Vol. 17. Basidiomycetes - Marasmieae. Singer, L. 1976
Vol. 17. Basidiomycetes - Marasmieae. Singer, L. 1976
Vol. 17. Basidiomycetes - Marasmieae. Singer, L. 1976
Monograph No. 17
MARASMIEAE
(BASIDIOMYCETES - TRICHOLOMATACEAE)
by
Rolf Singer
Cl _t, Of CANCER
TROPIC
^ - ---------
"^
FLORA:
NEOTROPICA|/
TROPIC OF CAPRICORN
Publishedfor
Organization for Flora Neotropica
by
THE NEW YORK BOTANICAL GARDEN
Bronx, New York 10458
1976
Published for
Organization for Flora Neotropica
by
The New York Botanical Garden
STAFF COMMITTEE
Ghillean T. Prance, Chairman
Enrique Forero
K. P. Dumont
Alvaro Femandez-Perez, Ex officio
EDITORIAL COMMITTEE
GEOGRAPHIC CONSULTANTS
MonographNo. 17
MARASMIEAE
(BASIDIOMYCETES - TRICHOLOMATACEAE)
by
Rolf Singer
-Vk\ L t*'11
of CANCER
TROPIC
--------------------------
FLORA
NEOTROPICA/
TROPIC Of CAPtICORN /
Publishedfor
Organization for Flora Neotropica
by
THE NEW YORK BOTANICAL GARDEN
Bronx, New York 10458
1976
Published by
The New York Botanical Garden
Bronx, New York 10458
ROLFSINGER1
INTRODUCTION
of the known species of the world flora, it has become obvious that there is a need
for a new treatmentincludingthe numerousCentraland North Americanspecieswhile
omitting the exclusively temperateones. At the same time there is a need to consider
new taxa, discoveredin regions where Marasmieaehave been collected before - and
especially in regions where mycologists have never been active until after 1965, such
as the Cordillerasof Colombia and the Amazonas region of Ecuador. Including,then,
the new discoveriesof the period between 1965 and 1973 but excluding the Oudeman-
siellinae, the Marasmieaeas representedin the present monographcontain a total of
12 genera and 322 species.2
These figures confirm the impression- not unexpected to those who have
collected fungi in the tropics that the Marasmieaeare among the most importantflor-
istic elements of that vast region, followed by the Collybieae(compare my monograph
of the neotropical species of MarasmiellusSinger 1973).
AND ECOLOGY
PHYTOGEOGRAPHY
where they are introduced with plant materialfrom the subtropicaland tropical zones.
Others frequently invade the greenhousesof the temperate zones e. g. Crinipellissipar-
unae. If I exclude the collections of adventitious species from the statistics, I may say
that of the approximately 300 well defined species of Marasmiusless than 10%are pan-
tropical and as many as 229 have been recognized as occurringin the neotropics. This
hlighpercentageis not only due to the relatively large, perhapsexceptionally large rep-
representationof Marasmiusin the neotropics but, to a certain degree, also to less exten-
sive collecting and relatively few monographicrevisions(Ceylon, Zaire) in the paleo-
tropics. A similarpreponderanceof neotropical species can be observed in other genera
such as Gloiocephala,Crinipellis,Chaetocalathus. The generaPhysocystidium,Manuripia,
Epictwphus,Hymnenogloeahave thus far been observed only in the neotropics and only
Palaeocephalaexclusively in the palaeotropics.
The statistics for the whole of the Marasmieaeshow that among the species of
Marasmieae(without Oudemansiellinae)treated in the present monographthere are
species belonging in the following four groups;
Group I. Species occurringexclusively in the subtropicsand/or tropics of the Americas
or in parts of it: 274 species or 85.9%
Group II. Species occurringaside from an area in the neotropics also in some parts of
the paleotropics,Oceania or Australia: 21 species or 6.6%
Group III. Species occurringaside from some parts of the neotropics also in the tem-
perate regions or parts of them: 21 species or 6.6%
Group IV. Species occurring in the neotropics but also in the paleotropics and in parts
of the temperate zones: 3 species or 0.9%
One may argue that as the palaeotropicsare graduallyexplored so that eventually
our knowledge of the Marasmieaeflora of the paleotropics will match the knowledge we
have now of the neotropical species, the percentageof the species belongingin group II
will turn out to be largerthan indicated above. But it may also be expected that as we
begin to discover further elements of the neotropic flora thus far unknown, we are likely
to deal mostly with species rare and restricted in their distribution,and therefore likely
to be strictly neotropical. Thus, the chances that the percentagesreported above will
be substantiallyaltered after future research,appear to be small.
MYCOSOCIOLOGY
forests just after the retreat of the flooding. In these localities extensive and inten-
sive collecting of Marasmieaehas taken place (other collections have been added in
brackets).
The results are interestinginsofar as surprisinglyfew species are common to the
five localities even within their groupingaccordingto inundability. This may mean
two things, one exclusive of the other: either Marasmieae(and obviously other agarics)
are, as a group of organisms,not characteristicfor the higher "taxa" of phytosociolog-
ically definable forest communities in the Amazonas region or else they are extremely
sensitive to minor differences in microclimate,host-distributionand geographicaliso-
lation. Only these extreme views can be taken, but which one of the two is difficult
to decide.
My own experience in other regions of the neotropics tends to favor the latter
alternative,since even a glance at the phytogeographicand ecological data in the keys
and species descriptionsshows that many species are selective as far as area and associ-
ation are concerned. On the other hand, the collections of parts of the vast Amazonas
region are not extensive enough to permit us to prove this hypothesis by the lists of
species indicated below (Tables I, II).
When flying low over the hylaea with obviously basically uniform vegetation -
and at a time when a given tree with spectacularaspect during its flowering period or
with recognizablefeatures (like Bertholletia excelsa Humboldt & Beaupland)can be
pointed out and is easily traced as far as its distribution in the forest is concerned -
one can often notice that many of these trees, obviously characteristicfor the partic-
ular forest type, are widely spaced, often with distances of over 1 km between indiv-
iduals. If it is assumed then, given the dependence of the heterotrophic organismson
their hosts, that fungus species may be equally scattered in the forest and that even
repeatedvisits and very intensive collecting will not registerevery population in fruit-
ing - species cannot be detected when in the vegetative state - it becomes clear that
the necessarilyreduced collecting area and the fungi observed in it, cannot by any
means be complete enough to produce fully comparablelists.
