentary, subcylindric to conical, hyaline; some of the superficial thick-walled hyphae showing as appressed thick-walled hyphal ends among the dermatocystidia, these with broad obtuse tip.

On rotten twigs (of coniferous trees?). MATERIAL STUDIED. CHILE. Valparaiso, Dusen P 320 (B), type. Unfortunately, the host is unknown. In the following subsection it comes close to

G. alvaradoi and G. confusa, differing from the first in spore measurements, from the second in habitat and color. Its position in subsection Religiosae is somewhat dubious. Aside from that, the species, known only from the type collection from Central Chile, may not be a truly neotropical element. The Asiatic species known to belong in this group, Gloiocephala caucasica (Singer) Singer, has considerably larger carpophores and more numerous lamellae.

Gloiocephala sect Gloiocephala subsect Confusae Singer, subsect nov Pileo plerumque albido vel albo; stipite centrali, excentrico, laterali; sporis satis ex-

iguis (usque ad 12 plongis);dermatocystidiis generatim conspicuis, capitatis; pills quam 200 ulongioribus absentibus. Ad Dicotyledones.

Pileus mostly whitish or white; stipe central or eccentric or lateral.



Gloiocephala 297

On Dicotyledones. Spores rather small, up to 12 llong. Dermatocystidia generally conspicuous and capitate, but hairs longer than 200 u absent.

TYPE SPECIES. Gloiocephala confusa Sing.

Key to the Species of Subsection Confusae

1. Clamp connections present. 2. On fallen dead leaves; cystidia thin- to thick-walled, capitate.

3. Hymenophore often slowly developing but in fully mature specimens usually present, lamellar (5-6 lamellae). 14. G. conJilsa.

3. Hymenium none. hymenial surface smooth, sometimes a few vein-like ribs formed in the immediate neighborhood of the stipe. 4. Spores more than 5 p broad. 15. G. mycenoides. 4. Spores less than 5 p broad. 16. G. epiphylla.

2. On wood; cystidia in hymenium none or few and then thick-walled, metuloid, derma- tocxystidia also metuloid; stipe lateral. 17. G. quitensis.

1. Clamp connections absent; lignicolous. 18. G. alvaradoi.

14. Gloiocephala confusa Singer, Sydowia 14: 278. 1960. Fig 107. TYPE. Singer B 426, from Brazil. Pileus white, smooth, eventually slightly sulculate above the lamellae and becoming

slightly buff when drying out, under a lens finely pubescent, obtuse, convex, 1-2.5 mm broad. Lamellae developing slowly, at first none and some mature basidia and spores found on the smooth hymenial surface which is white, later developing five veins or rather narrow lamellae, lamellae quite distinct and not vein-like when fully mature, usually five or six of these, distant to extremely distant, not intervenose or inserted, adnate, not collariate. Stipe white, chestnut to dark chestnut below, eventually often black with umber apex, finely pubescent from the dermatocystidia under a lens, insititious, filiform-equal, 839 X 0.1-0.3 P. Context thin, membranous, white, unchanging in the pileus, inodorous.

Spores 8-12 X 2.5-3.3 u fusoid to almost acicular, on thinner side mostly applanate when seen in profile, hyaline, but tending to become brown where in contact with the incrustations of the cystidia, smooth, inamyloid. Hymenium: basidia 18-24 X 4.8-6 , 4- spored: basidioles fusoid; cystidia moderately numerous on both edges and sides of lamellae but more numerous on edges, 30-68 X 6-15 u, ventricose with capitate apex, more rarely cylindrical and rarely non-capitate, with the capitulum mostly as broad or broader than high, 5.5-10 wide, with thin to slightly thickened wall (0.3-0.6 s thick), hyaline but with a conspicuous succineous brown to chestnut brown resinous incrustation, inamyloid. Hyphae of the pileus-trama loosely arranged and perhaps weakly gelatinized in places, not parallel, hyaline, filamentous with clamp connections; those of the stipe abruptly different, parallel, deep chestnut in KOH in the lower portion of the stipe, vaguely pseudoamyloid, with firm to slightly thickened wall. Cortical layers: epicutis of the pileus hymeniform, made up by vesiculose elements interrupted by some dermatocystidia like those of the hymenium; occasionally a few metuloid elements present, e g 28 X 5.5 u, fusoid to subulate, acute, thick-walled; occasionally the dermatocystidia showing some apical prongs e g 4.4 X 1.8 u, but both metuloids and prongs not constant; covering of the stipe with numerous dermatocystidia which are 16-39 X 2.8-7 u, rarely larger, shaped and incrusted like the hymenial cystidia but in the colored portion of the stipe more often with brown than with hyaline main wall; among these dermatocystidia, some filamentous inconspicuous ascendant terminal cells of surface-hyphae.

On dead dicotyledonous leaves in tropical coastal forest and subtropical marginal forest. Along the South Atlantic coast.




MATERIAL STUDIED. BRAZIL. Rio de Janeiro, Angra dos Reis, 1 Jan 1952, Singer B 426 (F), type. ARGENTINA. Buenos Aires, Punta Lara. I May 1966, Singer S 738 (BAFC).

15. Gloiocephala mycenoides Singer, Beil. Nova Hedwigia 29: 148. 1969. TYPE. Singer M 7161, from Chile. Fig 108. Pileus white, pilose at least under a lens, especially ciliate at the margin, smooth,

convex or with slightly depressed center, 1-3 mm broad. Lamellae none (smooth hyaline, hymenial surface), or with very few distant radial veins visible under a lens, white. Stipe white, under a lens lightly pilose near the apex, more densely pubescent in the central region and below it, filiform and equal, long, the pubescence often so heavy at the base that the latter appears mycelioid, 6-22 X 0.1 mm. Context entirely white, very thin, inodorous.

Spores 10-11 X 5.5-6.5 , ellipsoid, smooth, hyaline, inamyloid. Hymenium: basidia 26-30 X 8.5-9.5 , 4-spored; basidioles more or less fusiform; cystidia 36-76 X 7.5-16 , ventricose below, above an apical constricition (2.5-5.5 p across) mostly subcapitate or capitate (3.7 diameter), hyaline, thin-walled, not or not distinctly incrusted, inamyloid. Hy- phae filamentous, not or vaguely gelatinized, inamyloid, filamentous, with clamp connections. Cortical layers: epicutis of the pileus hymeniform consisting of vesiculose elements e g 22 X 15 A, interspersed with dermatocystidia, both elements often pedicellate, the dermatocystidia much like the hymenial cystidia, but more variable, sometimes broader with short neck, sometimes very long with rather firm but not thick walls. Covering of the stipe much like those of the epicutis.

On dead fallen leaves of Aextoxicum in coastal fog forest. MATERIAL STUDIED. CHILE; Coquimbo: Fray Jorge, 22 Jul 1967, Singer M 7161 (SGO), type.

16. Gloiocephala epiphylla Massee, Grevillea 21: 34. 1892. TYPE. Material sent from Jamaica (not located at K). Pileus white or whitish, but tending to turn brownish at least in places when dried,

or brown dotted (from the resinous exudations of the dermatocystidia), neither glutinous nor mealy but often sticky from the exudations, convex then concave, smooth, tending to become nutant while drying, sometimes becoming papillate when dried, 0.3-3.5 mm broad. Lamellae none; hymenial surface smooth or with some low veins around the attachment of the stipe, white. Stipe white, brownish to black or deep brown below, pruinate from the dermatocystidia, insititious, almost equal or equal, central, eccentric, or lateral, 1-10 X 0.1- 0.3 mm. Context very thin, inodorous.

Spores 8-11 X (3-)3.5-4(-4.5) ,, applanate near base or on entire inner side when seen in profile, with or without a slight suprahilar depression, oblong to subfusoid in frontal view, hyaline, but those sticking to the resinous incrusting masses becoming brown, smooth, inamyloid. Hymenium: basidia (15-)19-27 X 4.5-7.5 ̂ , 4-spored, but sometimes many of them 2(-3)-spored; basidioles fusoid; cystidioles similar but larger and sterile, present only in over-aged specimens; cystidia moderately numerous to very scarce, sometimes absent, 35-40 X 8.5-10.5 p, hyaline, non-incrusted, ampullaceous or ventricose-mucronate; oleocystidia few or none in the hymenium, if present ventricose to subcylindric (mostly ventricose) in the lower portion, capitate, 36-110 X 9.5-16 p, (capitulum 7-22 ,across, constriction below it 3.5-10 in diameter), with the wall proper hyaline and thin, more rarely up to 0.4(1.5) p thick, deeper rooting than the basidia, with a caducous, sheet-like resinous incrustation which is brownish to spadiceous (violet brown in NH40H) and dissolves partly in most media, especially chloral hydrate, inamyloid. Hyphae of the pileus-trama hyaline, not gelatinized, thin-filamentous with thin inamyloid wall, often wavy and interwoven towards the epicutis and gradually more densely septate and interlaced towards the



Gloiocephala 299

hymenium, with many true clamp connections, but many septa "secondary", without clamps; hyphae of the stipe parallel, inamyloid or vaguely pseudoamyloid, rather dense, 2-17 p broad (i e many inflated), brownish stramineous in the pigmentes portion. Cortical layers: epicutis of pileus hymeniform made up by broadly clavate to ventricose or subisodiametric, often short pedicellate cells, 10-40 X 9-29 u, these with thin or rarely slightly thickened wall, hyaline or becoming brownish walled and non-incrusted rarely incrusted like the dermatocystidia (if the latter are scarce), with thin to slightly thickened (0.4 ,u), inamyloid wall; among them a variable number (always rather numerous and conspicuous at the margin of the pileus) of dermatocystidia(oleocystidia 30-160 X 5-30,, mostly 57- 135 X 14-38 u, deeper rooting than the other elements of the epicutis, with hyaline to stramineous-hyaline, thin or firm, up to 0.2-0.5(-1) p thick wall, mostly with continuous, succineous-brownish to chestnut, resinous (often violet-brown in NH40H) incrustation which breaks up into irregular sheets, cylindric or ventricose, the capitate apex (5.5-30 , diameter) sometimes caducous because of deliquescence of the wall at the constriction underneath (3.5-1 5 across), the base often swollen (12-20 p diameter); covering of the stipe consisting of numerous dermatocystidia like those of the pileus, but also smaller ones without capitulum and incrustation (18-24 X 3.5-9 p), or even vesiculose ones present, some also with a resinous brown incrustation and likewise mostly thin- to firm-walled. The incrustation is soluble in chloral hydrate and forms brown drops in this solvent.

On fallen dicotyledonous leaves in the tropical and subtropical forest and along forest roads, and in gallery forest from dicotyledonous trees and shrubs (known hosts: Croton, Miconia, Cecropia). From Jamaica to Argentina.

MATERIAL STUDIED. VENEZUELA. Dto. Federal between Colonia Tovar and El Tigre, 29 Jul 1972, Dumont et al VE 6580 (NY); Bolivar: Along Rio Sacaica, 7 Aug 1972, Dumont et al VE 7287 (NY), ECUADOR. Tungurahua, Roi Topo (towards San Carlos), 30 Apr 1973, Singer B 7212 (F), B 7192 (F); Napo, Lago Agrio, 8 May 1973, Singer B 7308 (F). ARGENTINA. Tucuman: Chic- ligasta, Rio Cochuna, 11 Feb, 1952, Singer T 1824 (LIL), T 1824a (LIL); Tafi, Rio de los Sosas, 26 Feb 1952, Singer T 1872 (LIL).

I was unable to locate the type, but its description and illustration leaves no doubt but that Massee's fungus refers to the species described above. The supposedly 1-spored basidia are either a (common) erronerous observation or refer to the apiculus of the basidioles.


17. Gloiocephala quitensis Singer, sp nov TYPE. Sinlger B 7113, from Ecuador. Pileo albo, 1.5-2 mm diametro vel elongato usque ad 5 mm; lamellis nullis; stipite

albo, sub lente piloso, insititio, laterali. Sporis 8-10.2 X 4.5-5.5 p; hyphis fibulatis, in

epicute metuloideis dermatocystidiis62-169 1 longis praesentibus. Ad ramos dicotyledoneos delapsos et arboribus arbustisque affixos in Aequatoria. Typus in F conservatus est.

Pileus white, macroscopically seemingly glabrous, under a lens pilose from erect hairs projecting 1/10 mm, not viscid, smooth or subsmooth, circular-discoid or tongue- shaped with oval to fusiform circumference, 1.5-2 mm diameter or elongated to 5 mm. Lamellae none; hymenial surface white, glabrous, smooth, flat or almost so. Stipe white, pilose under a lens, equal, horizontal or at an angle with the pileus strictly lateral, insititious, 0.3-2 X 0.4-0.5 mm. Context white, very thin, inodorous.

Spores 8-10.2 X 4.5-5.5 p, ellipsoid, smooth, hyaline, inamyloid. Hymenium: basidia 30-34 X 6-7 p, 1-4-spored; basidioles fusoid; cystidia none or few, and then either of the metuloid kind as on the sterile surfaces or clavate-ventricose and broadly obtusely rounded, like the epicuticula non-cystidial elements, mostly absent. Hyphae hyaline,




thin-walled, inamyloid, with clamp connections. Cortical layers: epicutis of the pileus with long-hair-like metuloid dermatocystidia interspersed among the hymeniformly ar-

ranged elements of the epicutis, these 24-45 X 6.5-11 1, entire, clavate to clavate-ventricose, with broadly rounded apex, thin-walled, inamyloid, hyaline; metuloids 62-169 X 8-16 A, broadest in lower half, mostly attenuated to an obtuse tip, thick-walled (wall 1-4.5 sp thick), apex usually thin-walled, inamyloid or vaguely pseudoamyloid at times, with amorphous hyaline incrustation in many, deep rooting. On the surface of the stipe usually also numerous dermatocystidia of the same type.

On dicotyledonous branches fallen to the ground or on dead standing branches. Knownhost: Clibadium sp (Compositae).

MATERIAL STUDIED. ECUADOR. Pichincha: Quito, Quebrada de Miraflores, at 250 m, 24 Apr 1973, Singer B 7113 (F), type, B 7111 (F).

18. Gloiocephala alvaradoi Singer, sp nov Fig 109. TYPE. Singer T 3909, from Argentina. Pileo albo, 4-5 mm lato; lamellis albis, distantibus; stipite albo, usque ad 10 X 0.2

mm; mycelio basali albo. Sporis 7-8.5 X 3-3.7 ,; cystidiis capitatis; dermatocystidiis simillimis; hyphis defibulatis. Ad corticem Alni in Argentina. Typus in F conservatus est.

Pileus white, glabrous but under a lens pubescent, convex, soon flattened, 4-5 mm broad. Lamellae white, distant, sometimes intervenose when old, rather narrow to mod-

erately broad, at first very narrow, adnate, eventually decurrent. Stipe white, finely pubescent as the pileus, rather subequal, not insititious, up to 10 mm long and 0.2 mm broad. Basal mycelium finely fibrillose, white. Context white, thin, inodorous.

Spores 7-9.5 X 3-4 1, oblong-fusoid, smooth, hyaline, inamyloid. Hymenium: basidia 20-35 X 5.5-6 p, 4-spored, with some 2-spored ones intermixed, basidioles narrowly clavate or cylindric, few fusoid, without basal clamp; cystidia 32-60 X 4.5-11 u, ventricose

below, capitate, subcapitate, or rarely claviculate above, with a constriction 1.5-6 ,wide, apical swelling bilobed, 4-9.5 p across, wall moderately to distinctly thickened but inner outline of wall layer not clearly defined in KOH mounts and cystidia there appearing opaque. Hyphae hyaline, filamentous, not gelatinized, clamp connections, inamyloid. Cortical layers: epicutis of the pileus hymeniform, consisting of basidiomorphous elements 10-23 X (3-)5.5-8 M, hyaline, often roughened in the apical region, thin- to moder-

ately thick-walled, cylindric to clavate or ventricose. Dermatocystidia among these elements numerous, like hymenial cystidia. Surface of the stipe with dermatocystidia similar to the hymenial cystidia.

On living cortex and wood of standing Alius jorullensis, often among mosses in

company of Mycena yalae Sing., also on living trunk of Polylepis australis. MATERIAL STUDIED. ARGENTINA. Jujuy: Lagunas de Yala, about 2000 m alt, 9 Feb 1965,

Singer T 3909 (F), type, Singer T 3945 (F).

Gloiocephala sect Spathulariae Singer, sect nov

Carpophoris spathulatis vel cochleatis; hymenophoro plerumque venoso; sporis minutis, usque ad 6 s,longis.

TYPE SPECIES. G. spathularia Singer.

Key to the Species of Section Spathulariae

1. Pileus white. 19. G. spathularia. 1. Pileus (brownish-) yellow. 20. G. lutea.



Gloiocephala 301

19. Gloiocephala spathularia Singer, Sydowia 14: 279. 1960. Laschia merulioides Heim, Rev. Mycol. 10: 49. 1946 ex Heim, Rev. Mycol 31: 154. 1966.