On the other hand, if the Marasmieaewere, although widely scattered in the
Amazonas Basin, neverthelessevenly distributedamong the communities making up
the hylaea, one would expect at least some species to be common to all three terra-
firme localities. This, however, does not apply, except in the case of Marasmiius
haematocephalus, which is so common and adaptable that there is no tropical or
subtropicalforest type in the neotropics where this fungus does not occur, and the
same species has also been observed in the palaeotropicsand, adventitiously, in the
temperate zone of North America.
Somewhat more similaritybetween localities 1, 2 and 3, and 4 and 5 respect-
ively would be apparentif we were able to add all collections from the Napo river
region and compare them to all localities visited in the Beni, Mamore and Madrede
Dios regions, or all collections of the lower Para and all gathered in inundated for-
ests of the upper Amazonasand tributaries. Yet, at present this would be meaning-
less since most other localities in these regions, i e, those not included in the lists
given on Tables I and II, show deteriorationto a variabledegree; they are far from
virgin forest, bit are either second growth or partly destroyed by fire, selective wood-
cutting or cultivation. As we have shown in previous papers(Singer in Hacskaylo,
1971; Singer & Moser, 1965; Singer in Piterbarg,1966; Singer & Morello, 1960) such
stands tend to give shelter to a large number of species otherwise rare or absent, fre-
Introduction 5
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
TABLE II
Amazonas,Inundableriver-sideforest
4. Marchaspect in inundableforests along 5. May-Juneaspect of the inundablerain
the lower Madrede Di6s up to Conquista. forest in the lower Amazonas. BRAZIL:
Bolivia:Pando Para',Belem
Marasmiustereticeps Marasmiusrotuloides
M. guyanensis M. cladophyllus
M. pandoanus M. leoninus
M. dodecaphyllus [Marasmiellusstenophylloides]
M. panerythrus [Marasmiellusdefibulatusvarstrictior]
Gloiocephalalongifimbriata [Marasmiellusbisporiger]
Crinipelliseggersiivar flavipes [Marasmiellusaporposeptus]
C. excentrica [Marasmiellusnigripes]
[Marasmielluspandoanus] [Hydropusparaensis]
[Trogiacantharelloides] [Hydropusdepauperatus]
[Pluteus neophlebophorus] [Hydropussemimarginellus]
[Inocybe matris-dei] [Gerronemastrombodes]
[Pterulapalmicolal [Mycenaaosma]
[Mycenabiornata]
[Lepiota xanthophylla]
[Russulabatistae]
TABLE III
Marasmiusleveilleanus Marasmiuschiapasensis
Marasmiuscarpenterianus Marasmiusatroincrustatusvar inodorosus
Marasmiusfloriceps Marasmiusbrunneocinctus
Marasmiushaematocephalus Marasmiusdictoyledoneus
Marasmiusrubroflavus Marasmiusxerampelinus
Marasmiustrinitatis Marasmiuschrysoblepharis
Marasmiusneosessilisvar montepiensis Marasmiushaematocephalus
[Many species of Agaricales,Aphyllophorales Marasmiuslongisporus
and Gasteromycetes,see Singer(to be pub- [Micromphaleseparatum]
lished,none of them observedin area7.1 [Mycenamaculatavar]
[Pluteusnitens]
[Psathyrellacrinipellis]
[Favolaschiaselloana]
OF THE AREA
CIRCUMSCRIPTION
While I adhere in principle to the definition and limits of the area covered by
the neotropics as accepted by the Organizationfor Flora Neotropica, it may be useful
to point out here that the Marasmieaehave a mycelium protected by the host tissue
or substratumand the majority of species is found in spots with comparativelyfavor-
able microclimaticconditions. Consequently, many basically tropical species pass over
into the subtropical,a few even in the warm-temperatezone of both northernand
southern hemisphere. Because of such a relatively unalteredand gradualtransition of
the flora, it was believed to be preferableto admit a species to treatment in Flora Neo-
tropica even if it was found only in the subtropicalzone - provided its precise area
of distribution is insufficiently known - i e, occurrence in the intertropicalregion
Introduction 7
proper remains uncertain. However, in such cases a note is added stating that the par-
ticular species is marginalfor the neotropic mycoflora.
ACKNOWLEDGMENTS
The present monographhas been preparedand edited with the help of two grants,
GB 7477, GB 35471, from the National Science Foundation, WashingtonD.C.
8 Flora Neotropica
SYSTEMATIC TREATMENT
Tribe Marasmieae subtribe Oudemansiellinae Singer, Sydowia 15: 58. 1961 [1962]
TYPE GENUS.CrinipellisPatouillard.
3. Pileus with KOH turning green or (green-) black and in the latter case the hairs and/or
the hypotrichium pale greenish or greenish-melleous or greenish gray rather than pale
ochraceous-brownish to dull ocher or hyaline; (pleuro-) cystidia usually present,
scarce to numerous. sect 3. Grisentinae, p 37.
3. Pileus with KOH merely turning darker (in the same color as found in the dried
pileus, often reverting to the color of the fresh wet specimens), or not reacting; hairs
of the pileus in 5% KOH pale ochraceous brownish to dull ocher, or hyaline; (pleuro-)
cystidia present or absent.
4. Stipe at first central but soon becoming eccentric, mostly short and curved, 2-7
mm long; pileus (at first) white or whitish (excepting sometimes the center); the
great majority of the spores with Q<2,< 10.5p long. sect 2. Excentricinae, p 34.
4. If the stipe less than 7 mm long, strictly central and straight and not white, but
more frequently the stipe longer than 7 mm and then also usually central, more or
less straight, more rarely inconstantly slightly eccentric or curved when mature; if
stipe eccentric, spores as above or different. sect I. Crinipellis, p 10.
5. Spores more than 9,u broad, subglobose. subsect Macrosphacrigerae, p 10.
5. Spores less than 9u broad.
6. Cheilocystidia present and mostly crowded, but pleurocystidia very scarce
and inconstant and mostly, if at all present, in form of cheilocystidia
slightly removed from the edge, rarely pleurocystidia well differentiated in
old carpophores, but lacking in younger ones. subsect Stipitarinac, p 11.
6. Cheilocystidia and pleurocystidia both differentiated, the pleurocystidia
usually different from the cheilocystidia. at least many of them.
subsect Heteromorphinae, p 32.
Crinipellissect Crinipellis
Pileus not brightly colored and not turning green or green-blackwith KOH or NaOH,
and the hairs not greenish or greenish-melleousin 5%KOH;stipe generally central and
pigmented, insititious. The type is that of the genus.
loid, with the wall 1.4-2.8 , thick wall, not secondarilyseptate, very long, with rounded
tips.
On living tree in subtropicalforest
MATERIALSTUDIED. Known only from the type locality: BRAZIL. Rio Grande
do Sul: Estacao Sao Salvador, A. Sehnem 1917, 21 Jan 1946 (LIL: C 8339, type).
4M&P = Maerz & Paul (1930) - here as well as in all following keys and descriptions.
12 FloraNeotropica
11. Pileus with or without a small papilla or umbo, and not or in-
constantly and weakly concentrically walled-furrowed.
12. Spores 7-10 X 3.7-6p, Q 1.8-2. C. pseudostipitaria var rzesites.
12. Spores 7.7-10 X 3.5-4.5ui Q> 2.
9 C. pseudostipitariavarpseudostipitaria.
7. On dictoyledonous hosts.
13. Pileus about 40 mm broad; spores 7.5-9.3 X 5.4-6.8u; center of pileus less
or not pilose; surface more or less zoned. 13. C. missionenesis.
13. Pileus smaller, rarely reaching more than 20 mm in diameter; otherwise
as above or different.
14. Spores larger than 6.3 X 4.2p.
15. Spores up to 5.51 broad (or less broad); lamellae white or whitish
when fresh.
16. Growing on wood, woody sticks, branches or roots; cheilo-
cystidia appendiculate or not.
17. Hairs almost constantly with close secondary septa at least
in the great majority of the apical portions of the hairs of
the pileus (structure ladder-like), often with acute tips,
with walls 1.5-3, thick; cheilocystidia not crowned by
apical finger-like or thorn-like appendages, but often with
1-5 finger-like lateral and/or apical excrescenices; spores
6.5-12 X 3.5-6u, Q> 2 (rarely slightly < 2). 15. C. septotricha.
17. Hairs different, or spores and cheilocystidia different.
18. Pileus 9-19 mm broad, obtuse with depressed center;
stipe 8-16 mm long (i e about as long or not more than
twice as long as the diameter of the pileus), spores
(7-)9-10 X 4.5-5.3 p. Hygrophilous. 14. C. herrerae.
18. Pileus smaller or stipe (relatively) longer; spores as above
or different.
19. Cheilocystidia crowned with few to numerous apical
finger-like appendages or setulae, or with conspicuous
side branches or forked in lower portion, rarely a few
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 sub-
distant, spores very variable: 7-10 X 3-6.5,. 20. C. mexicana.
16. Growing on fallen leaves or leaf petioles of dicotyledonous trees
and shrubs; at least some cheilocystidia with several apical ap-
pendages.
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
Stirps Carecomoeis
What may be called stirps Carecomoeisis the group of obviously related species
no. 2-4 of this monograph.
or a mucro at the tip, thin-walledto very slightly thick-walled,or wall locally somewhat
thickened (to 1.5 u), hyaline to melleous-hyaline. Hyphae of the regularhymenophoral
tramawith clamp connections, hyaline, inamyloid. Coveringlayers: hairs of the pileus
3.5-11 p broad, with acute or rounded tips, with stramineousto melleous or (near mar-
gin also) hyaline 0.5-5.5 u thick (in KOH or NH40H), strongly to moderately strongly
(if strongly - purple) pseudoamyloid, some hairs with abrupt thickenings,all very long,
often bundled together with the longest hairs in the center of the strands,some hairs
with secondary septa but these scattered (and not ladder-like,crowded). Hypotrichium
of multi-septatehyphae which are broaderthan those of the trama proper (5-8 ,ubroad),
with hyaline, rather thin walls. Hairs of stipe much like the hairs of the marginof the
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 greg-
arious, fruiting from April until August.
MATERIAL STUDIED. CUBA, Wright, both type and authentic material (FH). MEXICO.
Chiapas: between Ixtocomitan and Solusuchiapa, 1 Aug 1969, Singer M 8805 (F), il 8811 (F).
COLOMBIA. Valle: Buenaventura, San Joaquin, 19 Apr 1968, Singer B 6230 (F).
In the neotropics we find only var carecomoeis. The varietiessubelata and
litseae which may also be separatespecies of the same stirps are confined to the South
Pacific (Philippines,New Caledonia);they grow on fallen leathery leaves.
The collection Singer M 8805 is the only one without clamp connections, with
only bisporous basidia seen and with numerousand conspicuous cystidioles on the sides
of the older lamellae. This may be a bisporous-parthenogeneticform of the same spec-
ies, or possibly a related species; it was also the only one on Gramineae. The charac-
ters of this form have not been included in the general description of the species, but
the specimen is cited under "Materialstudied."
covered by them, convex, eventually flattened and often with umbilicus, with a constant
central small papilla, the latter often inserted in the umbilicus when carpophoresmature,
1-2 mm broad. Lamellae pure white, rather narrow, subarcuate,close, free or subfree.
Stipe white, soon becoming chestnut-fuliginousfrom the base upwardsand dried entire-
ly cinnamon, white-pilose, straight, insititious, equal, 7-8 X 0.1 mm. Context very thin,
in pileus white; odor none.
Spores 9-13.5 X 5-6.5(-7)p, mostly 10-11.5 X 5.5-6.5 A,subfusoid, hyaline,
smooth, thin-walled,becoming thick-walledwhen overmature,some with a septum
near the apex when overmature,inamyloid. Hymenium: basidia 18 X 7-8.8 u;
basidioles fusoid or subfusoid;cystidia few (like cheilocystidia) or none on the sides
of the lamellae; cheilocystidia subventricose,with some knob-like short excrescencies
at the apex, hyaline, few, size of basidia. Hyphae of the stipe parallelwith each
other, filamentous, thick-walled,brown-walled,inamyloid. Coveringlayers: epicutis
of pileus consisting of hairs, these very long, 4-5(-9) broad, hyaline, thick-walled,
rounded or acute at the tip which is often thin-walled,pseudoamyloid,wall 1.5-3(3.5)
; thick, some hairs with a subapicalsecondary septum. Hairs of the stipe short to
long (the short ones subulate), thick-walled(wall 2-2.81 thick), 7-8.5 u in diameter,
all or most with acute tip.
KOH on pileus negative.
On dead leaf of a monocotyledonous plant, probably Araceae.