TYPE. Singer T 2308A, from Argentina. Pileus white, smooth, macroscopically glabrous, pruinate under a good lens, some-

what transparent, convex to flat, sometimes with a small papilla, circular to oval in circumference, 2-5(-11) mm broad and often tongue-shaped elongated to 4-11 mm. Lam- ellae at first none (smooth hymenial surface), but soon anastomosing veins or narrow lamellae formed (not strongly or not at all radially arranged) which form a anastomos-

ing, sometimes merulioid hymenophore. white when fresh, slightly pruinose when seen under a good lens, attingent or subdecurrent at the stipe. Stipe white, laterally or sub-

laterally attached but often continued as a rib-like continuation on the outer surface of the pileus, often curved or forming an angle at the lateral point of attachment so that the pileus remains in horizontal position with the hymenium looking downwards, more

rarely eccentrically attached, pubescent under a lens, smooth, mostly equal, insititious or with a minute white fibrillose basal mycelium (both observed in a single population) 3-1 2 X 0.3-1 mm. Context white, thin, unchanging, somewhat tough-flesh or flesh- membranous, inodorous.

Spores 3-4.8 X 1.8-2.5 ,, cylindrical, smooth, hyaline (inamyloid, in purely 4-

spored carpophores 3-3.7 X 1.8-2.5 ,). Hymenium: basidia 16-18 X 3.5-4.8, , 1-2-3-4-

spored or all 4-spored; basidioles fusoid; cystidia 20-60 X 6-23 p, ventricose with or without an apical capitulum or knob, or clavate to ventricose-vesiculose without a capitulum, or with a slightly widened apex, many becoming covered by an applicate very thin hyaline then melleous, rarely granular resinous incrustation, pedicellate and deep rooting. Hyphae of the pileus-trama thin to firm-walled and mostly filamentous some broader

(to 8 p diameter) interwoven but loosely arranged, not or scarcely gelatinized, inamyloid, with clamp connections; those of the stipe parallel, cylindric, hyaline; subhymenium narrow, of very small interlaced hyaline elements of variable shape, often with excrescences in old material. Cortical layers: epicutis of the pileus hymeniform but becoming somewhat irregular by formation of superimposed cells, consisting of mostly erect clavate or

vesiculose, but often distorted cells e g 14-25 X 8-17 u, interrupted occasionally by derm-

atocystidia about 23-42(-59) X 8-13(-19) p but more variable in shape than the hymenial cystidia, often ampullaceous with a short, broad neck, or clavate, ventricose, cylindric or fusoid, always broadly rounded above, rarely subcapitate or with a small appendage, hyaline, also ? incrusted in the same manner as the hymenial cystidia, with thin to firm wall, inamyloid; covering of the stipe also with versiform dermatocystidia of the same

type, these 10-49 X 5-17 p.

On wood (tree trunks, fallen branches and twigs) in tropical forest. MATERIAL STUDIED. ARGENTINA. Salta: Oran, 3-4 km W of Rio Blancobridge N of

Oran, 17 Mar 1955, Singer T 2308A (LIL), type. ECUADOR. Napo, Lago Agrio, 17 May 1973, Singer B 7523, 16 May 1973, B 7494 (F).

I have not seen material of Laschia merulioides Heim from the Malagasy Republic but the description fits our species well excepting the spores (allegedly "seeming to be amyloid"); but Pegler (1973) finds them inamyloid in the type.


20. Gloiocephala lutea Singer, sp nov Fig 110. TYPE. Singer B 7333, from Ecuador. Pileo brunneolo-flavo vel flavo-luteo; lamellis nullis, dein 2-7 veniformibus; pseu-

dostipite laterali sed superficiei sterili per partem pilei affixo, parte libera 4-8 mm longa.




Sporis 3.5-5.2 X 2.3-2.8 p. Ad palmas in hylaea Aequatoriae. Typus in F conservatus est.

Pileus brownish yellow, deep yellow, glabrous, oval in outline and narrowly margined near the lateral attachment of the pseudostipe, elongated in the direction of the

stipe, 6-7 X 4.5-5.5 mm. Lamellae none, but at maturity 2-7 longitudinal low veins present which generally do not reach either the stipe-near nor the distal end of the pileus, the hymenial surface concolorous with the steril surface somewhat concave. Pseudo- stipe lateral but continued on the sterile surface of the pileus to which it is attached like the mid-vein of a leaf but not reaching the distal end of the pileus, continued downwards in the same direction as the pileus and on the same plane, concolorous, glabrous or subglabrous, insititious or subinsititious, equal, the free portion 4-8 X 0.7-0.8 mm. Context very thin, inodorous.

Spores 3.5-5.2 X 2.3-2.8 p, ellipsoid, smooth, hyaline, inamyloid. Hymenium: covering the whole concave fertile side of the pileus and the marginal portion of the other; basidia 14-26.5 X 3-5 u, 4-spored; basidioles filamentous then often fusoid; cystidia of unequal distribution, 27-41 X 8.5-14 p, ventricose and sometimes long-pedicellate, smooth and entire, somewhat opalescent or normally diaphanous, some with a slight resinous incrustation, inamyloid, thin-walled, some imbedded and not reaching the level of the sterigmata. Hyphae of the trama of the pileus inamyloid, subhylaine or hyaline, with clamp connections (but some secondary septa present), thin-walled, radially arranged but somewhat interwoven, not gelatinized, the whole pileus trama 25-60 , thick, containing aside from the hyphae also some endocystidia like the hymenial cystidia. Cortical layers: epicutis of the pileus and surface of the attached part of the stipe hymeniform, or subhymeniform, consisting of cylindrical to subisodiametric to broadly ventricose, broadly rounded elements interspersed with dermatocystidia, the former 21-27 X 8.5-12 , with firm, inamyloid wall (about 0.3-0.5 t thick), some somewhat resinous-incrusted; dermatocystidia ventricose with a constriction underneath the claviculate to capitate apex, 30-32 X 7-10 , constriction about 2.5-6.5 p across, capitulum 6-7 1, diameter, otherwise like the other cells of the epicutis, and scarcely projecting.

On palms in hylaea. MATERIAL STUDIED. ECUADOR. Napo, Sacha 4 (Sacha Norte), 10 May 1973. Singer B

7333 (F), type. The carpophores are unique by their spoon like shape and their color.

Gloiocephala sect Sessiles Singer, sect nov Stipite pseudostipiteque absentibus atque pileo sessili: sporis maioribus quam 12.5

p et pro ratione latiusculis; cystidiis metuloideis amyloideis aut cellulis diverticulatis in

epicute praesentibus; oleocystidiis praesentibus; mycelio lignicola. Stipe and pseudostipe absent; pileus sessile; spores > 12.5 p, relatively broad; amy-

loid metuloids and/or diverticulate elements in epicutis present; oleocystidia present. Lignicolous.

TYPE SPECIES. G. sessilis Singer. The discovery of the sessile species of Gloioceplhala on one hand and the "pileate"

Physalacria subpeltata on the other, both having the characteristic oleocystidia, shows quite clearly that these genera, Gloiocephala and Physalacria, are related with each other and therefore marasmioid, not clavarioid fungi.



Gloiocephala 303

Key to the Species of Section Sessiles

1. Hymenium on inconspicuous veniform ridges or entirely smooth; pileus white with red punctations from resinous incrustations; metuloid cystidia on sterile surface absent; spores 6.5-8 p broad. 22. G. sessilis.

1. Hymenium on conspciuous, narrow, anastomosing, distant lamellae; pileus pale orange; metuloid dermatocystidia with amyloid wall present; spores somewhat narrower.

21. G. anastomosans.

21. Gloiocephala anastomosans Singer, sp nov Fig 111. TYPE. Singer B 7392, from Ecuador. Pileo pallide aurantiaco, 5-15 mm lato; lamellis angustis, anastomosantibus, dis-

tantibus; stipite nullo. Sporis 13-14 X 5.2-5.8 A; dermatocystidiis metuloideis amyloideis praesentibus. Ad lignum emortuum in hylaea in Aequatoria. Typus in F conservatus est.

Pileus pale orange, glabrous when seen macroscopically, with fine hair-like hyaline bodies pilose when seen under a strong lens, irregularly rugulose, convex, eventually applanate, 5-15 mm broad. Lamellae white, very distant, rather narrow, strongly intervenose-anastomosing but not densely so, concurrent. Stipe none. Context whitish, very thin.

Spores (10-)13-14.5 X (4.5-)5.2-5.8 p oblong, with suprahilar depression, smooth, hyaline, inamyloid. Hymenium: basidia 33 X 10.5 p, 4-spored; basidioles fusoid; cystidia 27-47 X 9-14 ,, ventricose, narrowly constricted underneath a capitate head, the latter 5.5-7 p wide, the constriction 3.5-6.5 p wide, the wall thin, hyaline, sometimes resinously incrusted, the head sometimes with finger-like excrescences (but rarely), inamyloid; metuloids in the hymenial surface also present although rare and smaller, sometimes completely absent, otherwise like the epicuticular ones. Hyphae hyaline, inamyloid, thin- walled, with clamp connections. Cortical layers: epicutis consisting of a hymenial layer which contains mainly a "pavement-like" hymenium of subisodiametric vesiculose elements, these thin- to firm-walled and inamyloid, often with a mucro 5-7.5 j broad, 22- 43 X 13-24 '; among these some dermatocystidia much like the hymenial cystidia but these rare and sometimes almost completely absent; besides rather numerous hair-like metuloid dermatocystidia, 63-130 X 6-9 u, with swollen (12-18 p) base, thick-walled (wall 2-3 p,thick) and amyloid (livid-violet) in most cells at least partially, the wall thinned at the obtuse tip which is often incrusted by resinous material, very few echinate hyaline thin-walled bodies sometimes present.

On dead wood in hylaea. MATERIAL STUDIED. ECUADOR. Napo, Shushufindi, 240 m alt, 13 May 1973, Singer

B 7392 (F), type. This species, obviously related to the type of the section, is characterized by its

anastomosing lamellae, the pale orange pileus, the narrower spores and, mainly, the amyloid metuloids in the epicutis. The macroscopical aspect of this fungus is much like a Marasmius of section Neosessiles, especially Marasmius cecropiae.

22. Gloiocephala sessilis Singer, sp nov Fig 112. TYPE. Singer B 7119, from Ecuador. Pileo albo, rubro-puncato, 5-6 mm lato; lamellis nullis vel veniformibus et incon-

spicuis; stipite nullo. Sporis 12.5-14.5 X 6.5-8 1; dermatocystidiis tenuitunicatis. Ad ramos et ramulos Clibadii in Aequatoria. Typus in F conservatus est.

Pileus white with red punctations from the resinous incrustations of the cystidia,




especially at the margin, smooth, glabrous but finely glandular-pilose under a strong lens, convex, obtuse, laterally attached, oval to reniform in outline, 5-6 mm broad. Lamellae none or 1-2 short veniform inconspicuous ridges present, hymenium basically smooth, white, red-spotted from the incrustations of the cystidia, concave. Stipe none, but sometimes pileus attached to a dark brown somewhat pilose rhizomorph. Context very thin, white, inodorous.

Spores 12.5-14.5 X 6.5-8 (F to 114; Q<2), ellipsoid, inner side often subapplanate, without a suprahilar depression, smooth, hyaline, inamyloid. Hymenium consisting of basidia, basidioles and oleocystidia; basidia 20-38 X 9-14.5 ,, 4-spored; basidioles

mostly + fusoid; cystidia in form of oleocystidia moderately numerous, 60-80 X 15-19 p,

narrowly ventricose, with 6-9 apical appendages e g 7 X 2 p1, making the cystidia in shape similar to Pluteus atricapillus, but exuding large amounts of resinous red (in NH40H red- brown) incrustation, with hyaline rather thin (0.6-0.8 p) wall, slightly opaque, with finely granular to filamentous internal bodies which are hyaline to pale reddish brown in ammonia. Hyphae hyaline, filamentous, not gelatinized, thin- to firm-walled, radially arranged, 1.5-4 pbroad, with clamp connections, but some secondary septa present, inamyloid. Cortical layer: epicutis of the pileus hymeniform or subhymeniform, consisting of subisodiametric to broadly ventricose, broadly rounded or rarely apically acuminate elements which are hyaline, more rarely pale succineous, 12-20 X 9-12 j, and dermatocystidia 30- 42 X 5-5.5 p, fusoid-ampullaceous with the tip often narrow and obtusate or else broadened and nodulose or elongated into a versiform hyaline diverticulate element with diverticula often irregularly broadened, 1.5-2.5 v high, at times connate and forming a crust- like covering over a thin to firm wall, the dermatocystidia rather numerous and hyaline, thin-walled; some of the epicuticular elements covered at the apex or over most of their surface with similar diverticula or with apically thickened wall (1-3 A), or apically or laterally (mostly apically) emitting setulae-like appendages, these usually single, sometimes one cell with up to three appendages, these subulate, 6-16 X 2-4 ;: in places all these elements, especially the dermatocystidia, covered by a red (brown-red in ammonia) resinous incrustation.

On branches and woody twigs of Clibadium sp in montane ravines. Ecuador. MATERIAL STUDIED. ECUADOR. Pichincha, Quito, Quebrada de Miraflores, at 2950 in

alt, 24 Apr 1973, Singer B 7119 (F), type.

8. Physalacria Peck, Bull. Torrey Club 1: 2. 1882. Eoagaricus Krieger, Maryland Acad. Sci. Bull. 3: 8. 1923.(superfluous name11). Baumanniella Hennings, Engler Bot. Jahrb. 22: 543. 1893. TYPE SPECIES. Physalacria inflata (Schweinitz) Peck. Habit of the carpophores specific, "physalacrioid," i e either the pileus deformed

to a globose or irregularly inflated (often short-cylindric, conical, or oval) head; if elongated, long-obclavate, or if piliform, fertile on the side opposite the insertion of the pseudostipe which is centrally attached and insititious; hymenial surface smooth, often in irregular fragments and concentrated in areas directed towards the ground or obliquely vert-

ically exposed; pseudostipe surface sterile, sometimes with Rameales-structure and mostly with dermato-(oleo-)cystidia (these also mostly present on the sterile parts of the pileus; hyphae of the thin, monomitic trama of the pileus filamentous, few somewhat inflated, often loosely arranged, those of the stipe parallel, all inamyloid, with clamp connections, not or slightly gelatinized; basidia normal; basidioles mostly fusoid, some ampullaceous,

1 iThe family Eoagaricaceae Krieger is therefore likewise not legitimate since it is based on a "superfluous name."



Physalacria 305

with 2-4 sterigmata: cystidia and/or oleocystidia (mostly the latter) present; cystidioles also often present; spores very small to large, ellipsoid, fusoid, oblong, acicular, or fusoid, sometimes somewhat curved, smooth, thin-walled, inamyloid, acyanophilic, hyaline in KOH, but may at times tend to become slightly brownish in overaged condition.

On dead and living leaves and wood. Development gymnocarpous, stipitocarpous.

Key to the Sections of Plysalacria

1. Pileus vertically elongated, long-obclavate. sect Hormomitaria p 305 (not represented in the neotropics).

1. Pileus not vertically eloneated. 2. Pileus subisodiametric, or short cylindric, conical or ovate, mostly hollow.

sect Physalacria, p 306. 2. Pileus hat- or disc- or shield-shaped. sect Pileolina,p 311 (not represented in the neotropics).

Physalacria sect Hormomitaria (Corner) Singer, comb nov Hormomitaria Corner, Monograph Clavaria Allied Genera, p 696, 1950. The type species, H. sulphurea Corner has been adequately described by Corner,

(1950, p 412) and should be transferred to Physalacria as P. sulphurea (Corner) Singer12 Two other species were referred to this genus by Berthier (1968) and Corner (1953) respectively. Only the latter is neotropical.

Horrnmoitaria albidula Corner, Ann. Bot. 17: 362. 1953. Pileus inverted, white then cream, solid, up to 2 mm high and 0.4-0.5 mm broad

below, 50-100 p wide at the acute apex, sterile at base (i e towards the pseudostipe). Hymenophore none. Pseudostipe subpuberulous, 0.2-0.35 X 0.1 mm.

Spores 3.5-4 X 2.5-3 ,, broadly ellipsoid, blunt, aguttate, thin-walled, the apiculus minute, (apparently hyaline, inamyloid). Hymenium: not thickening, consisting of basidia. 4-spored, acerosa when young, with basal clamp; sterigmata four, 3 ,ulong; oleocystidia absent. Hyphae 2.5-8.5 , wide, many of the cells becoming secondarily septate with septa at intervals of 6-70 p, not forming moniliform rows, thin-walled, with oily contents, not agglutinated at the surface; cystals 12 ,wide, tetrahedral, abundant in the flesh, particularly in the pseudostipe; hyphal tips 1.5-2.5 P wide. Cortical layer: pseudostipe surface with dermatocystidia to 20 j long, 2.5-6 p broad, subclavate or subventricose, with thin to slightly thickened wall, divergent, loose, smooth.

According to Corner (1953), the habit and general structure of the carpophore is the same as in Hormomitaria sulphurea, from which it differs in white fruiting bodies, smaller spores, narrower hyphae, not forming moniliform rows of cells on becoming secondarily septate, shorter caulocystidia, and absence of oleocystidia.

This latter character keeps me from transferring this species to Physalacria although Corner (1950, 1970) recognizes a species in Physalacria which is likewise devoid of oleocystidia, P. stilboides. I know neither this species nor Hormomitaria albidula. The description of the latter has been adapted from Corner (1953).