MATERIALSTUDIED. BRAZIL.Pernambuco:Mun.Sao Lourenco,Matado Sao, Joao,
16 Jul 1960, SingerB 3412 (F, typus).
This is similar to, though still smaller than C. careconzoeis,but has slightly
smallerspores, less pigmentationespecially in youth, and a glabrouspapilla;it grows
on monocotyledonous dead leaves in the tropical forest.
the entire cell cylindric to subfusoid or subclavate,if entire often strongly sinuose-
wavy or if branchedthe branchesmore rarely curved or crooked, and these rarely
subacute or acute, in dried material often strongly agglutinated,hyaline. Hyphae of
the trama filamentous, inamyloid, nongelatinizedwith clamp connections. Covering
layers: hairs of the pileus 4.3-10.5 broad, in KOH and ammonia stramineous-yellow-
ish to deep melleous, the paler ones in dried materialoften showing (necro-)pigment
precipitatedin brown granules,with distant secondary septa or without them, but
without ladder-likestructureeven near the mostly broadly rounded tip, very rarely
some with such a structrue,with 0.7-5.5 p thick wall, slightly but distinctly pseudo-
amyloid.
On grass culms and grass roots, often densely gregariouswith deeply immersed
stipes, in South America generally in gardens,especially seed beds, and apparently
adventitious. Fruiting in the rainy season, September and May in Asia, summer
(December to February)in South America.
MATERIALSTUDIED. ARGENTINA. Tucuman, gardenof the Instituto MiguelLillo,
Singer T 108 and 108a (LIL). NORTH VIETNAM. Hanoi,Bon 4362, 31 May 1890 (FH type):
Cai Kinhmassive: Bau Mau(Tonkin),Bouitan341b (FH, authentic).
melleous to tawny, without ladder-likesecondary septa, with the tip not acute, with
a diameter of 5-8 ,u,most frequently 5-6.5 ;,, with the wall 1.5-3 M,most frequently
1.5-2 Hthick, pseudoamyloid;hypotrichium consisting of repent broad (5-15 ,), rusty-
incrusted hyphae with clamp connections; hairs of the stipe very similar to those of the
marginof the pileus, stiff, more variableand less pigmented, pseudoamyloid but weaker
than the hairs of the pileus, 18-200 X 5-7 I in the apical, up to. 500 A,long in the lower
half, with hyaline wall 1.5-2 ,uthick, rarely with a short obtuse side-branch,otherwise
like the hairs of the pileus.
NaOH on dried pileus: More brightly ferruginous-brown,not green or gray.
On small woody twigs, apparentlyof dicotyledonous trees or vines, gregarious,
fruiting in the rainy season.
MATERIALSTUDIED. MEXICOVeracruz:7 km south of Montepio,Estacion Biologica
de las Tuxtlas,Herrera,21 June 1969, comm. SingerM 8125 (F, type).
The species is named for Dr. Teofilo Herrera,noted Mexican mycologist, collector
of the species.
trama below, and broader;hairs of the stipe (middle of the stipe) simple and long,
rarely with a short lateral branch, the tips subacute to acute, with or more rarely
without ladder-structure,with 6-10 ,udiameter, with 1.5-2.7 , thick wall.
KOH on the dried pileus darkerbrown, but not tending to gray or greenish.
On dead wood of trees and shrubs, fruiting from December to August, often in
subxerophytic vegetation or in plantations. Known hosts: Coffea arabica,Achatocarpus
nigricans.
MATERIALSTUDIED. BERMUDA.B & JDodge (NY paratypes). PORTORICO. Bruce
Fink (FH), probablybelonginghere. CUBA. Wright45, det. Berkeley& Curtis(A. stipitarius),
(FH, paratype). GUADELOUPE.Duss 484 (FH, paratypes). Duss 486 (FH, paratype):Basse-
Terre,Duss 513, 14 Jul 1901, (FH, typus). ARGENTINA. Tucuman:5 km S of Buruyacu,19
Feb 1957, Singer T 2989 (F); Posta de Lozano, 9 Feb 1966, Singer T 4995 (F).
Singer (1942) describedoriginallya "forma umbilicata ad int" differingfrom the
typical papillate form in the absence of a papilla in the umbilicus or on the disc, grow-
ing on smallersticks and twigs. Dennis (1951a) observed this form in the Arena For-
est on Trinidad(K) and described it with slightly longer spores (9-11 X 4-5 u),
This comes close to an extratropicalSouth Americanspecies which differs how-
ever in subglabrousto glabrous disc on the pileus and broaderspores. It is Crinipellis
mlolfinoalna (Speg.) Sing. which was described by Spegazzinias Marasmiusmolfinoanus
Speg., Bol. Acad. Nac. Ci. Cordoba29: 123, 1926, from dead roots of Fagaracoco. Only
the type (leg. Mlolfino212, LPS) is known. The spores are 9-11 X 6-7.5 u; the cheilo-
cystidia are entire or forked or with both apical and lateral short appendagesor diver-
ticula; the hairs are about 8 u broad and have the ladder-likestructureof C. septotricha,
with attenuate-obtuseto frequently needle-sharpacute tips, at least those at the mar-
gin of the pileus. It comes from the SierraGrande,provinciade Cordoba, at Yaci,
February 1912. This species, certainly different from C. septotricha, is mentioned
here because its host extends north into the subtropicalzone, together with the "Monte"
-vegetation characteristicfor the Sierrasde Cordoba.
ILLUSTRATION.Dennis (1951a), p1 20, fig 6.
16. Crinipellispodocarpi Singer in Singer & Digilio, Lilloa 25: 225, 1952. Fig 6.
TYPE. Singer T 833 (LIL), Argentina.
Pileus in the broad marginalzone light brown to ochraceous-ferrugineous (reach-
ing "Arab"M&Pfading to "terrapin"M&P),the entire disc region darkerchestnut to
fuliginous or the disc proper, a deep chestnut to ferruginous-fuliginousdot surrounded
by a narrowzone which is light rusty brown fresh and grayishpallid dried, and this
zone separatedfrom the marginalzone by a deeper colored (deep brown when dried)
concentric line, the extreme marginoften stramineous-pallid,hairy all over, but the
hairs long, often fasciculate in the broad marginalzone and frequently projectingbeyond
the margin,whereas in the disc zone the hairs are short and the surface appearsmore
tomentose to almost subglabrous,with some erect bristle-likehairs occasionally showing
on the papilla, the latter at first very prominent and reaching 1 mm in height, later
becoming inconspicuous and appearinglike a tiny umbo under a lens, convex with
applanatezone around the papilla or slightly depressedin the center in adult material,
the marginalzone strongly declivous for a long time and often radially sulcate, even-
tually more repandbut with circularcircumference,4-13 mm broad. Lamellaewhite
or whitish, broadertowards the stipe but still narrowto moderately broad there, close
to subdistant, mostly close or subclose, free. Stipe chestnut brown (e g 8 J 12, M&P)
below and somewhat paler (light brown) above, strongly pilose and hirsute below, less
hairy above, straight or curved, central very rarely slightly eccentric,equal or more
26 Flora Neotropica
rarely slightly broadenedat or near the insititious base, 9-28 X 0.8-1 mm. Context
white, thin, fleshy (but seemingly tough because of the hairy covering), inodorous.
Spores 7-8.2 X 5-7.2 u, with Q = 1.3-1.8, ellipsoid to subglobose, without sup-
rahilarapplanationor depression, hyaline, smooth, thin-walled,later often somewhat
thickish walled, inamyloid. Hymenium: basidia 19-36 X 7-8.8 ,, 4-spored;cystidia
none; cheilocystidia 12-35 X 6-8.5 p, with clavate, ventricose, or cylindricalmain
body and with four to nine erect or obliquely erect apical finger-likeappendages
which are 3.3-15 X 1-7 p,, mostly 3.5-5 X 1-2, , simple, more rarely forked, hyaline
(as the main body), inamyloid. Hyphae of the trama of the pileus and hymenophore
filamentous, non-gelatinized,thin-walled,hyaline, inamyloid, with clamp connections.
Coveringlayers: hairs of the pileus (marginalzone) long-filamentousor graduallytaper-
ing to a narrow-obtuseor broadly rounded tip, melleous in KOH, with thick (0.6-2 p)
wall which towards the apex often becomes thinner and more hyaline, with or without
sparseto numerous secondary septa (but these rather distant from each other, extra-
ordinarilyweakly and slowly pseudoamyloid, 3-8 , broad, mostly about 3.3-4.3 Abroad:
hypotrichium cutis-like, consisting of pale melleous to subhyaline short, rather broad
(2-8.3 u), often curved hyphal cells with clamp connections and with up to 1 Athick
walls, often with hyaline incrustationbut not pigment-incrusted.
On living and dead trunks and branchesof Podocarpusparlatorei, often ascend-
ing up to 3.5 m, gregarious,fruiting in the summerrainy season.
MATERIAL STUDIED.ARGENTINA. Jujuy:15 km NEof Postade Lozano,20 Feb 1966,
SingerT 5309 (F);Tucuman: Taficillo,6 Jan1950,SingerT 833 (UIL,types).
This species is often parasiticon Podocarplusin the subtropical-montaneto trop-
ical-montanezone of the Selva Tucumano-Boliviana. It is apparentlyclosest to C. stip-
paria.
like or forked apical appendageswhich are either erect or obliquely upright rarely
distributedon a cylindric elongation of the apex of the cheilocystidium and branch-
ing at different levels of the apex so that the apex looks like a fir tree, much more
commonly all at the tip, arisingat the same level, rarely the main body sending out
a long lateral filamentous branch, branchesand appendagesoften secondarily diverti-
culate by short wart- or knot-like protuberances,rarely the whole cheilocystidium
entire and smooth (in some preparationsno such cells found), the apical appendages
1.5-4.5 X 1-2 p and obtuse, the others 1-lOp long, the entire cell hyaline, often more
than 11 broad if appendagesor branchesare spreadingout above, not thick-walled,
inamyloid. Hyphae of the tramahyaline, inamyloid, with clamp connections. Cover-
ing layers: hairs of the pileus long and straight or flexuous, 4-11 broad, the walls
melleous-brownor melleous-fuscousto (more rarely) subhyaline, strongly pseudoamy-
loid and cyanophilous, walls 1.5-4s, thick, taperingat the apex to an obtuse tip, or
with a broadly rounded tip, without secondary septa, rarely with one or very few
scattered secondary septa, never with ladder-structure;hypotrichiumconsisting of
ratherbroad, cylindric, sometimes branching,almost thin- to moderately thickish-
walled hyphal cells which are inamyloid and melleous to brown pigmented, the
pipnent intraparietalbut also here and there incrusting the walls.
KOH on the dried pileus negative.
On sticks, branches,small twigs and fallen leaves of the evergreentough type,
always on dicotyledonous never on living host tissue, growingusually gregariously
in shaded ravinesand cloud forest vegetation but also in subtropicaland tropical
forest of the plains from Cuba to South Brazil.
MATERIALSTUDIED. CUBA. Wright(FH, type); Wright15 (part) (FH). VENEZUELA.
Dto,Federal:SierraCostera,El Junquito, + 1800 m alt, Deinnis1800, 10 Jun 1958 (K). COL-
UMBIA.Valle: Mun.Call: CerroLa Horqueta,+ 2000 -malt, 2 May 1968, R. SingerB 6903 (F).
BRAZIL. Rio Grandedo Sul: Sao Leopoldo, ex HerbarioTheissen,Fungi Austroamericani,Rick
213 (FH).
The Braziliancollection has been determined by Theissen as Marasmiusthwait-
esii Berk. & Br., but it is not identical with that species.
but with glabrousapex, insititious, equal, 23-32 X 0.5-0.7 mm. Context white, in-
odorous.
Spores 7.5-9 X 4-5.3 , most frequently 8.5 X 4.5 p, ellipsoid, hyaline, some
pale melleous in KOH, inamyloid, smooth. Hymenium: basidia 17-22 X 6-7.5 p,
4-spored; basidioles fusoid, with basal clamp; cystidia none; cheilocystidia 16-28 X
5-7.5 p, ventricose to clavate and with two to six apical finger-likeappendageswhich
may all arise at the same level, but some also often at a deeper level, or cheilocystidia
forked near the base (where they are 2.5-3.5 p broad), some slightly constricted in the
middle, appendages2-13 p long and some secondarilyforked. Hyphae hyaline, more
or less interwoven in the trama, with clamp connections, inamyloid. Coveringlayers:
hairs of the pileus acute to broadly rounded, most frequently broadly rounded or
attenuate-obtuse,stramineousto melleous-brownishin KOH,psuedoamyloid, 3.5-6.2Ip
broad, with walls 1-21 thick, without ladder-structure;hypotrichium consisting of
rather thick-walled,mostly not swollen elements which are stramineous-hyalineor
hyaline, without any incrustation,or only with a pale melleous one; hairs of the
stipe like those of the pileus, but more often acute.
KOH on dried pileus darker,not green or gray.
On dicotyledonous twigs and branches(various species of trees and shrubs)
fallen to the ground, fruiting in the rainy season.
MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yunlas, Coroico, 26 Jan 1956. Singer B
558 (F, type).
This species differs from C. herreraein relatively longer stipe and in the lack of
the rusty incrustationof the hypotrichiumnit differs from the C. patoitillardiiof the
South Pacific in the lack of glabrouszones in the less differentiateddisc region of the
pileus, the more appendiculatecheilocystidia, and in the habitat. It differs from C.
stuppariain size of the carpophoresand shape of the spores, and from C co/iiiiiuta
in the absence of pleurocystidia.