While I have no doubt but that Hormomitaria as originally described is congeneric with Phvsalacria, I am not certain about the species described above, nor am I any more convinced than Corner with regard to the position within the genus Physalacria of Phy- salacria stilboides.

Nevertheless, for completeness sake, Hormomitaria albidula is treated here. It was

12Basionym. Hormomitaria sulphurea Corner, Monograph of Clavaria and Allied Genera p 649, 1950.




collected by Corner (769) in Brazil, Rio de Janeiro, Corcovado, 28 Aug 1948, on dead wood.

ILLUSTRATION. Corner (1953), fig 3.

Physalacria sect Physalacria Characters: Those of the genus, but with neither elongated and solid nor with hat-

or shield-like pileus. See fig. 118. TYPE SPECIES. Physalacria inflata (Schweinitz) Peck.

Key to the Species of Section Physalacria

1. Spores reaching 17-22 , in length. On leaves of Clusia. 1. P. clusiae. 1. Spores smaller, not longer than 15 p.

2. On conifers (mostly on dead hanging leaves, cones, twigs, etc). 2. P. tenera. 2. On Angiospermae (if on Monocotyledones, see "6" below), mostly on dead wood of

Dicotyledones. 3. Head broader than 2.5 mm. 3. P. brasiliensis. 3. Head not broader tahn 2.5 mm (but may be longer).

4. Spores reaching up to 15 p in length or over 3.5 , in breadth. 5. Cystidia not dimorphic (no appendiculate ones present). 4. P. andina. 5. Some cystidia with finger-like appendages at apex. 5. P. conciinla.

4. Spores smaller. 6. Spores 4.5-6.5 X 2-3.5 p mostly 5-6 X 2.5-3 p; pseudostipe not insititious.

6. P. aggregata. 6. Spores up to 9, long or only up to 5(-6)p long; pseudostipe insititious.

7. Spores up to 5(-6),u long. 8. Head smooth or somewhat irregular; spores 3-5(-6) X 2-3.2 . 9. P. langloisii. 8. Head in age mostly distinctly folded-rugose; spores extremely small;

2.5-4.5 X 1.5-2.5 . 8. P. sanctaemarthae. 7. Spores up to 9g long. 7. P. orinocensis.

1. Physalacria clusiae Sydow, Ann. Mycol. 28: 35. 1930. Fig 113. TYPE. Sydow 205, from Venezuela. Pileus yellowish whitish to ochraceous-yellow, smooth, entire, 0.4-0.8 mm wide,

lentiform and broader than high or globose and slightly depressed, hollow, often with a depression around the apex of the stipe when mature. Pseudostipe concolorous, eventually tending to become bister at the base, solid, up to 0.8 X 0.1 mm.

Spores (9-)7-22 X 2(-3) ,, straight or somewhat curved with applanate or concave inner side, cylindric-subfusoid in frontal view, almost needle-shaped, hyaline, smooth, inamyloid, acyanophilous. Hymenium: basidia absent on the side opposed to the ground, 25-30 X 7-8 P (according to Sydow), 2-4-spored; basidioles fusoid, 25-34 X 6-7 ,, hyaline, some opalescent, apically mucronate or with one or two filiform appendages; cystidioles basidiomorphous, with a wall 0.5-2.5 p thick; oleocystidia (20-)47-90 X 9-11 A, slightly ventricose to strongly ventricose below, ampullaceous or with a claviculate broadened, obtuse apex, moderately numerous, often constricted, constriction to 5.5 , wide, with 0.5 p thick wall, with homogeneous or granular contents (blue in cotton blue mounts), with a granulose or more often continuous resinous incrustation all over, this pale stramineous to stramineous, both incrustation and wall inamyloid. Hyphae of the pileus interwoven or in parallel strains, hyaline in the inner layer and slightly gelatinized there, narrow-filamentous and wavy, in the outer layer with firmer inamyloid wall, with clamp connections; those of the stipe parallel with each other, inamyloid, filamentous, not gelatinized, hyaline to pale yellowish in KOH. Covering layers: stipe covered with sparse oleocystidia and numerous very small setula-like outgrowths which are short filamen-



Physalacria 307

tous, often forked, or ventricose-mucronate about 5-6.5 iphigh. Trama monomitic. On fallen leaves of Clusia (type on Clusia rosea), hypophyllous. MATERIAL STUDIED. VENEZUELA. Puerto La Cruz, Sydow 205 (LE); Aragua; Parque

Nacional Henry Pittier, 12 Jul 1971, Dumont et al VE 1919 (NY).

ILLUSTRATION. Dennis (1970) fig 4 L.

2. Physalacria tenera Sydow, Ann. Mycol. 28: 33: 1930. Fig 114. TYPE. Svdow 360, from between El Lim6n and Colonia Tovar, between Distrito

Federal and the state of Aragua, Venezuela. Berlin-type destroyed. Pileus pure white, later white or light yellowish-white, at first seemingly finely

pruinate-mealy, smooth or almost so, ovate-globose later more ovate or obtusely conical witli broadly rounded base, 0.5-1 mm high, 0.5-0.8 mm broad, distinctly differentiated from the stipe. (Pseudo)stipe concolorous, at first finely mealy, equal, only slightly widened at the apex, terete, 0.4-1(-1.4) mm long, 75-90p broad.

Spores 6-11(-12) X 2.5-3.5(-4) , obliquely ovate-ellipsoid, or oblong, gradually attenuate towards the rounded apex, with a small hilar appendage, mostly broadest below the middle, hyaline, with minutely granular plasmatic interior. Hymenium: basidia 20-25 X 5-6 y 4-spored, sterigmata 3-5 p long; (basidioles fusiform); oleocystidia scattered, 40-55 X 8-10v1, fusoid-obtuse, broadest in the middle, with light brown resinaceous incrustation at the apex. Hyphae in the pileus 1.5-3 p broad, some inflated to 8 *p, thin- walled, with sparse clamp connections: those of the pseudostipe 3-6 p broad, with firm walls. Cortical layer of the pseudostipe consisting of conical projecting escrescences.

On both sides of dead but still attached needles of Podocarpus coriaceus. MATERIAL STUDIED. None. The description is edited according to the data given by H. Sydow (1930). Phlisalacria concinna Sydow has later (Corner, 1953) been interpreted as being

identical with the species described here under no. 5, in the sense of Corner. 1 have not seen the original material of P. conlcinna which has been destroyed. However, the des-

cription of the type as given by Sydow seems to suggest identity with P. tenera. Sydow's P. concinnla grew on the same host, sometimes intermingled with specimens determined P. tenera by Sydow, at the same locality and seems to represent younger material. The

cystidioles are in my opinion merely basidioles. The smaller size of the fruiting bodies is in agreement with the hypothesis that the specimens were not fully developed. The

oleocystidia are described as sometimes capitate, a shape one would also expect to be noticeable in Physalacria tenera where it has perhaps been overlooked because of the incrustation covering the apex of these cells.

3. Physalacria brasiliensis (Rick) Corner, Monograph Clavaria Allied Genera, p 464. 1950.

Baulannleiella brasiliensis Rick, Broteria 5: 11. 1906.

TYPE. Rick, from Brazil (not studied). Pileus pale stramineous white, drying reddish brown, globose, often depressed,

hollow, 3-5 mm in diameter. Pseudostipe at first white pruinose then sordid, up to 3 X 0.5 mm.

Spores 6 X 3.5 p, ovoid, apiculate, smooth, hyaline. Other details unknown. On wound of trees, type on Anchietea salutaris (Violaceae). MATERIAL STUDIED. none. Rick indicates monosporous basidia, but this probably refers to the basidioles.

I have not studied this species. It is keyed out here and described according to Rick's




data. Corner believes this to be a Physalacria, and he is obviously right. Baker (1946) indicates a specimen of P. inflata (Schweinitz) Peck found in southern Brazil. I suspect that this might refer to P. brasiliensis and if Baker's identification is correct, P. brasiliensis might be conspecific.

4. Physalacria andina (Patouillard & Lagerheim) Patouillard, Essai taxonomique Hymenomycetes, p 50. 1900.

Phvsalacria orinocensis var anldina Patouillard & Lagerheim. Bull. Soc. Mycol. France 9: 136. 1893.

TYPE. Patouillard Herbarium from Ecuador. Pileus white to deep cream when moistened, appearing mealy, smooth or slightly

rugose, conical to acuminate, 1-2.4 mm high and 0.35-1.2 mm broad. Pseudostipe concolorous, relatively short at maturity, 0.3-0.75 mm.

Spores 12-15 X 3-4 (Litschauer in Sydow), 7-10 X 4-4.5 (Baker), ellipsoid to fusiform, smooth, hyaline. Hymenium: basidia 18-24 X 7-8 , (Litschauer in Sydow), consistently 2-spored and sterigmata about 3 u long (Baker); cystidia numerous, 30-40 X 12-15 p (Baker), according to Litschauer up to 70 , long and 12 ' broad, obtusely fusiform or ventricose-rostrate, projecting, with dark resinous contents being secreted at the apex: surface with definite fertile and sterile parts. Hyphae of the pileus thin- walled, hyaline, 2-4 p broad, in part inflated to 7-8 ,; in the hollow portion some hyphae are penetrating "radiating from the pseudoparenchymatous end of the stipe" (Baker) stipe-hyphae parallel, 3-5 p broad, somewhat thick-walled; subhymenium 15-20 P thick, poorly defined (Baker). Cortical layer: Unknown.

On dead wood. Ecuador, according to Baker also in Colombia. MATERIAL STUDIED. None (description according to Sydow and Baker).

5. Physalacria concinna Sydow, Ann. Mycol. 28: 36. 1930. sensu Corner, Ann. Bot. 17: 364. 1953.

TYPE. Sydow, from Venezuela (not studied, see note below). Pileus white, ovoid-conic, very minutely hispid when dried, up to 0.6 mm high

and 0.25-0.35 mm broad. Pseudostipe slightly brownish at base minutely pruinose when dried, 0.5 X 0.08-0.12 mm.

Spores 8-10.5 X 3.5-5 ,, ellipsoid to subcylindric, with a small apiculus, smooth. Hymenium: basidia 22-30 X 7-8.5 p, clavate, 4-spored: cystidioles 18-23 X 3.5-6 p, very abundant, set with 1-6 sterigma-like processes up to 8 p long, irregularly arranged at or below the apex; oleocystidia 35-75 X 6-13 p, ventricose-lanceolate, with a short obtuse or subcapitate neck imbedded in a globose oily-resinous mass, 10-17 , wide, thin-walled or with firm walls, usually a long base, projecting up to 30 p; transitions to cystidioles few. Hyphae of the head 2-4.5 u wide, thin-walled, (?) subgelatinous, not inflated, branched at a wide angle, hyphae of the stipe composed of compact longitudinal hyphae 2-7 p wide, clamped. Cortical layers: base of the head apparently not differentiated as a sterile zone though some of the cystidia of this part rather larger and slightly thick-walled; superficial hyphae of the stipe 2-5 , wide, the walls thin or thickened to 2 p, scarcely agglutinated, densely papillate with processes up to 5 , (rarely 15 p)

long and 1-1.5 p,broad; dermatocystidia as short oleocystidia, up to 30 Pulong, 12 p

broad, ventricose to subcylindric, colorless. On dead petioles, and leaflets of Rhus copallina and on twigs of Salix and Tilia

in Florida, U.S.A., C.G.S. (BPI). MATERIAL STUDIED. none.



Physalacria 309

I have not seen the type of this species but suspect strongly that Sydow's original collection does not refer here but to P. tenera. In this case, the Florida species described by Corner from whom I have adapted my description above would have to be renamed. This problem can be solved by studies of the type locality of P. concinna.

ILLUSTRATION. Corner (1953) fig 4.

6. Physalacria aggregata Martin & Baker, Bull Torrey Club 68: 280. 1941. Fig 115.

TYPE. Martin 2260 (IA), from Panama (not sectioned). Pileus white, drying ochraceous yellow, in form of a subglobose (slightly longer

than broad) to ovoid or oblong, frequently obtusely conical head which is often somewhat lobed or with plicatelongitudinal furrows and sometimes rugose at the stipe-near base which is truncate when mature, hollow with a thin fragile shell, 0.5-4 mm high, 0.5-2.5 mm broad. Pseudostipe white, finely pubescent but glabrescent, more or less equal, solid, often oblique in relation to the axis of the pileus, not insititious but with white basal mycelium, 1-3.5 X 0.1-0.2 mm. Context white, thin, odor none; taste mild.

Spores (4.5-)5-6(6.5) X (2-)2.5-3(-3.5) ~, ellipsoid or subcylindric to oblong or ovoid, mostly ellipsoid, but often flattened on inner side, even slightly reniform, hyaline, smooth, inamyloid. Hymenium: basidia 14-22-(32) X 6.2(-8) , 4-spored, often with a few 2-spored ones intermixed; basidioles fusoid, few narrowly clavate-subcylindric; basidia and basidioles absent or very scarce in the stipe-near zone of the head; cystioles 22-30 X 3-15 ,u, basisiomorphous but often mucronate, hyaline; oleocystidia deeper rooted than basidia and cystidioles, 27-90 X 6-12.5 A, ventricose with mucronate apex, some with subcapitate to claviculate apex, hyaline with abundant resinous stramineous to succineous-brown incrustation, some also hyaline-granular incrusted, interior at first finely granular, hyaline, later "empty," wall proper hyaline and inamyloid, thin, projecting. Hyphae of the trama of the pileus hyaline, inamyloid, with clamp connections, loosely arranged and becoming obsolete except for a thin tier below the subhymenium, with clamp connections. Hyphae of the stipe parallel, hyaline, thin-to somewhat tick (0.5-1 u) walled, 2-18 broad, inamyloid, but metachromatic (purplish red) in cresyl blue mounts, with clamp connections. Cortical layers: covering of the stipe with numerous dermatocystidia similar to the cystidia of the head, but often small and inconspicuous; stipe-near portion of the head sterile or almost so.

On dead wood (e g trunks of trees), gregarious to cespitose. Known host: Phoe- be porphyria. From Mexico and Panama south to Northwestern Argentina.

MATERIAL STUDIED. MEXICO. Vera Cruz: Estaci6n Biol6gica de las Tuxtlas, 7 km south of Montepfo, 20 Jun 1969, Singer M 8055 (F). BOLIVIA. Beni, Vaca Diez, Guayarmerin, 6 Mar 1956, Singer B 1622 (F). ARGENTINA. Tucuman, Sierra de San Javier, about 900 m alt, 17 Apr 1949 10 Jul 1949 (fruiting continuously), Singer T 458, T 458 a, (LIL), 25 Feb 1962, Singer T 3779 (BAFC).

7. Physalacria orinocensis Patouillard & Gaillard, Bull. Soc. Mycol. France 4: 41. 1888. Fig 116.

TYPE. Gaillard, from Venezuela. Pileus white, tending to cream or buffish when dried, appearing pubescent under a

lens, smooth, subglobose or campanulate to truncate-conic with broadly rounded or flattened tip, 1-2 mm broad and high. Pseudostipe concolorous, macroscopically glabrous, under a lens pubescent, equal, solid, insititious, 0.5-2 X 0.1 mm.

Spores 6-8.5 X 3.7-4.2 X 2.5-3.8 u, ellipsoid to fusoid and some (not all) laterally compressed, smooth, hyaline, inamyloid. Hymenium: basidia 18.5-30 X 6.8-8.5 u, either




2-spored or 4-spored; basidioles fusoid as in Marasmiuzs; oleocystidia numerous, much like those of Marasmius sect Epiphylli, 31-60(-80) X 7-11(-13) A, ampullaceous with a neck 4-12 X 2.5-5 g, thin-walled, rarely slightly thicker-walled and then appearing opaque obtuse, the projecting portion of the cystidia mostly incrusted by a melleous-hyaline, thin

incrusting sheet which falls off early in preparations. Hyphae filamentous, with clamp connections but some septa without clamp; those of the stipe 3-8.5 p broad, parallel with each other, all inamyloid. Cortical layer: surface of the stipe consisting of the end- cells of the cortical hyphae of the stipe but these often distorted or knotty to form some sort of Ranzeales-structure; also some dermatocystidia present, these like the hymenial cystidia, but often scarce or smaller.

On dead wood (branches, vines) from the Orinoco region south to Northern Ar- gentina. Known hosts: Celtis sp; Iresina sp.

MATERIAL STUDIED. ARGENTINA. Jujuy, Juglans-dominated forest at 1500 m alt, near Posta de Lozano, 21 Feb 1966, Singer T 5417 (F): Tucuman: Pie del Periquillo, about 1000 m alt, 30 Dec 1951, Singer T 1692 (LIL).

The first of these collections was 2-spored; the second 4-spored. The type, as reported on by Gladys E. Baker (1941) is both 2- and 4-spored. The collection T 1692 was determined P. bambusae H5hnel by Corner (1957), but the non-gramineous hosts, longer oleocystidia and larger size cause me to identify it with P. orinocensis so that P. bambusae apparently does not occur in the neotropics. The asperulate spores of P. banl- busae Hohnel and P. sasae Imai are not in agreement with the generic characters of Phlv- salacria and suggest that these species belong in some other genus, probably in or near Pistillaria. PhYsalacria bamlbusae Hohnel sensu Corner (1950) is a true Phvsalacria, closely allied to P.. orinocensis Patouillard & Gaillard and apparently sometimes misdetermined. It should be given a new name.

8. Physalacria sanctaemarthae Martin & Baker in Baker, Bull. Torrey Club 68: 286. 1941. Fig 117S.

TYPE. Martin 3485 (IA), (not sectioned). Pileus white, becoming yellow or yellowish on drying, smooth but soon becoming

rugose-furrowed with almost gyrose folds, in shape of a globose stuffed head wlich may become somewhat conic or slightly elongate-oblong and usually somewhat irregular and hollow in age, 1-2.5 mm high and 0.8-1.5 mm broad. Pseudostipe centrally attached, white, macroscopically glabrous but pubescent under a lens, equal, insititious, 1-3 X 0.1-0.2 mm, rarely shorter or longer.

Spores 2.5-4.5 X 1.5-2.5 u, ellipsoid, hyaline, smooth, inamyloid. Hymenium: basidia 11-12 X 2.5-3 ,, 4-spored (said to be 2-spored by Baker 1941); oleocystidia 33-45 X 7-9.8 p, ventricose to clavate, with firm to thick (0.4-1 p), hyaline main wall which is inamyloid and incrusted by a resinous sheet-like hyaline to yellowish incrust- tation. Hyphae with thin to thick walls, which may gelatinize slightly, non-gelatinized and parallel in the stipe, with clamp connections, inamyloid.

On woody matter in the forest. Gregarious. Known host: Palmae sp. From Co- lombia south to Bolivia, possibly also in Florida, U.S.A.

MATERIAL STUDIED. COLOMBIA: Sierra Nevada de Santa Marta, Hacienda Cincinnati, 8 Aug 1935, Martin 3485 (IA). BOLIVIA. Beni: Vaca Diez, Riberalta, 170 m alt, 31 Mar 1956, Singer B 2395 (F).

This is not a bisporous form of P. langloisii as may be suspected from the descriptions published by Baker (1941) although P. sanctaemnarthae and P. langloisii are closely related to each other and to P. aggregata. It is possible that my collections



Physalacria 311

from Miami, Fla. (F 992 and F 1259, both determined as P. langloisii (F) on coconut

shell) belong here. A collection by Rick (MO 140328) and one by Furtado both from southern

Brazil are mentioned by Corner (1970) under P. aggregata, but the carpophores are somewhat larger (head 2-3 mm wide). I have not seen this material which has the spores of P. sanctaemartae and the size almost of P. brasiliensis.

9. Physalacria langloisii Ellis & Everhart, Jour. Mycol 4: 73. 1888. Fig 117L. TYPE. Langlois 350, from Louisiana, U.S.A. (not sectioned). Pileus white to pale yellowish, in shape of a globose or subglobose head which

may become irregular, somewhat depressed or elongated, hollow, with rarely flattened stipe-near base, macroscopically glabrous, finely pubescent from oleocystidia when seen under a lens, 0.3-2 mm broad. Pseudostipe centrally attached, macroscopically glabrous, finely pubescent under a lens (from dermatocystidia), insititious, 0.3-2.5 X 0.07-0.1 mm.

Spores 3-5(-6) X 2-3.2 p, ellipsoid, oblong, or ovoid, hyaline, smooth, inamyloid. Hymenium: basidia 16-20 X 4.5-5.5 p, clavate, 4-spored. Basidioles fusoid; cystidioles 14-16 X 5.5-8.5 p, basidiomorphous but often subvesiculose; oleocystidia 28-85 X 7.5-25 p, mostly 30-55 X 10-17 1, covered by a melleous resinous incrustation, the wall proper hyaline and thin, rarely firm, inamyloid, ampullaceous or ventricose with a short neck or a constriction underneath a subcapitate apex (and the constricition 2.5-6 P wide, capitulum 3.3-7 s, in diameter), with a long pedicel reaching below the level of the base of the basidia and sometimes showing a basal inflation. Hyphae of the trama of the pileus thin-walled, loosely arranged and filamentous, some slightly inflated (near stipe), otherwise 1-3 , broad, with clamp connections, inamyloid. Hyphae of the pseudostipe parallel, many with slightly thickened wall which is inamyloid. Cortical layers: covering of the stipe consisting of dermatocystidia which are oleocystidia (see above).

On all kinds of woody material, mostly dead fallen branches of dictyledonous trees. Densely gregarious, sometimes associated with an Ozoniun-like mycelium, southern North America (Louisiana, Florida) and central Chile; probably also in Jamaica.

MATERIAL STUDIED. U.S.A. Florida: Coconut Grove, R. Thaxter 1891 (several collections), (FH): Louisiana; Langlois 350 (FH), not sectioned. CHILE. Santiago: Laguna Aculeo, 24 Sep 1967. Singer MA 7991 (SGO).

Material from Jamaica mentioned by Corner (1970) under P. aggregata (Welden 639) coincides perfectly with our description of P. langloisii according to Corner's data. I have not seen this material.

Physalacria sect Pileolina Singer, sect nov

Carpophoro pileiformi, pseudostipitato. Pileus hat- or shield-shaped with the fertile portion on the side opposite to the

side on which the pseudostipe is attached, not hollow or scarcely hollowed, consequently much like Gloiocephala from which it differs in growing on the lower side of the substratum, with the surface around the pseudostipe directed upwards and sterile (see fig. 118)

TYPE SPECIES. Physalacria subpeltata Redhead. This section might be represented in the neotropics but the few specimens thus

far found and referable to the section come from the temperate zone of North America and from Hawaii.




9. Manuripia Singer, Sydowia 14: 279. 1960. TYPE SPECIES. Manuripia bifida Singer Characters: See key p 61; (same as in the species described below).

1. Manuripia bifida Singer, Sydowia 79: 279. 1960. TYPE. Singer B 2272, from Bolivia. Pileus brownish yellow, becoming cinnamon brown, smooth, dry, with a darker

umbo, at first conic-papillate, then convex to applanate with a slight wart-like center, not umbilicate, 1-2 mm broad. Lamellae none; hymenial surface smooth or with up to three low veins, subconcolorous with the pileus surface, or paler. Stipe black, in primordium with white apex, shining, simple or branched, glabrous, equal, arising from a

rhizomorph, 1-3 X 0.1 mm; rhizomorph black, dried becoming brownish fuscous, hirsute from (under a lens) brownish hairs, thicker than the stipe, about 2 mm thick, insititious where attached to the substratum.

Spores 8-9.7 X 3.8-4.8 ,, ellipsoid, smooth, hyaline, inamyloid. Hymenium: basidia 20-21 X 5.5-5.7 1; cystidia none. Hyphae of the pileus filamentous, interwoven, hy- line, inamyloid, with clamp connections; those of the interior of the rhizomorphs hyaline, not or scarcely pseudoamyloid. Cortical layers: epicutis of the pileus hymeniform or subhymeniform, consisting of broom cells of the Rotalis-type, i e balloon shaped or vesiculose-clavate, upper portion covered by short divergent setulae, main body 12-22 X 10-18 ,u, often pedicellate, setulae about 1.5 u high, often aggregating to form crust- like surfaces over the apex of the main body, deep melleous: covering of the stipe none or inconspicuous; covering of the rhizomorphs consisting of setoid hairs, 15-95 X 8-8.5 I, fuliginous or fuliginous-chestnut, with thick (3.5 p) wall, with occasional small thorn-

like excrescencies on the sides, or angular verruculose or with several very small appendages at the apex, the larger ones cylindric to cylindric-subcapitate, the smaller ones cylindric to cone-shaped, numerous.

On dead fallen woody stick of a dicotyledonous plant in rain forest. Bolivia. MATERIAL STUDIED. BOLIVIA. Pando, Manuripi, Conquista, 26 Mar 1956, Singer B

2272 (LIL), type. ILLUSTRATION. Singer (1960), p 40, fig 9.

10. Epicnaphus Singer, Sydowia 14: 279. 1960. Characters (see key p 61); same as those of the species described below. TYPE SPECIES. Epicnaphus phalaropus Singer.

1. Epicnaphus phalaropus Singer, Sydowia 14: 279. 1960. TYPE. Singer, B 2175, from Bolivia. Pileus orange, glabrous, smooth, dry, convex or campanulate, then applanate and

tending to be papillate especially when dried, 3-4 mm broad. Lamellae none. Hymen- ial surface smooth, white. Stipe blackish, smooth, glabrous, central, equal, 5-6 X 0.1 mm; rhizomorphs none; basal mycelium white, sometimes scanty so that the stipe appears to be insititious when seen macroscopically, but always distinctly fibrillose when seen under a lens. Context extremely thin, inodorous.

Spores 6.8-7 X 3-3.8 u, ellipsoid-oblong, smooth, hyaline, inamyloid. Hymenium: basidia 20-21 X 3.8-4.2 p; cystidia none. Hyphae of the trama of the pileus filamentous, hyaline, subparallel-subinterwoven, with clamp connections, pseudoamyloid. Hyphae of the stipe parallel with each other, most thick-walled, having the aspect of Crinipellis- hairs when seen individually, especially in Melzer's reagent, hyaline to melleous in KOH,



Hymenogloea 313

strongly pseudoamyloid, weakly pseudoamyloid in the thin-walled hyphae of the connective tissue, all filamentous, not gelatinized, distinctly brown pigmented in the rind-layer. Cortical layers: epicutis of the pileus hymeniform to subhymeniform, consisting of broom cells of the Rotalis-type, with the main body 1 2-20 X 6.5-10 j, hyaline, clavate to clavate-subvesiculose or obpiriform, in the upper portion beset with setulae which are 1.5- 2 , high, divergent, pale melleous. Hypodermium not containing swollen or pigment- incrusted elements, poorly developed.

On dead dicotyledonous branch in rain forest. MATERIAL STUDIED. BOLIVIA. Pando, Manuripi, Conquista, 24 Mar 1956, Singer B

1 75 (LIL). type. ILLUSTRATION. Singer (1960), pl 40, fig 10.

11. Hymenogloea Patouillard, Essaei Taxonomique, p 146. 1900. Libellus Lloyd, Mycological Writings, Letters 45: 6. 1913. Characters see Key p 61; (same as in the species below). TYPE SPECIES. Hymenogloea riofrioi (Patouillard) Patouillard.

1. Hymenogloea papyracea (Berkeley & Curtis in Berkeley), Singer, Lilloa 22: 343. 1951.

Craterellus papyraceus Berkeley & Curtis in Berkeley, Trans. Linn. Soc. 22: 129. 1859. Libellus papyraceus (Berkeley & Curtis in Berkeley) Lloyd, Mycol. Writings. Letters 45: 6.

1913. Stereiiu riofiioi Patouillard, Bull. Soc. Mycol. France 8: 117. 1892. Hvn1ielogloea riofrioi (Patouillard) Patouillard, Essai Taxonomi que, p 146. 1900. TYPE. Fendler, from Venezuela Pileus "'cadmium orange," later "ta ming" (M&P), after drying becoming bright

red ("holly berry" M&P), glabrous or subglabrous, smooth, convex, later applanate and often helvelloid contorted, with acute, entire margin, 32-118 mm broad. Lamellae none. HymIenial surface smooth, at first basically concolorous with the pileus and likewise turning red on desiccation, but later some pallid spots forming irregularly over the surface. Stipe light yellow with "cadmium orange" (M&P) apex and bright ferruginous ("rust", 6 A 1 2 M&P) base, the apex later becoming "ta ming" (M&P), otherwise becoming deep chestnut brown ("hindu", 8 J 9 M&P), glabrous, smooth, cylindric, 50-125 X 2.2-4 mm; basal mycelium pallid, tomentose and strigose, abundant. Context of the pileus and stipe cartilaginous, flexible, thin (1 mm in the pileus), odor almost nil, taste distinctly raphanaceous.

Spores 8.3-9.7(-12 according to Dennis) X 3-4 u, cylindric to subfusoid, smooth, hyaline, inamyloid. Hymenium: basidia 19-24.6 X 6-7 ,, 4-spored; basidioles fusoid; cystidia in young specimens not differentiated from the basidioles, later (cystidioles?) extremely numerous in relation to the basidia, but not clearly differentiated from them except by their shape since they are broader and occasionally rather irregularly shaped and opaque from somewhat thickened walls, but clavate, hyaline, not projecting and only slightly or scarcely deeper rooting than the basidia, rarely with a constriction in the middle or with horn-like excrescences or bulges, 23.5-29 X 7-9.7 ,. Hyphae of the pileus-trama subparallel with each other and readially distributed, hyaline (in NH40H), rather dense, mostly filamentous, occasionally somewhat inflated, with thin to slightly thickened wall, strongly pseudoamyloid, with clamp connections. Hyphae of the stipe parallel with each other, brownish in the pigmented lower portion, not gelatinized, with with clamp connections. Covering layers: epicutis of the pileus hymeniform or subhymeniform, consisting of balloon-shaped or clavate elements, these 38-29 X 8-12.5 A, rarely constricted in the middle, with mostly very slightly thickened wall, not or scarce-




ly pseudoamyloid, smooth, hyaline in KOH and ammonia, forming a continuous and

homogeneous layer. No covering layer on the stipe. NH40H and KOH bleaching the surfaces slowly to glassy hyaline or pale yellow-

ish. On leafmold in tropical-montane forests. MATERIAL STUDIED. VENEZUELA. (Aragua?), Fendler (FH, K), type; Merida, above

Mucuy, 8 Aug 1958, Dennis 1787 (K). COLOMBIA. Cundinamarca, Salto de Tequendama, 24 Jul 1960, Singer B 3516 (LIL, COL). ECUADOR. Pichincha: Gualea, Riofrio, Jan 1892 (FH), type of S. riofrioi BOLIVIA.. La Paz, Nor-Yungas, Carmen Pampa, 17 Feb 1956, Singer B 1219 (LIL), 26 Feb 1956, B 1537 (LIL).

ILLUSTRATION: Singer (1960), pl 40, fig 11; Dennis (1970), pl 7, fig 26.

12. Rimbachia Patouillard, Bull. Soc. Mycol. France 7: 159. 1891. Characters: See key (p 61). See also note after Rimbachia paradoxa, and fig 118. TYPE SPECIES. Rimbachia paradoxa Patouillard.

Key to the Species of Rimbachia

1. Spores 10.5-11 X 3.8-6p,; on rotting fragments of mosses. 1. Rimbachia paradoxa. 1. Spores 11-14.5 2.7-4 on leaves and petioles of palms. 2. R. palmigena.

1. Rimbachia paradoxa Patouillard, Bull. Soc. Mycol. France 7: 159. 1891. TYPE. Rimbach, from Ecuador. Pileus white, buffy yellowish dried, glabrous, transparent when fresh, with straiglt

to slightly undulated margin, inversely cup-shaped, the marginal portion becoming more applanate, 4-7 mm across. Lamellae none; hymenial surface white, smooth or with some very low and obtuse radiating veins not reaching the margin, concave. Pseudostipe attached to the central portion of the sterile surface of the pileus, white, flexuous or somewhat straight, stuffed, equal, 2-4 mm long, 1 mm broad. Context thin, white.

Spores 10.5-11 X 3.8-6 , oblong to oblong-subovate, in profile with applanate inner side (according to Patouillard ovate, 6-7 X 3 iA), smooth, hyaline, thin-walled, inamyloid. Hymenium: basidia 26-28 X 6.3-6.8 p, 4-spored; cystidia none. Hyphae of the trama of the pileus filamentous, inamyloid, non-gelatinized. Epicutis of pileus not hymeniform.

On fragments of mosses in the montane region of the Andes. MATERIAL STUDIED. ECUADOR. Pichincha: Egido near Quito, Rimbach (FH), type,

Authentic material (FH). Some data obtained are from the better preserved authentic material which is id-

entical with the type; they are somewhat incomplete since the material is scanty. Thus, the position of the genus had to be judged by the more detailed data obtained from the second species described below as Rimbachia palnzigena Singer. The position of the car-

pophore in nature (i e the direction of the fertile surface with regard to the ground) is somewhat doubtful unless its development includes a later stage in which the pseudostipe bends over so that the hymenial surface is directed towards the ground or else the substratum was at least vertical if not corresponding to the lower surface of exposed earth-covered wood (roots?). Otherwise, assuming the hymenium in vivo is at maturity directed upwards, this fungus would be unique among the Agaricales as well as related forms in the Aphyllophorales.

Corner (1966) indeed describes a nutant fungus where the hymenium at maturity points downwards. However, his description refers to a collection from the Amaz-



Rimbachia 315

onian lowlands, not compared with the type, and is in my opinion not identical with the type.

The position of the genus is, consequently, still somewhat uncertain unless, as we have done in the second and third edition of "Agaricales in modern taxonomy" Singer (1962, 1974), it is derived from the characters of the second species described below. If so, it would turn out to be so close to Physalacria section Pileolina that it cannot be classified in any tricholomataceous tribe except Marasmieae.

An effort to collect topotypical material in Ecuador was unfortunately unsuccess- ful.

ILLUSTRATION. Patouillard (1900), fig 65, but see fig 118.