19. Crinipellistenuipilosa Singer in Singer & Digilio, Lilloa 25: 228. 1952.
TYPES:Sitger T 1119 (LIL), from Tucuman, Argentina.
Pileus ochraceous-brown,almost ferruginous("gold brown" M&P),with "cinnamon
br." M&Pfascicles of hairs in the marginalzone, center more sparselypilose, on black
ground visible among the hairs under a lens in dried condition, whereasthe ground
color appearingunder the same circumstancesamong the hairs of the marginalzone is
white, but without any naked zones, nor distinctly concentricallyzonate, conic-appla-
nate, with a prominent papilla, not umbilicate, about 17 mm broad. Lamellaewhite,
subdistant, almost broad, free. Stipe brown below, concolorous with the pileus in the
middle portion, and white at the apex, flocculose-pilosebelow, appressedlvhairy in the
middle portion, and subglabrousat the apex, subequal, or slightly taperingdownwards,
34 X 2 mm. Context white; odor of Mlicroiimplale foeticlzdn, i e, of sauerkraut.
Spores 6.8-8.3 X 2.7-3.5 ; varying from as broad as 6.8 X 3.4p (Q=2) to as
narrowas 8.3 X 2.7 (Q=3), oblong or sub-fusoid, smooth, hyaline, inamyloid, thin-
walled, some over-agedones eventually thick-walled, 1-celled. Hymenium: basidia
4-spored;cystidia none, or extremely few near the edges and like the cheilocystidia;
cheilocystidia 16-30 p long, with two to severalapical cylindricalappendagesof 5.5-8.5
X 2 p1over a clavate main body, entirely hyaline. Hyphae with clamp connections,
inamyloid. Coveringlayers: hairs of the pileus agglutinatedin bunches, pseudoamyloid,
with broadly rounded tip, 4-8.3 p broad, wall 0.5-0.7 1 thick: hypotrichium of ramose
broad sometimes incrusted hyaline elements.
Crinipellis 29
hyaline to hyaline in the apical portion, and there often with ladder-likeor at least
dense secondary septa, in the lower portion melleous to melleous-umberin KOH,
sometimes somewhat flexuous, rarely somewhat nodose, generally quite smooth and
non-incrusted,strongly pseudoamyloid, 2.7-7.5 p broad, but sometimes broadened to
up to 8 p diameter at the base, with walls, 0.5-1.8 p thick and towards the apex often
even thinner;hypotrichiumhyaline, pigment-less,more rarely pale melleous, multi-
septate with ratherbroad (12-40 X 6-16 ) hyphal cells and/or a single cell 4.5 X 4.5
p at the base of the hairs;hairs of the stipe 5-7 1broad, with walls 2-3 p thick, tips
rarelyrounded, most frequently acute, with or without ladder-structure.
KOH on dried pileus deeper rusty brown, not greeningor graying.
On fallen leaves, rarely passing over to small dead woody twigs, in subtropical
and tropical-montaneforests, gardensand plantations, always on Dicotyledones, soli-
tary or gregarious. Known hosts: Smilax sp., Quercussp., Coffea arabica.
MATERIALSTUDIED. U.S.A. Florida:HighlandsCo., HighlandsHammockState Park,
2 Sep 1942, SingerF 444a (F, type), 30 Aug 1942, SingerF 444 F. paratype). MEXICO. Oaxaca:
SierraMazateca:between Huautlade JimenezandTeotitlandel Camino,km 20-21, 11 Jul 1969,
SingerAl 8444 (F); Rancho del Cura 7 Jul 1969, SingerM 8288 (F); Road to Huautlade Jimenez,
CerroVerde, 6 Jul 1969, SingerM 8274 (F); Chiapas, 15 km W of San Cristobalde las Casas,2400
m alt, 2 Aug 1969, SingerM 8820 (F). VENEZUELA. Dto.Federal:CaracasBotanicalGarden19
Jun 1958, Dennis 1071A (K).
This is very close to C.phyllophila and C. dipterocarpi,the latter from Asia, the
former from Mexico. Crinipellisphyllophila differs in slightly largercarpophores,
slightly broaderspores, almost exclusively acute hairs on the margin of the pileus which
is more often obtuse than papillate and longer hyaline cheilocystidia. The extreme forms
of both species are easy to distinguish,but the more elongated forms of C foliicola
with longer spores - connected with the small-sporedtypical form by transitions- are
often difficult to distinguishfrom C phyllophila. It is not impossible that the latter
is merely an extreme form or variety of C foliicola, but our data on C. phyllophila are
somewhatincomplete with regardto the anatomy of the coveringlayers, and therefore,
for the time being, I describethe two as different species.
24. Crinipelliscommixta Singer in Singer & Digilio ex Singer, Rev. Mycol. 18: 11.
1953. Fig 12.
of the pileus long (e g 260 p long), 3-5.5 p broad, with 1.5-2.5 p thick wall, with mostly
slightly attenuated apex but with rounded-obtusetip (there diameter 2-3 1). Between
the hairs occasional (not dense) hyphal outgrowths of the hypotrichium are visible
(the hypotrichium otherwise not strongly differentiated from the trama of the pileus),
these outgrowths cylindrical,ventricose, or conical, 10-13 X 2-5 A; hairs of the stipe
not differentiated from those of the pileus, but still more scattered.
On woody stick (probably monocotyledonous), gregarious.
MATERIALSTUDIED. known only from the type locality. MEXICO.Veracruz:
Near Montepfo,Estacion Biologicade las Tuxtlas, 20 Jun 1969, SingerM 8083 (F, type).
Crinipellissect Grisentinae(Singer)Singer, stat nov
Crinipellissect Eu-CrinipellissubsectGrisentinaeSinger,Lilloa 8: 497. 1942.
TYPE. species. CrinipellismirabilisSinger.
Pileus either normally brown, rust-brown,chestnut, etc. and fading by de-
hydration, or else brightly pink to red, in the former case with pleurocystidia,in the
latter case without or with scarce pleurocystidia, in both cases distinctly turning gray
to green when a drop of 5%KOH or NaOH is applied to the dried surface of the
pileus and/or the hairs are greenish or green incrusted when seen under the micro-
scope in KOH or NaOH mounts.
dried condition and under a lens) dark and abrupt papilla, 2-7 mm broad. Lamellae
cream white to white, narrow,intermixed, subclose to moderately close, at first sub-
ascendant, soon horizontal, free. Stipe"russetbrown," the apex sometimes concolorous
with the center of the pileus or almost pallid, eventually almost bay, tomentose-pilose,
central or more rarely slightly eccentric, with widened but insititious base, 3-9 X 0.3-1
mm. Context white, in the region of the papilla relatively fleshy, otherwise extremely
thin, reviving,inodorous.