2. Rimbachia palmigena Singer, Darwiniana 14: 11. 1966. TYPE. Singer B 2503, from Bolivia. Pileus white, later cinerous-pallid, eventually cinereous, dried castaneous-pallid,

subglabrous to subpruinose, inverse cupuliform, eventually more repand, 3-7 mm broad. Lamellae none; hymenial surface smooth, white, dried tending to light chestnut color. Pseudostipe white, later becoming cinereous, on drying becoming brownish-sordid with fuliginous-brown base which is very slightly disc-shaped and widened, insititious, subglabrous to subpruinate, more or less oblique and the apex often very slightly recurved so that the carpophores are slightly nutant and the hymenial surface, at first directed upwards or sidewards, becomes strictly directed downwards, filiform, 10-12 X 0.4-0.9 mm; basal mycelium none, more rarely present in form of an applicate, white, membranous crust around the base of the pseudostipe. Context white, concolorous with the surface in old and dried material, very thin in the pileus; odor none.

Spores 11-14.5 X 2.7-4 1, cylindrical allantoid, thin-walled, smooth, hyaline, inamyloid. Hymenium: basidia 15-20 X 4-7 ,, clavate, hyaline, in the herbarium some brownish-grayish discolored from an intracellular necropigment, 4-spored; cystidia none, but near the margin of the pileus occasionally some sparse forked or nodulose hyphal ends entering the hymenium. Hyphae hyaline, in herbarium specimens sometimes brownish grayish from an intracellular necropigment, parallel with each other and the surfaces, or very slightly interwoven, radially arranged in the pileus-trama which is non-gelatinized, inamyloid, with filamentous hyphae with thin, rarely slightly thickened wall, monomitic, rather dense but towards the surfaces more loosely arranged; hyphae of the stipe parallel or subparallel with each other, with thin to slightly thickened inamyloid to slightly pseudoamyloid wall, all hyphae with numerous clamp connections; subhymenium dense, subramose, of minute hyphal cells. Epucutis of pileus consisting of repent, filamentous hyphae forming a cutis, some with external rather short branches or nodes as in a weak Rameales-structure, without conspicuous dermatocystidia and not hymeniform.

On palm fronds and inflorescences, petioles etc. rotting on a very humid ground of the hylaea.

MATERIAL STUDIED. BOLIVIA. Pando., Madre de Dios, Las Piedras, 5 Apr 1956, Singer B 2503 (LIL, BAFC), holotype in BAFC.

This species is in my opinion clearly congeneric with the type species of the genus Rimbachia. Both species of Rimbachia are very much like Physalacria section Pileolina which differs in the presence of characteristic oleocystidia such as found in many species of Phvsalacria and Gloiocephala. Patouillard (1900) and Corner (1966) treated Rimbachia as "cantharelloid" but both these authors include in this highly artificial group represen- tatives of Aphyllophorales as well as Agaricales.




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(extreme.. :- left.. an right) cys i di (y, adplu 2. 17. Crnieli dei . Capphr, o n cy- .... .L tidium' (c e ner) t hree--;. chioytii. 18. Crinipellis. rbd.z Apex: of . hai, elotii, soe(vr matre. 19.I. T. -rnpli purure. Chioy sii (-h) on niefo erte agno h ies (m);' s p o r e . (s p ) ; pro i l o f . p i l e u s . (,ca X: 1)~ andcr o h r eteergt) 0 rplinins

Capohoe, f iehilocsida soe.adapxofhi.2.'Fiipli egeri._ Celoytdi n spr. 2 Crnpli s

ipau na. Celcsii,spr,crohr. 2. C h e oaahsclmlie.. -Che...iocysidia (ch ) L p r s(p,. : ' cappoe (c) aro tie(s n ar fplu h)



Chae tocalathus-Marasmius 319

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FIGS 24-33. 24-26. Chaetocalathus liliputianus Cystidia (cy, pl), cheilocystidia (ch), apex of hair X 2/2. 25. Hymenium with basidia (ba) and cystidium (cy), spores, apices of hairs of pileus (sp respectively hs). 26. Cystidia and apex of hair (cy respectively hs). 27. Lachnella cecropiae. Spore, apices of hairs, cystidium. 28. Amyloflagellula pseudoarachnoides. Carpophore, spore, elements of epicutis and one pseudoamyloid appendage X 2000. 29. Marasmius yalae. Carpophore, cheilocystidium, spore, element of the epicutis. 30. Marasmius cyrillidis. Two carpophores, cheilocystidium, two spores, fragment of the epicutis, surface elements of the stipe. 31. Marasmius earlei. Carpophore, spore, cheilocystidium, element of epicutis. 32. Marasium liquidambari. Young and adult carpophore, two spores, cheilocystidium (left) and four elements of the covering of the stipe. 33. Marasmius splachnoides. Two spores and six elements of the covering of the stipe.

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of the

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X. 2~-. stucur of stp surface, spore.. 40,~:,., Maasiu crsc.-a (C and_' M:-. exustus (E. Above:j,_ smot c Ls t idi um; broom cell; caropo. Be o w s p o L e ; bro cell. . - . . 4. -Mars_u spardru.:

Two c y s t i d ia.. .. , broo m cell and spr...,



Marasmius 321

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cell, spore cheilocystidia.e 48 Maasicuts vleorotals. BroomX245 cells,aspore, cappchotre.ph49.sMarasmiu

niebuilanimy. Carpophore, upper right profile of pileus X 11/2, spore, and broom cell. 50. Marasmius hlajleicola. Carpophores, spores and broom celL 51. Marasmzius variabdliceps var derubnicans. Carpo- phore, spore, two broom cells. 5 2. Marasmius baeocephalius (B) and Ml. peckii (P). Cheilocystidium (ch), broom cell of epicutis, spores, carpophore. 53. Mlarasmius nmagnisetulosus. Carpophore and elements of epicutis, type 1 and type 2. 54. Mfarasmius polycladus. Young carpophore attached to black rhizomorph (slightly enlarged), mature pileus, two spores and two broom cells.




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FIGS 55-69. 55. Marasmirus microdendron. Carpophores rising from black rhizomorph, X 5; spore and elements of the epicutis of type 1 and type 2. 56. Marasrnits xerampelinuts. Carpophore, broom cells

;il~~~~~~~~~~~~~~~~~~~~~~~~~~~-R

and spore 57. Marasnus brnneocinctus Carpophore spore and three broom cells 58 Marasmius pallidocinctus var latisporus. Carpophore, spore, broom cell. 59. Maras,imus bam butsimiformis. Carpo- phore and three cystidioles. 60. Marasinius mazatecus. Carpophore, spore. 61. Marasmthus coiwvolu- ticeps. Dermatocystitida of stipe, carpophore, three spores, two broom cells. 62. Marasmtius neglectits. Basidioles, broom cell, two carpophores, spore, another broom cell from the epicutis (far right). 63. Marasmnius helvoloides. Carpophore, spore, broom cell. 64. Mfarasmius proletarits. Upper row: spore, cheilocystidia; two elements of the covering of the stipe. Lower row: carpophore, broom cell of the covering of the stipe, at right two broom cells of the epicutis of the pileus. 65. Marasintius bezerrae var bezerrae. Carpophore, pileus, spore. 66. Marasmtius bezerrae var griseoferrugineus. Carpophore, spore, broom cells. 67. Marasmius napoensis. Carpophore, spore, broom cell. 68. Marasmiws leomntus var aberrans. Carpophores, spore, four broom cells. 69. Maras,tius floniceps. Carpophores, broom cell, spores.

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FIS556.55 aasismirdndo.Capporsrsigfombak hzmop X ;sor n elmnt f h eiutso tp Iad ye .5. aasis eameins arohoe bom el an sor. 7.Maasiiisb3iieointu. aropor, poe ndtheebromcels 5. arsnhi

palioints a ltspnt.Capphr, pre rom el.5. aasnusbaiusnfords aro phr adtre ytiils.6.Maainu aztcs.Cropoe soe.6.Afrsnisctiou ties.Dratcsitd o tpe apohre heesoes wobom el. 2 Mrsiu elets Baidols,bro cll to apohoes soe,anthr romcel ro heepcuis(arriht. 3 Marasmus heloloids. Carophore spor, broo cell.64. Mrasnzis proltarits. Uppr row:spore cheiocytidi; to elmens ofthecoveingof te sipe.Lowr ro: crpopore brom cll o th coern o hesip,atrgh woboo elsofte piuiso tepies.6. aasnusbzere a beera. arohoe ples sor.66 laasis ezraevr riefertgiiis.Capphre poe broom cells. 67. Marasinius napoensis. Carpophore, spore, broom cell. 68. Marasmilis leollinus var~~~~~~~~~~:Ei: abrrns Crpphre, poe,for rom els.69 Aarsi u fo?ces.Capohoes boo cll spores.



Marasmius 323

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Marasmius 325

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FIGS 91-98. 91. Marasnijus perlongispermus. Above: two cheilocystidia. Center: spore, element of the epicutis. Below: carpophore. 92. Marasmius batistae. Elements of the epicutis (ep), basidiole (b), dermatocystidia (of the pileus, dce, of the stipe, dcs), carpophore. 93. Marasmius latiusculospermus. Left: two cystidia; right: spore, and an element of the epicutis. 94. Marasmius hinnuleiformis. Two carpophores according to sketch made from fresh material; cystidia (cy), dermatocystidium (dc). 95. Marasmius inyocephalus. Elements of the epicutis, carpophore, six elements of the covering of the stipe. 96. Marasmius strictipes. Spore, carpophore. 97. Gloiocephala capillata. Various elements of the epicutis (ep); cystidium of the hymenium (cy); spore (sp); carpophore (ca); long hairs of the stipe (hs); other elements of the covering of the stipe (st). 98. Glofocephala longicrinita. Hair of the stipe (st); hair of the pileus (h'p'); dermatocystidia of the pileus (-ep-); cheilocystidia (ch); smaller elements of the covering of the stipe (below).

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(mdl) de m toytii an te lmnso te e i ui rgt, larges deratoysid i u m . X.1 60,.- other. fiue X 50, capphr ~A.I. 01. G icphllogsea. Hi h;. L" o re (sp; rpphr (c) yt du ) eleet of t h e p iuti (ep. 10. G l i c p a a a l mrh. Ha- . ir,elmnso epicutis, cheilocystidium, three carpophore s X 2 ~~~.10.Gliocpaapdcro!. Hair ... st.ipe.. ( h) spore~~~~~~~~~~~~~~~~~~~~~~.. (s) carp op hor (,a) demtcytdu (d) te eeet fepcts(p.



Gloiocephala 327

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32(369).



List of Taxa 331

NUMERICAL LIST OF TAXA

1. Crinipellis 1-1. C. macrosphaerigera 1-2. C. carecomoeis 1-3. C. gracilis 1-4. C. urbica 1-5. C. subtomentosa 1-6. C. megalospora 1-7. C. perpusilla 1-8. C. bisulcata 1-9. C. pseudostipitaria

a. var pseudostipitaria b. var mesites

1-10. C. catamarcensis 1-11. C. schini 1-12. C. atrobrunnea 1-13. C. missionensis 1-14. C. herrerae 1-15. C. septotricha 1-16. C. podocarpi 1-17. C. stupparia 1-18. C. coroicae 1-19. C. tenuipilosa 1-20. C. mexicana 1-21. C. foliicola 1-22. C. phyllophila 1-23. C. dicotyledonum 1-24. C. commixta

a. var commixta b. var junia

1-25. C. excentrica 1-26. C. myrti 1-27. C. albipes 1-28. C. alcalivirens 1-29. C. trichialis 1-30. C. sapindacearum 1-31. C. tucumanensis 1-32. C. dusenii 1-33. C. rubida 1-34. C. austrorubida 1-35. C. purpurea 1-36. C. insignis 1-37. C. eggersii

a. var eggersii b. var flavipes c. var epiphyllus d. var lilaceipes

1-38. C. sublivida 1-39. C. siparunae 1-40. C. trinitatis 1-41. C. perniciosa

2. Chaetocalathus 2-1. C. columellifer 2-2. C. liliputianus 2-3. C. niduliformis 2-4. C. asperifolius 2-5. C. carnelioruber 2-6. C. aurantiacus

3. Lachnella 3-1. L. cecropiae 3-2. L. subfalcispora 3-3. L. caracassana 3-4. L. villosa

4. Amyloflagellula 4-1. A. pseudoarachnoidea

5. Flagelloscypha 5-1. F. polylepidis

6. Marasmius 6-1. M. yalae 6-2. M. polylepidis 6-3. M. cyrillidis 6-4. M. bactrosporus 6-5. M. leguminosarum 6-6. M. earlei 6-7. M. liquidambari 6-8. M. splachnoides 6-9. M. hakgalensis

6-10. M. cryptotrichus 6-11. M. chiapasensis 6-12. M. radicellicola 6-13. M. dysodes 6-14. M. atroincrustatus

a. var atroincrustatus b. var inodorosus

6-15. M. atlanticus 6-16. M. eorotula 6-17. M. defibulatus 6-18. M. tomentellus 6-19. M. llicis 6-20. M. crescentiae 6-21. M. sphaerodermus 6-22. M. magnoliae 6-23. M. echinosphaerus

6-2 3A. M. kroumirensis 6-24. M. leveilleanus 6-25. M. scotophysinus 6-26. M. euosmus 6-27. M. martini 6-28. M. carpenterianus 6-29. M. munyozii 6-30. M. sanctixaverii 6-31. M. caliensis 6-32. M. multiceps 6-33. M. cupressiformis 6-34. M. rotuloides 6-35. M. panamensis 6-36. M. pararotula 6-37. M. vergeliensis 6-38. M. apatelius 6-39. M. rotalis 6-40. M. castellanoi 6-41. M.pandoanus 6-42. M. oaxacanus 6-43. M. manuripiensis 6-44. M. psychotriophilfis



332Flora Neotropica

6-45. M. cundinamarcae 6-46. M. leucorotalis

a. var leucorotalis b. var discopallescens c. var leucozonites

6-47. M. arimanus 6-48. M. griseofuscescens 6-49. M. scototephrodes 6-50. M. leucozonitiformis 6-51. M. tanyspermus 6-52. M. carneotinctus 6-5 3. M. violeorotalis 6-54. M. carminis 6-55. M. buzae 6-56. M. populiformis 6-57. M. minimus 6-58. M. dodecaphyllus 6-59. M. platyspermus

b. f scandens 6-60. M. nebularum 6-61. M. aequatorialis 6-62. M. nothomyrciae 6-63. M. hylaeicola 6-64. M. vigintifolius 6-65. M. idroboi 6-66. M. eucladopus 6-67. M. tetrachrous 6-68. M. tereticeps 6-69. M. variabiliceps

a. var variabiliceps b. var mesites c. var tucumanensis d. var separatus e. var heptachroinus f. var nothotereticeps g. var derubricans

6-70. M. baeocephalus 6-71. M. peckii 6-72. M. chrysochaetes 6-73. M. aspilocephalus 6-74. M. pallenticeps 6-75. M. conicopapillatus 6-76. M. nigrobrunneus 6-77. M. fuligineorotula 6-78. M. magnisetulosus 6-79. M. misionensis 6-80. M. schultesii 6-81. M. conquistensis 6-82. M. boliviae 6-83. M. pallipes 6-84. M. aciculaeformis

a. var aciculaeformis b. var albus

6-85. M. polycladus 6-86. M. rubromarginatus 6-87. M. marthae 6-88. M. sanguirotalis

a. var sanguirotalis b. var paucilamellatus

6-89. M. microdendron 6-90. M. hippiochaetes 6-91. M. xerampelinus 6-92. M. hiorami

a. var hiorami b. var austroamericanus

6-93. M. aripoensis 6-94. M. puttemansii

a. puttemansii b. oligocladus

6-95. M. edwallianus 6-96. M. bromeliacearum 6-97. M. megalospermus 6-98. M. chrysocephalus 6-99. M. xerophyticus

6-100. M. guyanensis a. var guyanensis b. var erythrocephalus c. var guayarensis

6-101. M. trichorhizus 6-102. M. crinis-equi

a. var monocotyledonum 6-103. M. robertsonii 6-104. M. docotyledoneus 6-105. M. rhizomorphogeton

a. f rhizomorphogeton b. f arrhizomorphicus

6-106. M. graminum a. var graminum b. var culmisedus c. var schini d. var subalpinus

6-107. M. praecox a. f nraecox b. f rubroumbonatus

6-108. M. ruforotula 6-109. M. brunneocintus 6-110. M. pallidocinctus

a. var pallidocinctus b. var latisporus

6-111. M. rufomarginatus 6-112. M. foliicola 6-113. M. inaequalis 6-114. M. subrotula 6-115. M. amazonicus 6-116. M. tucumanus 6- 117. M. tageticolor 6-118. M. poecilus 6-119. M. phaeus 6-120. M. bambusiniformis 6-121. M. mazatecus 6-122. M. longisporus 6-123. M. onoticus 6-124. M. rubricosus 6-125. M. rhabarbarinus 6-126. M. berteroi

a. var berteroi var major

6-127. M. convoluticeps 6-128. M. matrisdei 6-129. M. neglectus 6-130. M. helvoloides 6-131. M. graminicola 6-132. M. haediniformis 6-1 33. M. proletarius 6-134. M. microhaedinus 6-135. M. trinitatis

a. var trinitatis b. var immarginatus

6-136. M. olivascenticeps 6-137. M. epelaeus



List of Taxa 333

6-138. M. digilioi 6-139. M.ater 6-140. M.ruber 6-141. M. bellus 6-142. M. beniensis 6-143. M. bezerrae

a. var bezerrae b. var chitepecensis c. var griseoferrugineus

6-144. M. napoensis 6-145. M. leoninus

a. var leoninus b. var aberrans

6-146. M. floriceps 6-147. M. cuatrecasasii 6-148. M. corrugatus

a. var corrugatus b. var aurantiacus c. var lacustris d. var portonovensis

6-149. M. araucariae 6-150. M. cladophyllus

a. var cladophyllus b. var glaberripes c. var intermedius

6-151. M. pusio a. var pusio b. var guatopoensis c. var poliobasis d. var graminivorus