Spores 7.3-9.7(-10.2) X 4-6.7 j, mostly 8.5-9 X 4.7-5.7 A,,ellipsoid, few oblong, the
majority with a Q smaller than 2, with or without a suprahilarapplanation,thin- then
thick-walled(0.7 s) and if over-agedoften developinga central or eccentric septum,
smooth, hyaline, inamyloid. Hymenium: basidia 24-33 X 7-8 ,, 4-spored;cystidia 16-49
X 6.8-8.3 t, ventricose, fusoid or clavate, entire, with thin to moderately thickened wall
which is sometimes pale melleous but usually hyaline, entire or with (rather rarely) one
or two apical prongs, in some specimens incrusted by an amorphousincrustation;others
are cystidiole-like and resemble largerbasidioles, these often mucronate or with 1-2
apical prongs, hyaline, with mostly thickish wall, rarely solid. Cheilocystidianumerous
but often with mixed-in basidioles or basidia, 13-48 X 5.5-8.3 ,, varying from being
shaped like the pleurocystidiato more branchedor appendiculateand then ventricose
to clavate with one to severalapical, finger-likeappendages,others with lateral branch-
lets or knots, sometimes constricted, sometimes (rarely) capitate, sometimes incrusted.
Hyphae of the trama filamentous with clamp connections, inamyloid, thin-walled. Cov-
ering layers: hairs of the pileus golden melleous to subhyalineor hyaline, the pig-
mented ones often (but not all of them) turningolive brown, greenish or green in
KOH, but eventually, after long exposure to the alkaline medium becoming pale
greenish melleous, subhyaline or hyaline, not incrusted, 3.5-8(-11)u broad, with a
majority of acute or a majority of obtuse tips, but always both types present, with or
without a ladder-structurefrom secondary septa, the walls 1.3-2.8(-4), thick, often
flexuous, strongly pseudo-amyloid,some hairs at times intermixed with the others and
these more thin-walledand less strongly pseudoamyloid,more often obtuse. Hairsof
the stipe similar.
KOH on the surface of the dried pileus weakly or partiallygreenish,but not con-
stantly so, sometimes merely deeper brown, not black-brown,nor distinctly green or
gray.
On dead Monocotyledones, particularlyon culms of Bambuseae(Chusquea,Anrn-
dinaria), and on dead grass culms in subtropicaland tropical forests. Gregariousfruiting
in the rainy seasons.
MATERIALSTUDIED. VENEZUELA.Distrito Federal:Botanic Gardenof Caracas,19 Jun
1958, Deinnis1071 (K, F). BRAZIL.SSoPaulo: Cipo, 1 Sep 1964, Skvortzov& Alim 71393, (NY).
ARGENTINA. Tucuman;Las Lenguas,20 Feb 1951, SingerT1310 (LIL), Singer T 1291 (LIL),Sin-
ger T 1291a (LIL). INDONESIA. Java:Hohnel (FH), Zollinger2078 as "A. trichophorus"(FH, type).
The cystidia of this species are characteristicand generally numerous. In some
specimens true cystidia are formed, whereas in another collection a second, cystidiole-
like, type of pleurocystidiummay be predominant. This species is closely related to the
precedingone and should not be confused with such small species of section Crinipellis
as C. perpusilla,C. stupparia,or C. mexicata or C. commixta.
the top of the papilla, but the papilla becoming naked in many cased in old specimens,
even at its tip, the papilla in older specimens often in the middle of a shallow umbilicus
or a slight depression,4-13 mm broad. Lamellaewhite, not cream, remainingwhitish in
age, intermixed-tridymous,almost narrowto (more frequently) broad, 1.2-1.8 mm broad,
close to subdistant, mostly subclose, mostly slightly ventricose, free, more rarelyvery
narrowlyadnexed. Stipe at first slightly paler than the pileus, later "kis kilim" to "co-
coa" or "leaf mold" or 15 E/H 12, M&P,but with a particularlybright reddishchestnut
color ( e g "chutney") at the apex, or even "Maracaibo"or "Java"there, sometimes at
the apex or entirely fading to "butterscotch"(M&P),with more or less appressedrather
dense hairs over most of the surface, but towards the base often with bristle-likehir-
sute hairiness,insititious,,equal, at the apex sometimes vertically furrowed, 13-36 X
0.3-1 mm. Context white, thin, inodorous.
Spores 5.5-8.5 X 4-6 ,, most frequently 7.5 X 4.8ju, ellipsoid, with Q < 2, hyaline,
thin-walled,when over-agedthick-walled,and averaging8.2 X 5.5 A1,inamyloid, smooth.
Hymenium: basidia 26-30 X 6.7-8.8 ,, 4-spored;cystidia moderately numerous,basidio-
morphous, well differentiatedfrom the basidia in phloxine-KOHpreparations,27-45 X
7-10.7 ,, optically "empty," hyaline, smooth and entire, thin-walled, clavate or clavate-
subvesiculose;cheilocystidia 18.7-40 X 5.3-11 sj, crowded at the edges of the lamellae,
simple, forked or branching,most frequently entire at the apex, and then ventricose or
ventricose-subcapitateor subclavate,very rarely crowned with more than two appendages,
but more frequently subvesiculosebelow with a mucro, mostly somewhat opaque, but
not thick-walled,sometimes many or all with a resinousmelleous-hyalineincrustation
or hyaline incrusted at the apex only. Hyphae of the trama hyaline, with clamp connect-
ions, inamyloid. Coveringlayers: hairs of the pileus between center and extreme margin
long and brown, a few among them subhyaline to hyaline, in 5%KOH turninggreenish
or greenish melleous, 4-6 ubroad, with acute, even needle-sharptip or with attenuate-
obtuse tip, hardly ever with broadly rounded tip, the walls 1.3-2.7 u thick, pseudoamyloid,
smooth, with or without secondary septa, but very rarely approachinga ladder-like
structure,occasionally some with acute apical appendage.
KOH on the dried pileus "olive green" M&P. NH4OH negative.
On fallen branches, sticks and woody vines, rotting logs, etc. of dicotyledonous
trees, shrubs or vines in subtropicalforests and in shady wet ravines,fruiting in the
summerrainy season (December until February). Known hosts: Iresina sp (Amaran-
thaceae);Boehmeeriacaudata (Urticaceae);Solatlni sp.