6-152. M. setulosifolius 6-15 3. M. pleuracanthus 6-154. M. musicola 6-155. M. allocystis 6-156. M.oleiger 6-157. M. megistosporus 6-158. M. pallescens 6-159. M. panerythrus 6-160. M. haematocephalus

a. var haematocephalus b. var purpureomarginatus c. var pseudotageticolor d. var oenechinus e. var rubicundus f. var macrocephalus g. var transiens h. var atroviolaceus i. var leucophyllus

6-161. M. haedinus a. var haedinus b. var minor

6-162. M. splitgerberi 6-163. M. hypophaeus 6-164. M. guzmanianus 6-165. M. montagneanus 6-166. M. tenuisetulosus 6-167. M. anomalus 6-168. M. phaeocystis 6-169. M. ferrugineus

a. var ferrugineus b. var gardneri

6-170. M. bambusinus 6-171. M. nogalesii 6-172. M. wilsonii

6-173. M. dennisii 6-174. M. helvolus 6-175. M. confertus 6-176. M. yarizae 6-177. M. hyalaeae

a. var hyalaeae b. var hypsizygus

6-178. M. praeandinus 6-179. M. hinnuleus 6-180. M. pseudocorrugatus 6-181. M. spegazzinii 6-182. M. aztecus 6-183. M. chrysoblepharis 6-184. M. personatus 6-185. M. rubroflavus 6-186. M. glaucopus 6-187. M. bahamensis 6-188. M. actinopus 6-189. M. atrorubens

a. var cystidifer b. var atrorubens c. var dumontii

6-190. M. pseudoniveus a. var pseudoniveus b. var amylocystis

6-191. M. heterocheilus 6-192. M. spiculosus 6-193. M. cohaerens

a. var cohaerens b. var americanus

6-194. M. flammans 6-195. M. echinatulus 6-196. M. venezuelanus 6-197. M. jalapensis 6-198. M. isabellinus 6-199. M. polycystis 6-200. M. paulensis 6-201. M. cecropiae 6-202. M. griseoroseus 6-203. M. ustilago 6-204. M. sessiliaffinis 6-205. M. tenuissimus 6-206. M. spaniophyllus

a. var spaniophyllus b. var igazuensis

6-207. M. sejunctus 6-208. M. neosessilis

a. var neosessilis b. var montepiensis

6-209. M. pleurotelloides 6-210. M. perlongispermus 6-211. M. smaragdinus 6-212. M. batistae 6-213. M. oligocystis 6-214. M. aimara 6-215. M. ionides 6-216. M. latiusculospermus 6-217. M. murrillianus 6-218. M. hinnuleiformis 6-219. M. viegasii 6-220. M. myocephalus 6-221. M. rhyssophyllus 6-222. M. ditopotrama 6-223. M. poromycenoides




6-224. M. oreades 6-225. M. cibarius 6-226. M. albogriseus 6-227. M. niveus 6-228. M. strictipes 6-229. M. heliomyces 6-230. M. pseudocollinus 6-231. M. cohortalis

a. var cohortalis b. var hymeniicephalus c. var arenicolor d. var alachuanus

6-232. M. riparius 6-233. M. silvicola

7. Gloiocephala 7-1. G. occidentalis 7-2. G. capillata 7-3. G. longicrinita 7-4. G. longifimbriata 7-5. G. lamellosa 7-6. G. cinnamonlea 7-7. G. allomorpha 7-8. G. longisperma 7-9. G. inobasis

7-10. G. palmarum 7-11. G. podocarporum 7-12. G. religiosa 7-13. G. ciliata

7-14. G. confusa 7-15. G. mycenoides 7-16. G. epiphylla 7-17. G. quitensis 7-18. G. alvaradoi 7-19. G. spathularia 7-20. G. lutea 7-21. G. anastomosans 7-22. G. sessilis

8. Physalacria 8-1. P. clusiae 8-2. P. tenera 8-3. P. brasiliensis 8-4. P. andina 8-5. P. concinna 8-6. P. aggregata 8-7. P. orinocensis 8-8. P. santaemarth-: 8-9. P. langloisii

9. Manuripia 9-1. M. bifida

10. Epicnaphus 10-1. E. phalaropus

11. Hymenogloea 11-1. H. papyracea

12. Rimbachia 12-1. R. paradoxa 12-2. R. palmigena



Exsiccatae 335

INDEX OF E XSICCATAE

Baker, R. 1488 (6-160).

Balansa, B. 3379 (6-83); 3380 (6-231b); 4284 (6-181).

Bartholomew, E. 1935; 1735 (1-5)

Batista. Chaves 2224 (6-141)

Bertero, C. G. (M. berteroi)

Bon, H. F. 4362 (1-12); 4397 (6-76)

Boutan 341b (1-12)

Britton, E. G. 611 (6-187); 612 (6-160)

Britton, N. L. & Seaver, F. J. 1352; 1540 (1-9b)

Brown, A., Britton, N. L. & Seaver, F. J. 1350 (2-3)

Bruch, C. 67 (6-167).

Buza, J. J. & Dennis, R. W. G. 173 (6-2)

Cardenas, M. 243 (6-226)

Carpenter, J. B. 136 (6-28)

Cook, 0. F. 1538 (6-160)

Corner, E. J. H. 769 (Hormomitaria albidula)

Curtis, M. A. (see Wright) Demange

67 (6-76) Dennis, R. W. G.

42 A (6-173); J40 (6-3); 56 (6-161a); 101 A (6-106a); 101 B (6-106a); 137 (4-1); 158 (6-135); 164 (6-162); 231 (6-603); 232 (6-34); 246 A (6-140); 255 (6-lOa); 265 B (6-227); 274 (6-211); 339 (6-86); 376 (1-40); 391 (6-47); 397 (6-188); 409 (6-76); 411 (6-106a); 414 1011 (6-159); 1020 (6-112); 1022 (6-184); 1023 (6-84b); 1052 (6-196); 1968 (6-57); 1071 (1-29); 1071 A (1-21); 1103 (6-151b); 1107 (6-175); 1157 (6-112); 1165 (2-2); 1172 (6-158); 1427 (6-201); 1472 (3-2); 1775 (6-55); 1787 (11-1); 1800 (1-17); 1809 (5-1); 1876 (3-3);

de Witte 9432 (6-132)

Digilio, P. L. & Singer, R. T 488 (6-138)

Dodge, B. G. J. (no number) (1-15)

Donk, M. A. 11554 (2-1)

Dumont, K. P. et al. VE-234 (2-2); '.'6-llOa); 725 (6-207); 1919 3859 (2-2); 3S -2); 3938 (6-189b); 4194 (6-148); (6-206b); 4300 (6-117); 4676 5a); 4877 (6-189c); 5124 (6-19); 6-160); 5145 (6-38); 5173 ( ); 5980 (6-139); 5867 (6-188); (6-78); 6555 (6-31); 6580 ( 6707 (6-31); 7287 (7-16);

" 6-5); 7375 (6-170)

Dusen, P. 320 (7-13)

Duss, P. 35 (2-4); 67 C 93 (2-2); 445 (2-2); 484 (1-15); 48o ti-i5) 492 (1-9); 513 (1-15); 539 (6-215); 594 (6-186); 1765 (1-9); (6-189b); 1769 (6-221); 1877 (1-9b); 1888 (1-9)

Earle, F. S. 36 (6-231d); 85 (6-154); 184 (6-20);

269(6-6); (6-6); 501 (6-114) Eggers, H.

s n (1-371) Eiten, G. & L. T. & Felipe, G. M.

5254 (6-127) Gaillard, A.

21 (6-122); 52 (6-160b); 116 (6-145a); 128 (1-25); 136 (1-8)

Ghesquihre, J. 1355 (6-33)

Glaziou, A. F. M. 9171 (6-231a)

Goossens-Fontana, M. 109 (6-75); 605 (1-32)

Goretta, Pedro 92 (6-145a)

Grisol 31 (1-33)

Guerrero, R. T. 50 (6-148b); 61 (6-168); 68 (6-150a); 71 (6-231b); 73 (6-125); 74 (6-150a); 91 (6-181); 92 (6-149); 102 (6-148d); 114 (6-148d); 118 (6-83); 132 (6-223); 133 (6-126b)

Guzmain, G. 948 (6-228); 1582 (6-216); 1869 (6-180); 2601 (6-164); 2786 (6-228); 2800 B (6-143b); 3666 (6-7); 4524 (2-2); 4580 (6-52); 4630 (6-8); 6148 (6-182)




Holden, G. C. 35 (1-32)

Hostmann 297 (6-189b)

JeEek 4 (1-4)

Johnston, J. R. 556 (6-158); 579 (6-158); 580 (6-169b)

Kermes 732 (6-101)

Krieger, K. W. Fungi Saxonici 1457 (3-4)

Langlois, A. B. 350 (8-9)

Lazo, W. PU-124 (1-24)

Lebrun, J. 3009 (6-102)

Leprieur, F. R. 989 (1-4); 990 (6-165); 1018 (6-202); 1023 (6-165); 1038 (6-227)

Libert, M. A. 119 (6-106a)

Lopez Gonzalez, A. 119 (6-182)

Lorentz, P. G. 3035 (6-142)

Louis, J. 11010 (6-24); 14862 (1-32); 14934 (6-39); 14935 (6-75); 15216 (6-188)

Lowy, B. 365 (6-223); 367 a(6-175); 423 (6-180); 447 (6-181)

Mc Farlin 56 (6-3)

Maguire, B. & Wurdack, J. 34796 (6-118)

Mains, E. B. 3628 (6-150a); 3660 (6-126b)

Martin, G. W. 2260 ; 3469 (6-27); 3485 (8-8); 8360 (6-150a)

Martin, G.W. & Bouche, A. M. 7549 (6-160c)

Martin, G. W. & Welden, A. 7279 (2-1); 7553 (6-152); 7606 (6-204); 7673 (6-150a); 8406 (6-204); 8562 (2-1); 8580 (2-1); 8682 (1-2); 8702 (6-26); 8733 (6-35)

Martin, G. W. et al. 7552 (6-150a)

Martinez, M. 89 (6-145a)

Mexia, Ynes 4429 (6-160)

Mille, L. 14 (2-2)

Murrill, F. 19885 (6-231d)

Murrill, W. A. & Murrill, E. 84 (6-196); 142 (1-20); 163 (6-218); 632 (6-217); 1047 (1-38)

Noack, F. 34 (1-7)

Ocampo 516 (6-181)

Occhioni, P. 11 (RB 5.2.04) (6-227); 15 (RB 5.2.17) (6-1 50a)

Petch, T. 3920 (16-9); 4171 (6-198); 4238 (6-76)

Piedras, M. 108 (6-182)

Prance, G. T., Pefia, B.S. & Ramos, J. F. 3461 (6-160)

Puiggari, J. I. 1644 (1-7)

Puttemans, A. 860 (6-94a)

Ravenel, H. W. 816 (6-15a)

Rick, J. 213 (1-17); 422 (6-150a); 477 (1-39); 576 (6-148d); 642 (6-83)

SaInchez 132 (6-180)

Sandwith, N. Y. 1580 (6-190)

Schmitz-Levecq 169 (6-23)

Schultes, R. E. & Cabrera, I. 12869 (6-80)

Sehnem. A. 1917 (1-1)

Singer, R. B 53 (6-233); 54 (6-198); 88 (6-19): 88a (6-19); 403 (6-213); 426 (7-14);

436 (6-150a): 437 (6-169b); 531 (6-195): 532 (6-195); 535 (6-11Ob); 549 (6-198): 558 t1-18): 590 (6-161a); 492 (6-160g); 656 (6-55); 668 (6-138); 669 (6-141): 670 (6-160i); 676 (6-148b); 686 693 (6-126b); 714 (6-231b); 720 (1-8): 722 (6-151c); 725 (6-126a): 734 (6-160d); 735 (6-170); 736 (6-113); 737 (6-160e): 738 (6-160i); (6-165); 739 (6-160i); 740 (6-160); 743 (6-160e); 747 (1-37c) 755 (6- 114); 758 (6-19); 776 (6-156); 779 (1-37a); 782 (6-231b); 800 (6-64); B 800a (6-69a); 860 (1-35); 871 (6-64); 873 (6-64); 935 (1-35); 944 (6-170); 945 (6-160e); 958 (1-35); 980 (6-207); 1043 (6-126b); 1045 (6-160); 1048 (6-2) (6-3); 1067 (6-160); 1068 (6-175); 1079 (6-195); 1098 (6-82); 1106 (6-170); 1121 (6-188); 1126 (6-231c); 1134 (6-214); 1168 (6-145a); 1180 (6-165a); 1181 (1-37a); 1193 (6-77); 1194 (6-176); 1206 (6-192); 1210 (6-69b); 1219 (11-1); 1245 (6-64); 1270 (6-169a); 1318 (1-35); 1323 (1-20); 1369 (6-176); 1378 (2-2); 1394 (6-126b); 1395



Exsiccatae 337

B (6-69g); 1407 (6-178); 1411 (6-69f); 1414 (6-46a); 1433 (6-55); 1444 (6-25); 1477 (6-54); 1478 (6-66); 1483 (6-92b); 1484 (6-54); 1485 (6-67); 1489 (6-176); 1496 (6-201); 1499 (6-199); 1528 (6-67); 1529 (6-69e); 1530 (6-67); 1531 (6-69a); 1535 (7-4); 1537 (11-1).; 1544 (6-76); 1545 (6-76); 1547 (6-69d); 1595 (6-162); 1597 (6-150a); 1609 (6-159); 1620 (6-160h); 1622 (8-6); 1631 (6-175); 1636 (6-181); 1638 (6-140); 1648 (6-190a); 1649 (6-152); 1666 (6-204); 1690 (6-13); 1693 (6-181); 1700 (6-189a); 1704 (6-92b); 1706 (6-115); 1712 (6-140); 1718 (6-145a); 1722 (6-145a); 1722a (6-145a); 1723 (6-181); 1727 (6-150a); 1733 (6-152); 1744 (7-10); 1748 (6-189a); 1755 (6-160e); 1759 (6-150a); 1762 (6-90); 1776/III (6-9); 1779 (6-9); 1780 (6-189a); 1785 (6-175); 1785a (6-175); 1806 (6-192); 1820 (6-92b); 1821 (6-135); 1824 (6-189a); 1828 (6-189a); 1829 (6-150a); 1830 (6-145a); 1831 (6-145a); 1866 (6-152); 1868 (6-140); 1870 (6-203); 1873 (6-63); 1874 (6-63); 1875 (6-155); 1876 (6-lOOb); 1879 (6-153); 1883 (6-135b); 1884 (6-150a); 1885 6-152); 1886 (6-36); 1888 (6-145a); 1889 (6-165); 1889 A(6-190a); 1893 (6-135b); 1896 (6-145a); 1898 (6-141); 1899 (6-148); 1919 (6-143c); 1920 (6-153); 1926 (6-170); 1927 (6-97); 1928 (6-97); 1935 (6-lOa); 1936 (6-98); 1946 (6-115); 1952 (6-152); 1956 (6-64); 1963 (6-160e); 1966 (6-152); 1967 (6-160b); 1971 (6-157); 1975 (6-205); 1978 (6-105a); 1980 (6-189a); 1980 (6-6-189a); 1981 (6-118); 1982 (6-140); 1984 (6-124); 1985 (6-180); 1990 (6-145a); 1996 (6-90); 2000 (6-142);2014 (6-lOOc); 2016 (6-150a); 2027 (6-140); 2040 (6-lOOa); 2045 (6-1600; 2047 (6-208a); 2049 (6-180); 2052 (6-51); 2068 (6-222); 2069 (6-90); 2076 (6-191); 2089 (6-165); 2093 (7-10); 2094 (6-9); 2101 (6-lOOa); 2122 (6-151c); 2145 (6-41); 2146 (7-4); 2147 (6-159); 2151 (6-58); 2166 (6-135); 2172 (6-151a); 2173 (6-180); 2174 (6-148); 2175 (10-1); 2178 (6-231b); 2179 (6-227); 2189 (6-160h); 2190 (6-81); 2201 (2-6); 2214 (6-73); 2220 (6-233);

2225 (6-135); 2227 (6-43); 2228 (6-202); 2235 (6-174); 2238 (6-202); 2241 (6-160h); 2252 (6-227); 2272 (9-1); 2282 (6-174); 2295 (1-25); 2296 (7-9); 2299 (6-68); 2305 (6-lOOa); 2310 (1-37b); 2320 (1-36); 2330 (6-94b); 2349 (6-103); 2378 (1-37b); 2395 (8-8); 2413 (6-140); 2428 (6-119); 2429 (6-152); 2470 (6-196); 2478 (6-189a); 2480 (6-83); 2481 (2-1); 2499 (6-92b); 2503 (12-2); 2518 (6-105b); 2523 (6-128); 2527 (6-lOOa); 2528 (6-75); 2531 (6-105a); 2554 (6-135b); 2558 (6-171); 2565 (6-86); 2566 (6-151a); 2578 (6-134); 3047 (6-137); 3072 (6-148b); 3108 (6-162); 3109 (6-150a); 3111 (6-9); 3112 (6-170); 3112a (6-170); 3117 (6-9); 3119 (6-170); 3125 (6-86); 3250 (6-86); 3251 (6-94b); 3251 B(6-94a); 3281 (6-120); 3282 (6-86); 3342 (6-9); 3377 (1-30); 3394 (6-5); 3409 (6-143a); 3412 (1-3); 3436 (6-212); 3448 (6-89); 3489 (2-2); 3506 (2-2); 3510 (1-28); 3516 (11-1); 3560 (6-39); 3599 (6-65); 3637 (6-60); 4006 (6-64); 4029 (6-44); 4033 (7-4); 4078 (6-12); 4099 (6-200); 4103 (7-8); 4199 (6-40); 4212 (6-161a); 4224 (6-34); 4231/1 (6-34); 4240 (6-209); 4248 (6-150a); 4252 (6-34); 4269 (6-34); 6009 (6-45); 6057 (6-3); 6104 A(6-202); 6105 (6-50); 6105 A(6-50); 6119 (6-37); 6126 (6-111); 6149 (6-31); 6151 A(135); 6152 (6-134); 6154 (6-195); 6163 (6-178); 6166 (2-2); 6200 (6-152); 6206 (6-146); 6228 (6-161a); 6230 (1-2); 6234 (6-16); 6253 (6-135b); 6258 (6-147); 6262 (6-163); 6273 (6-190a); 6280 (6-161a); 6282 (7-3); 6283 (6-162); 6290 (6-16); 6295 '6-126b); 6297 (6-190b); 6298a (6-130); 6298b (6-130); 6322 (6-i61a); 6345 (6-170); 6354 (6-166); 6355 (6-163); 6380 (6-16); 6381 (6-129); 6398 (2-2); 6426 (6-37); 6444 (6-50); 6450 (6-53); 6451 (7-7); 6455 (6-196); 6462 (6-37); 6463 (2-2); 6489 (6-210); 6511 (6-146); 6512 (6-134); 6517 (6-147); 6903 (1-17); 6925 (6-65); 6927 (2-2); 6945 (6-148b); 7113 (7-17); 7119 (7-22); 7120 (6-122); 7160




B (6-150b); 7170 (1-32); 7172 (6-111); 7175 (2-2); 7192 (7-16); 7208 (6-23A); 7212 (7-16); 7224 (2-2); 7225 (6-126b); 7262 (6-148c); 7274 (7-4); 7293 (2-2); 7298 (6-63); 7301 (2-2); 7303 (6-148b); 7308 (7-16); 7311 (6-70); 7316 (1-37d); 7317 (6-106b); 7328 (6-160); 7333 (7-20); 7336 (6-233); 7340 (6-177b); 7342 (6-177a); 7343 (6-150c); 7348 (6-144); 7361 (6-48); 7363 (6-151a); 7366 (6-148c); 7377 (7-10); 7378 (6-106b); 7379 (6-150c); 7382 (6-208b); 7392 (7-21); 7394 (6-94a); 7394 (6-160b); 7400 (6-189b); 7411 (6-160b); 7412 (6-174); 7414 (6-145b); 7415 (6-208a); 7416 (6-145a); 7422 (6-148c); 7446 (6-132); 7454 (6-163); 7460 (6-69g); 7461 (6-189b); 7465 (6-189b); 7480 (6-120); 7490 (6-150a); 7494 (7-19); 7504 (1-37a); 7506 (6-108); 7512 (6-86); 7518 (6-126b); 7519 (6-144); 7523 (7-19); 7527a (1-37b); 7538 (6-61); 7541 (6-198); 7833 (6-156); 8804 (6-117); 12250; 17?25 (6-170): 15285

(7-5); 24901 (3-1); 24995 (6-92b)

C 3137 (6-205); 3165 (6-94a); 3172 (6-160a)

F 73 (6-229); 347 (6-15); 395 444a (121); 449 (6-135); 496 (6-160); 513 (6-148d); 416 (6-229); 444 (1-21); 449 (6-135); 496 (6-160); 513 (6-229); 521 (6-150a); 570 (6-145a); 596 (6-136) (6-148b); 688 (6-231d); 693 (6-145a); 741 (2-2); 992 (8-8); 1152 (6-229); 1196 (6-111); 1196a (6-111); 1219 (6-111); 1219a (6-111); 1456 (6-108); 1597 (6-187); 1906 (6-22); 1906a (6-22); 2511 (6-231d); 7348 (6-144)

M 98 (2-2); 1012 (6-206b); 1047 (6-79); 1082 (6-149); 1090 (6-160); 1123 (6-47); 1147 (6-149); 1507 (1-22); 1510 (6-228); 1519 (6-42); 1528 (1-22); 1544 (2-2); 6511, 6515 (6-62); 6517 (6-29); 7161 (7-15); 7991 (8-9); 8020 (6-108); 8027 (6-122); 8043 (6-108); 8055 (8-6); 8064 (6-160); 8066 (2-2); 8083 (1-27); 8084 (6-185); 8088 (6-146); 8104 (6-135); 8115 (6-28); 8116 (6-146); 8125

(1-14); 8137 (6-24); 8142 (6-131); 8188 (6-145b); 8201 (6-183); 8208 (6-9); 8225 (7-12); 8250 (6-10); 8251 (6-20); 8266 (6-172); 8274 (1-21); 8288 (1-21); 8299 (6-49); 8320 (6-14a); 8330 (6-42); 8331 (6-228); 8356 (6-3); 8368 (6-121); 8418/8420 (1-21) (6-7); 8444 (1-21); 8498 (6-172); 8741 (6-146); 8757 (6-208b); 8776 (6-160); 8783 (6-133); 8784 (6-187); 8785 (6-133); 8793 (2-2); 8805 (1-2); 8806 (2-2); 8811 (1-2); 8816 (6-14b); 8820 (1-21); 8827 (6-172); 8921 (6-111); 8943 (6-11); 8948 (6-122); 8956 (6-106a); 8960 (6-183); 8961 (6-91); 8962 (6-91); 8966 (6-160c); 8969 (6-11); 8971 (7-6); 8976 (6-160); 8979A (7-2); 8982 (6-109)

MA 10 (7-12) N 700 (6-193b); 4670 (6-106a) S 26 (6-224); 159 (6-106d); 201

f6-19); 229 (6-224); 590 (6-131); 632 (6-231b); 738 (7-14)

T 108 (1-12); 108a (1-12); 143 (6-107); 171 (6-160g); 184 203 (6-59); 208 (6-107b); 222 (6-160); 241 (6-160); 259 (6-59); 277 (6-233); 313 (6-106b); 458 (8-6); 458a (8-6); 516 (3-4); 540 (6-126b); 802 (6-198); 833 (1-16); 870 (6-167); 881 (6-138); 917 (1-34); 1084 (6-231b); 1114 (1-31); 1119 (1-19); 1130 (6-195); 1140 (1-23); 1182 (6-167); 1191 (6-233); 1193 (6-223); 1200 (6-231b); 1221 (6-114); 1223 (6-160c); 1233 (6-88a); 1240 (6-88a); 1244 (6-232); 1246 (6-231b); 1290 (6-69c); 1291a (1-29); 1291a (1-29); 1310 (1-29); 1329 (6-230); 1368 (6-126b); 1408 (6-59); 1428 (6-114); 1445 6-231c); 1457 (6-96); 1460 (6-116); 1477 (6-16); 1479 (6-233); 1488 (6-138); 1500 (1-24); 1527 (6-231a); 1641 (6-108); 1645 (6-107); 1647 (1-37a); 1666 (1-6); 1667 (6-230); 1669 (6-59); 1691 (1-31); 1692 (8-7); 1694 (6-195); 1707 (1-24); 1710 (1-24); 1730 (6-87); 1731 (6-160a); 1745 (1-10); 1755 (6-59); 1759 (1-24); 1782 (1-10); 1787 (6-106a); 1788 (6-106c); 1793 (1-11); 1816 (6-106a); 1822 (6-148); 1824 (7-16); 1824a (7-16); 1837 (6-198); 1854 (6-104); 1855 (1-11); 1872 (7-16); 1915 (6-226); 2018 (6-46c); 2029



Exsiccatae 339

(6-195); 2033 (6-107); 2059 (6-231c); 2070 (6-230); 2071 (1-9b); 2077 (6-76); 2103 (6-231c); 2119 (6-231a); 2126 (6-126b); 2147 (6-112); 2149 (6-76); 2150 (6-76); 2155 (6-151b); 2157 (6-160a); 2158 (1-31); 2159 (6-126a); 2161 (6-107); 2178 (6-107a); 2179 (6-88b); 2180 (6-107a); 2182 (6-107); 2183 (6-160c); 2190 (6-88b); 2191 (6-160); 2200 (6-107); 2218 (6-138); 2219 (6-126a); 2270 (6-231c); 2302 (6-83); 2308A (7-19); 2310 (6-126a); 2311 (6-112); 2791 (1-24b); 2852 (6-107); 2853 (6-88b); 2859 (1-24); 2879 (6-126b); 2886 (6-126b); 2887 (6-126b); 2902 (6-59); 2913 (6-126b); 2938 (6-160); 2948 (6-99); 2956 (6-160); 2989 (1-15); 2992 (6-160); 2995 (6-123); 2997 (6-114); 3026 (6-195); 3049 (6-59); 3073 (6-126b); 3080 (6-126a); 3156 (6-30); 3210 (6-233); 3422 (6-198); 3471 (6-88b); 3473 (6-107a); 3520 (7-4); 3521 (6-4); 3550 (6-226); 3591 (6-59b); 3668 (6-178); 3705 (6-69c); 3772 (6-46b); 3779 (8-6); 3903 (7-18); 3909 (7-18); 3945 (7-18); 4058 (6-15 Id); 4081 (6-76); 4995 (1-15); 5003 (6-106c); 5059 (6-1); 5127 (6-76); 5131 (6-1); 5202 (6-2); 5300 (7-11); 5309 (1-16); 5363 (6-224); 5417 (8-7); 5429 (6-23); 30671 (6-160)

Singer, R. & Digilio, P. L. M 23 (6-206b); 26 (6-48); 53 (6-48);

57 (6-208a); 84 (6-206b); 123 (1-13)

T 195 (6-59); 278 (6-138); 418 (6-126b); 501 (6-59

Skvortzov, B. V. & Alim 71393 (1-29)

Sparre, B. F 56 (6-62);

9587 (6-226) Splitgerber

1280 (6-162) Spruce, R.

31 (6-161a); 37 (6-117); 75 (6-33); 89 (6-150a); 97 (6-194); 112 (6-145a); 139 (6-118) (6-169a) (6-174); 140 (6-125)

Staner, P. C 325 (6-102)

Steyermark, J. A. 15605 (6-90)

Sultan Ahmad 509 (6-76)

Sydow 205 (8-1); 360 (8-2)

Thwaites, G. H. K. 810 (6-39)

Torrend, C. 56 (6-95)

Trail, J. 161 (6-125)

Ule, E. Herbarium Brasiliense 2699 (6-150a)

Vanderyst, H. 695 (6-38); 699 (6-188); 4015 (6-166)

Vargas, C. 6315 (6-227); 10050 (6-225); 13519 (6-126b)

Viegas, A. P. 7800 (6-219)

West, E. & Murrill, W.A. F 18267 (6-193b); 18277 (6-229)

White, L. W. 846 (6-179)

Wilson, P. 297 (6-172)

Wright (including Fungi Cubenses Wrightiani)

4 (6-160); 10 (6-184); 15 (1-17); 18 (6-119); 31 (6-146); 45 (1-15); 78 (2-2); 114A (6-113); 121 (6-84a); 129 (6-163); 155 (6-179); 160 (6-32); 162 (6-72); 855 (1-9); 858 (1-39)

Zamaolla 202 (6-226); 203 (6-226)

Zollinger, H. 2078 (1-29)




HOST INDEX

Abies (religiosa) 218, 295 Acer 82 Achatocarpus nigricans 25, 164 Aechmea 129, 159 Aextoxicum 93, 114, 298 Agropyrum 152 Allophylus edulis 90, 111, 253, 283 Alnusjorullensis (var. spachii) 59, 173, 300 Amaranthaceae 41 Amaranthus retroflexus 221 Anchietia salutaris 307 Andropogon 20 Arundinaria 39 Asara 114

Baccharis 150, 15 3 Bambusa 127, 129, 159 Bambuseae (bamboo) 18, 20, 35, 38, 227 Bauhinia 159 Bertholletia excelsa 4 Boehmeria 37, 111 Boehmeria caudata 39 Bowdichia virgilioides 68 Bromeliaceae 129, 144, 250, 292 Bursera 227

Caprifoliaceae 41 Carapa guianensis 147 Carpinus 277 Carya 156 Cecropia 58, 205, 212, 216, 243, 256, 299 Cecropia pamata 233 Celtis 310 Chrysophyllum 240 Chusquea 20, 29, 55, 129, 224 Clibadium 300, 304 Clibadium erosum 55 Clusia 307 Clusia rosea 307 Clusiaceae 142 Coccolobis 156 Cocos (coconut) 311 Coffea (arabica) 25, 31, 67, 253, 270, 277 Compositae 150, 300 Crescentia 82 Croton 299 Cynodon 20, 152 Cyphellopsis subglobispora 59 Cyrilla racemosifolia 67

Dryopteris 213 Dunalia breviflora 33 Durantia serratifolia 112, 25 3

Enterobolium 283 Ericaceae 116 Eryngium 81 Eugenia 137 Eugenia pungens 25 3

Fagara coco 25 Fagus 248 Ficus 75, 156, 241 Ficusdoliaria 135

Gramineae (grasses) 11, 14, 15, 16, 17, 22, 39, 89, 173, 176, 201,213,221

Hedera 90 Heliconia 198 Hibiscus 233

llex 81 Inga 101, 133, 141, 250, 277 Iresina 37, 41, 310

Juglans 310 Juniperus barbadensis 56

Lauraceae 45 Leguminosae (leguminosous trees) 69, 113,

190, 198 Ligustrum 81 Liquidambar 64 Liquidambar styraciflua 70, 267

Magnolia 80 Magnolia grandiflora 83 Malvaceae 233 Melanostomataceae 45, 213 Metahybe 37, 39 Miconia 299 Morinda citrifolia 56 Musaceae (Musa, banana) 141, 159, 206, 224 Myrtaceae 68, 142, 288 Myrtus 36, 253

Nectandra 241 Nectandra coriacea 158 Nothomyrcia 114

Oleaceae 41

Palmae 9, 198, 247, 293, 302, 310, 315 Panicum 224 Panicum maximum 243 Panicum mexicanum 19 Paritium 233 Paspalum 14 Persea 81 Phoebe porphyria 103, 104, 119, 173, 253,

309 Pinus (pine) 236 Piperaceae 30 Pithecollobium scalare 148 Platanus occidentalis 238 Podocarpus 11, 173, 250, 294 Podocarpus coriaceus 307 Podocarpus parlatorei 26, 295



Host Index341

Polylepis australis 67, 300 Polylepis sericea 61 Polypodiales 140 Populus 82, 162, 176 Prosopis torquata 33 Protium 224 Psychotria 102 Pteridophyta (Polypodiales, ferns) 151, 154,

192, 200, 213, 250, 288

Quercus (oak) 31, 32, 71, 73, 77 Quercus columbiana 99 Quercus humboldtii 67 Quercus laurifolia 77 Quercus virginiana 277

Rhus copallina 308 Richeria grandis 55 Rubiaceae 116

Sabal 156 Saccharum officinale 140 Salix 308

Salix humboldtiana 83 Sapindaceae 39 Schinus 20, 111, 192 Siparuna 5 1 Smilax 31 Solanaceae 150 Solanum 37, 39, 150, 192 Spathodea campanulata 83 Swietenia mahogani 43 Syringa 82

Taxodium 162, 176 Theobroma 51, 52, 100 Theobroma bicolor 52 Theobroma cacao 49, 52 Theobroma speciosum 52 Tilia 308 Triticum 152

Uvaria chamae 56

Violaceae 307 Vitex 51




INDEX OF GENERA, SPECIES AND VARIETIES13

Agaricus atrorubens 241 bambusinus 224 calopus 221 carecomoeis 13 ceratopus 247 chortophilus 17 cohaerens 247 columellifer 53 ferrugineus 222 fulvus 172 griseoroseus 256 haematocephalus 210 hymeniicephalus 278 liliputianus 54 oreades 273 spinuliferus 247 splachnoides 70 splitgerberi 216 tenuissimus 258 trichialis 38 trichophorus 39 urbicus 15

Amyloflagellula 9, 60 pseudoarachnoides 60

Androsaceus 62 corrugatus 191 epodius var. microsprus 220 glaucopus 240 haematocephalus 210 leoninus 186 longisporus 169 nigrobrunneus 128 rhodocephalus 210 splachnoides 70

Baumanniella 304 brasiliensis 307

Chaetocalathus 1, 3, 9, 53 africanus 53 aurantiacus 53, 57 carnelioruber 53, 57 columellifer 5, 53, 56 craterellus 53 liliputianus 53, 54, 56 niduliformis 53, 56

Chamaeceras 62 Collybia 272

albogrisea 274 bisulcata 18 carecomoeis 13 excentrica 35 fimicola 274, 275 hymeniicephala 278 nivea 276

setulosa 268 strictipes 276 stupparia 26 urbica 15

Collybiopsis 62 CrinipeUis 1, 3, 9

albipes 35, 36 alcalivirens 37 asperifolius 56 atrobrunnea 11, 21 austrorubida 37, 43 bambusae 17 bisulcata 11, 18 calosporus 54 carecomoeis 11, 13, 14, 15 catamarcensis 10, 20 commixta 12, 28, 32, 33, 34, 39, 41 coroicae 12, 18 corticalis 41 dicotyledonum 13, 32 dipterocarpi 31 dusenii 37, 41 eggersii 6, 43, 45, 49, 51 epiphyllus (v.) 47, 48, 49 excentrica 6, 34, 35 flavipes (v.) 47, 48 foliicola 12, 30, 31, 32 ghanaensis 23 gracilis 11, 14, 18 herrerae 12, 23, 28 hirticeps 29 insignis 45 iopus 43, 44 junia (v.) 32, 33, 34 lilaceiceps (v.) 47, 49 litseae (v.) 14 macrosphaerigera 10 megalospora 11, 17 mesites (v.) 12, 19 mexicana 12, 29, 32, 39 minutula 12, 34 mirabilis 37 missionensis 12, 22 molfmoana 13, 25 myrti 35, 36 nidulus 54 occidentalis (ssp) 19 orientalis (ssp) 20 patouillardii 28, 30, 41 pemiciosa 2, 45, 51, 52 perpusilla 11, 17, 18, 39 phyllophila 31 podocarpi 11, 25 pseudoarachnoides 60 pseudostipitaria 12, 19 purpurea 44

13Varieties are marked "(v)" after the epithet. Page numbers in italics refer to the place where the taxon is treated in taxonomic order.



Index 343

rubida 37, 42, 51, 52 sapindacearum 37, 39 schini 13, 20, 21 septotrichia 12, 24, 25, 29 siparunae 2, 45, 50, 52 stipitaria 9, 11, 16 stupparia 12, 26, 28, 39 subelata (v.) 14 sublivida 45, 50 subtomentosa 11, 16 tenuipilosa 12, 28 trichialis 37, 38 tucumanensis 37, 40, 41 trinitatis 43, 45, 51, 52 urbica 11, 15, 18 zonata 23

Cymatella longipes 88 Cyphella dochmiospora 59

minutissima 60 villosa 59

Cyptotrama 266

Dictyoploca rhyssophylla 271 Discocyphella ciliata 296

Eoagaricus 304 Epicnaphus 3, 61, 312

phalaropus 5, 312

Favolaschia magnifica 196 Flagelloscypha 9, 60

polylepidis 60

Gerronema 2 Gloiocephala 3, 61, 284, 302, 315

allomorpha 285, 290 alvaradoi 296, 297, 300 anastomasans 293, 303 capillata 285 caucasica 295, 296 ciliata 294, 296 cinnamomea 285, 289 confusa 296, 29 7 epiphylla 284, 291, 297, 298 inobasis 291, 292 lamellosa 285, 288 longicrinita 285, 286 longifimbriata 6, 285, 287 longisperma 291 lutea 300, 301 mycenoides 297, 298 occidentalis 285, 292 palmarum 291, 293 podocarporum 294 quitensis 297, 299 religiosa 294, 295 sessilis 302, 303 spathularia 300, 301 sulphurea 305

Gymnopus albogriseus 274 ludovicianus 278, 282 setulosus 268 strictipes 276

Heliomyces 62 berteroi 161 leveilleanus 85 plumieri 283

Hormomitaria 305 albidula 305 sulphurea 305

Hydropus 266 Hymenogloea 3, 61, 313

papyracea 313 riofrioi 313

Lachnella 9, 58 alboviolascens 58 caracassana 59 cecropiae 58 subfalcispora 58 villosa 59, 60

Laschia magnifica 197 merulioides 301

Lentinus minutulus 34 perpusillus 17

Libellus papyraceus 313

Manuripia 3, 61, 312 bifida 5, 312

Marasmiellus 2, 46, 225, 263 Marasmius 1, 8, 61, 62

aberrans (v.) 5, 164,189 aciculaeformis 123, 134 actinopus 237, 241 aequatorialis 94, 112 aimara 264, 266 alachuanus 278, 280 albogriseus 269, 274 albus (v.) 134 alliaceus 263 allocystis 5, 201, 206 amazonicus 161, 165, 225 americanus (v.) 24 7, 248 amylocystis (v.) 245. 246 androsaceus 63 anomalus 202, 220, 221 apatelius 92, 98, 99 araucariae 164, 196, 236 ardesiacus 283 arimanus 93, 104 aripoensis 124, 141 arrhizomorphicus (f) 151 asperifolius 56 aspilocephalus 122, 126 ater 162, 182 atlanticus 65, 77 atroincrustatus 6, 65, 76 atropurpureus 210 atrorubens 5, 237, 241 atroviolaceus (v) 213 aurantiacus 193 aurantiicolor (v) 154 austroamericanus 140 auriformis 283 aztecus 204, 235 bactrosporus 64, 68




baeocephalus 94, 121 bahamensis 237, 240 balansae 234 bambusae 17 bambusiniformis 161, 167 bambusinus 5, 202, 223, 224, 225 batistae 263, 265 bellus 5, 161, 163, 183 beniensis 163, 184 berteroi 162, 169, 170, 171 bezerrae 163, 185 boliviae 123, 132 brasiliensis (v) 249 brevispora (f) 216 brevisporus (v) 151, 153 bromeliacearum 124, 125, 143 bruchianus 283 brunneocinctus 6, 125, 156 bulliardii 110, 112 buzae 93, 109 caliensis 90, 91 calosporus 54 campanella 253 cantharelloides 283 capillaris 98 capillipes 82 carminis 93, 108 carneotinctus 93, 107 carpenterianus 6, 88 caryophylleus 273 castaneus 241, 243 castellanoi 92, 99 cecropiae 254, 255, 303 ceratopus 247 chiapasensis 6, 65, 73 chitepecensis (v) 185 chordalis 266 chrysoblepharis 6, 236, 23 7 chrysocephalus 5, 124, 144 chrysochaetes 122, 126 cibarius 269, 274 cladophyUus 5, 6, 163, 196, 236 clitocybiformis 283 cohaerens 237, 246, 247 cohortalis, 5, 270, 278 columellifer 53 confertus 204, 228, 229 conicopapillatus 122, 127 conquistensis 123, 132 convoluticeps 163, 173 coprophilus 220, 221 corbariensis 80, 81 corrugatus 5, 163, 191 , 194 crescentiae 80, 81, 82 crinisequi 125, 148, 149 cryptotrichus 65, 72 cuatrecasasii 163, 191 culmisedus (v) 125, 151, 152 culmiseduS 152 cundinamarcae 93, 99, 102, 104 cupressiformis 92, 95 curreyi 151 cyperinus 283 cyphellopsis 5

cyrillidis 64, 6 7, 68 cystidifer (v) 242 dasypus 16 defibulatus 65, 78 dennisii 203, 227 derubricans (v) 5, 120 dicotyledoneus 6, 125, 133, 150 digilioi 162, 181 discopallescens (v) 103 dodecaphyllus 6, 94, 110 dumontii (v) 243 dysodes 65, 75 earlei 64, 69 echinatulus 237, 249, 251 echinatus 246 echinosphaerus 80, 83, 84 edwallianus 124, 141, 143 eggersii 46 elaeocephalus 180 elongatipes 266 eorotula 65, 77 epelaeus 162, 180 epiphylloides 89 epiphyllus 87, 88 epodius 220 equicrinis 148, 150 erythrocephalus (v) 5, 147 eucladopus 94, 116 eufoliatus 90 euosmus 88 exustus 80, 81, 159 ferrugineus 5, 175, 202, 218, 222,

224, 227 flammans 237, 246, 248 flavofuscus 172 floriceps 6, 164, 190 floridanus 234, 235 foliicola 125, 158, 159 fuligineorotula 104, 122, 129 fulvovelutinus 247 fulvus 283 fusicystis 251, 252 gardneri (v) 222, 223 gardneri 222 glaberripes (v) 198, 199 glaucopus 237, 240 gordipes 159 graminicola 162, 1 76 graminivorus (v) 201 graminum 121, 148, 151 grandisetulosus 220 griseoferrugineus (v) 185, 186 griseofuscescens 9 3, 104, 105 griseoroseus 254. 257 griseoviolaceus 128, 129 guatopoensis 199, 200 guayarensis (v) 147 guyanensis 5, 6, 124, 146 guzmanianus 202, 218 haediniformis 5, 161, 177 haedinus 201, 215 haematocephalus 3, 4, 5, 6, 202, 209,

210, 214,217,218, 223 hakgalensis 65, 71, 72, 73



Index 345

heliomyces 270. 277 helvoloides 162, 1 75, 228 helvolus 5, 162, 176, 203, 214 hemibaphus 229 heptachroinus (v) 120 heterocheilus 237, 245 hinnuleiformis 268 hinnuleus 203, 232 hiorami 124, 139, 141, 145 hippiochaetes 123, 131, 138 hyalinotrichus 71 hygrometricus 79, 81 hylaeae 203, 230 hylaeicola 5, 94, 114 hymeniicephalus 278, 279 hymenofallax 205 hypophaeus 5, 202, 217, 218 hypsizygus (v) 231 idroboi 94, 115 iguazuensis (v) 259 iguazuensis 259 ilicis 80, 81, 82 immarginatus (v) 179 inaequalis 160 inodorosus (v) 6, 76 intermedius (v) 199 ionides 264, 26 7 isabellinus 252, 253, 255 jalapensis 237 jamaicensis 241 kroumirensis 84, 85 lacustris (v) 194 latisporus (v) 157 latiusculospermus 264, 26 7 leguminosarum 64, 69 leoninus 5, 6, 160, 163, 186, 188, 189,

190 leucophyllus (v) 202, 214 leucorotalis 93, 99, 102, 106, 114 leucozonites (v) 94, 103, 104 leucozonitiformis 93, 106 leveilleanus 6, 85 liquidambari 64, 70 longisporus 6, 161, 169, ludovicianus 278 macrocephalus (v) 212 magnisetulosus 122, 129 major (v) 80, 171, 172, 173, 176 manuripiensis 5, 92, 101 marthae 123, 136 martini 88 matrisdei 163, 174 mazatecus 161, 168 megalospermus 5, 124, 144 megistosporus 202, 207, 208 megistus 208 melinocephalus 164 mesites (v) 118 microdendron 123, 131, 137 microhaedinus 161, 178, 216, 217 microsporus (v) 220 minimus 93, 109, 114 minor (v) 216 minutissimus 285

molfinoanus 25 monocotyledonum 148 montagneanus 5, 202, 218, 219, 220 montagnei 219 montepiensis (v) 5, 6, 262 multiceps 92, 95 munyozii 88, 89 murrillianus 268, 264 musicola 201, 206, 214 myocephalus 269, 271 myrti 36 napoensis 164, 186, 190 nebularum 94, 112, 113, neglectus 162, 175 neosessilis 5, 6, 253, 261 nidulus 54 nigrobrunneus 122, 128 niveus 5, 269, 276 nogalesii 204, 225 nolaneiformis 16 nothomyrciae 94, 113 nothotereticeps (v) 120 nummularius 239 oaxacanus 92, 100, 101, 107 ochraceopapillatus 283 oenechinus (v) 212 oleiger 201, 207 oligocladus 141, 142 oligocystis 264, 266 olivascenticeps 162, 180 onoticus 162, 169 oreades 2, 69, 273, 274 orinocensis 186, 188 orinocensis (v) 5, 189 pallenticeps 122, 127 pallescens 202, 208 pallidocinctus 125, 157 pallipes 123, 133 panamensis 92, 96 pandoanus 6, 92, 100, 107 panerythrus 6, 202, 209 pararotula 5, 92, 97 paucifolius 222, 223 paucilamellatus (v) 137 paulensis 253, 255 peckii 122, 126 perlongispermus 263, 264 perniciosus 52 personatus, 236, 238 phaeocystis 202, 221 phaeus 161, 167 platyspermus 94, 104, 110 pleuracanthus 5, 201, 205 pleurotelloides 254, 263 plicatus 159 plumieri 269, 283 poecilus 161, 166, 167, 223 poliobasis (v) 200 polycladus 123, 135 polycystis 253, 254 polylepidis 64, 65 polyporoides 196 poromycenoides 269, 272 portoricensis 241




portonovensis 195 praeandinus 203, 232 praecox 125, 154, 155 proletarius 161, 177 pruinatus 151 pseudoarachnoides 60 pseudocollinus 270, 278 pseudocorrugatus 5, 203, 233, 236 pseudoniveus 237, 244 pseudotageticolor (v) 5, 211 psychotriophilus 93, 101 pusio 5, 164, 199 puttemansii 5, 124, 141 pyrinus 82 radicellicola 65, 74 repens 148 rhababerinus 162, 170, 171, 173 rhizomorphogeton 124, 150 rhodocephalus 210 rhyssophilus 269, 2 71 riparius 270, 282 robertsonii 125, 149, 155 rotalis 92, 105, 106 rotula 91 rotuloides 6, 92, 96 ruber 161, 182 rubicundus (v) 212 rubricosus 162, 170 rubroflavus 6, 237, 239 rubromarginatus 123, 135 rufescens 253 rufomarginatus 125, 158 ruforotula 125, 155 sanctixaverii 90 sanguineus 210 sanguirotalis 123, 136 scandens (f) 111 schini (v) 125, 133, 153 schultesii 124, 131, 148 scotophysinus 86 scototephrodes 93, 105 sejunctus 254, 260 semipellucidus 210 separatus (v) 119 sessiliaffinis 254, 257 setulosifolius 201, 204, 205 setulosus 247, 268 siccus 160 silvestris 205 silvicola 5, 270, 282 smaragdinus 263, 265 spaniophyUus 257, 259 spegazzinii 204, 234, 235 sphaerodermus 80, 82, spiculosus 237, 246 splachnoides 64, 70 splitgerberi 201, 216, 217 strictipes 270, 276 subagricola 88, 269 subalpinus (v) 125, 153 submulticeps 95 subrotula 161, 164, 217 subtomentosus 16 subtrichorhizus (v) 142, 143

superbus 165 symbiotes 284 tageticolor 161, 166 tanyspermus 93, 101, 106 tenuisetulosus 202, 220 tenuissimus 254, 258 tereticeps 6, 94, 117 tetrachrous 94, 117 thwaitesii 27 tomentellus 65, 79 tomentosus 16 tortipes 270, 284 transiens (v) 213 trichorhizus 124, 127, 147, 148, 149 trinitafis 5, 6, 162, 179, 181 tritici 151 tucumanensis (v) 119 tucumanus 161, 165, 211 umbraculum 85 ustilago 254, 256 variabiliceps 94, 114, 118 venezuelanus 237, 250 vergeliensis 92, 98 viegasii, 2, 269, 270 vigintifolius 94, 115, 116 vinosus 46, 210 violeorotalis 93, 107 virginianus 235 viridifuscus 265 weddelianus 284 wilsonii 203, 226, 227 wynnei 269 xerampelinus 6, 123, 138 xerophyticus 124, 145 yalae 64, 65 yarizae 203, 229

Micromphale 284 Mycena 273

cohaerens 247 Mycenella 1, 2, 8 Mycenites 62

Naucoria trichialis 38

Omphalina 2 Oudemansiella 1, 8

Palaeocephala 3 Peziza villosa 59 Physalacria 61, 302, 304, 315

aggregata 306, 309, 310, 311 andina 306, 308 brasiliensis 306, 307, 311 clusiae 306 concinna 306, 307, 308, 309 inflata 304, 306 langloisii 306, 310, 311 orinocensis 306, 309 sanctaemarthae 306, 310, 311 stilboides 305 subpeltata 302, 211 tenera 306, 307, 309

Physocystidium 1, 3, 8 Pistillaria 310



Index347

Pleurotopsis asperifolia 56 calospora 54 niduliformis 56

Pleurotus griseoroseus 256 liliputianus 54 niduliformis 56

Polymarasmius 62 multiceps 95 submulticeps 95

Poromycena 273

Rimbachia 61, 314, 315 palmigena 314, 315 paradoxa 314

Scorteus 62 Stereum riofrioi 313 Strobilurus 1, 8

Tephrophana 62 Trichopeziza villosa 59

Vanromburghia silvestris 205



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Marasmieae (Basidiomycetes - Tricholomataceae) || Marasmieae (Basidiomycetes - Tricholomataceae)