MATERIAL STUDIED. ARGENTINA. Tucuman: Yerba Buena, 12 Feb 1955, Singer T
2158 (LIL, type); Pie del Periquillo, 30 Dec 1951, Singer T 1691. (LIL); Quebrada de Lules, 24
Jan 1951, Singer T 1114 (LIL).
This species was formerly not distinguishedfrom C. patouillardiiand C. conmmrita.
However, the KOH reaction in C. tucumanensisis characteristicand the bright color at
the apex of the stipe has not been observed in C. commixta.
The EuropeanCrinipelliscorticalis (Desmazieres)Singer & Clemen9onis closely
related and differs from C. tucumanensisSinger in darkerKOH reaction, somewhat
largerspores, more scattered fusoid-subampullaceouscystidia, and the host range
(Caprifoliaceaeand Oleaceae).
32. Crinipellisdusenii (P. Hennings)Singer in Pegler, Kew Bull. 21: 518. 1968.
Fig 17.
Marasmniusdusenii Hennings, Bot. Jahrb. 22: 100. 1897.
TYPE.Dusen, from Cameroons(specimen lost).
42 Flora Neotropica
33. Crinipellisrubida Patouillard& Heim, Ann. Crypt. Exot. 1: 273. 1928 ("rubidus").
Fig 18.
TYPE. Venezuela (PC).
Pileus bright pink when fresh, vinaceous when dry or dried, with a more dull col-
ored center, scarcely concentrically zonate, with the radialribs strigose clothed, almost
squamulose,with incised-denticulatemargin,convex, then applanateor depressed,with
a slight papilla in the center, reaching20 mm broad. Lamellaewhite, russet when dried.
Stipe concolorous with the pileus all over, strigose-hispid,central or equal, (5-)40-60 X
(0.7-)1 mm. Context white, thin, Odor none.
Spores 7.3-8 X 4-4.8 p, ellipsoid, smooth, hyaline, thin-walled,the over-agedones
thick-walledand with a central or eccentric septum so that the spores become two-
celled before germination,and then often rectangular,inamyloid. Hymenium: basidia
(l-)4-spored; cystidia none seen; cheilocystidia 20-28 X 6-6.7 p, clavate or ventricose,
with a crown of numerous sterigma-likeappendagesin the apical region, these erect or
somewhat oblique. Hyphae inamyloid. Coveringlayers: hairs of the pileus (marginal
zone) long, in bunches of subparallelhairs runningradiallyon the pileus, hyaline to
pale vinaceous in NH40H, 6.5-7 , broad, wall 1.5-2.7 , but thinner at the attenuated
Crinipellis 43
and not broaderthan 4.7 p, or else ellipsoid and then not more than 4 p broad, Q = 2
or larger.
TYPESPECIES.Crinipellisiopus Singer.
Only neotropical species of the subsection:
seemingly insititious or not insititious, often curved, solid, subequal or with a basal
bulb (3-5)5-13 mm long, 0.8-2 mm broad; basal mycelium at times scarcely visible,
but sometimes distinct and strigose or partiallyascendant on the lower part of the
stipe and then tomentose-vellereousand pallid. Context inodorous.
Spores 10-13.7 X 5.5-6.2p1,ellipsoid, about twice as long as broad, with thin
to slightly thickened wall, without suprahilardepression, not curved, hyaline, inamy-
loid. Hymenium: basidia 18-28 X (7-)8-9E; basidioles subfusoid or subcapitate;
cystidia none; cheilocystidia 25-90 X 7-25 I,, mostly 30-50 X 12-14 p, ampullaceous
with an apex 25-50 X 2.5-7 1, cylindric or subulate and obtuse, often capitate (cap-
itulum up to 12 ,u broad), sometimes oblique or with slight constriction(s), from one
third to equal in length as compared with the ventricose base, hyaline, with thin to
thick wall, 0.3-4 I some with a colorless, granular,thin incrustation,inamyloid,
making the edge of the lamellae heteromorphous. Hyphae thin-walledwith clamp
connections, hyaline, inamyloid. Coveringlayers: hairs of the pileus scattered or in
bunches, moderately long (35)60-200 X 5.5-12.5(-17)lp, irregularlyshaped or filament-
ous, some subventricoseor capitate, but simple, with attenuated or equal apex and
obtuse to rounded tips, with 1-1.7 , thick walls, in the lower portion hyaline to pale
violet in NH40H, more hyaline towards the tip, slowly and moderately strongly
pseudoamyloid(pinkish cinnamon in the Melzer reagent), in the lower portion often
brown or hyaline incrusted;hypotrichium with a strong violet intraparietaland in-
crusting pigment.
KOH on the dried pileus no color change, eventually slate blue or black, H2S04
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-waterlevel (to 3 m above ground).
MATERIAL STUDIED. ECUADOR. Napo, Lago Agrio, 17 May 1973, Singer B 7527a
(F) BOLIVIA. Pando: Madre de Dios, 28 Mar 1956, Singer B 2310 (LIL, Holotype; BAFC,
isotype of variety); Beni: Vaca Diez, Riberalta 31 Mar 1956, 170 m alt, Singer B 2378 (F).
Crinipelliseggersiivarflavipes Sing. may be merely a form of the type variety
since it has once (B 7527a) been found side by side with specimens showing no yel-
low pigment on the stipe. On the other hand, the following varietiesappear to be
quite constant.
40. Crinipellistrinitatis Dennis, Kew Bull. Add. Ser. 3. Fungus Flora of Venezuela
and Adjacent Countries,p 465, 1970.
TYPE.Deiinis 376 (K), from Trinidad.
Pileus "Vandyke Red" to "Acajou Red" (Ridgway), clothed with short, sub-
erect hairs, with undulatingand indistinctly striate margin,convex with a small um-
bilicus, then flattened and often depressedat the center, up to 25 mm broad. Lamellae
"vinaceousbuff" with reddish edges, ratherclose, didymous, adnate. Spore print
pure white. Stipe "Acajou Red" (Ridgway), darkerand brownerbelow, downy, cy-
lindric or slightly enlargedupwards,solid, (14-23 X 1.5-2.5 mm). Context "Vinaceous
Buff" (Ridgway), dark brown on the base of the stipe, thin, thicker and brownerat
the disc of the pileus.
Spores 5-7 X 3-4 1, ellipsoid, inamyloid, [smooth]. Hymenium: cheilo-
cystidia 30 X 8 1, cylindricalor irregularand forked, thin-walled,red. Coveringlayers:
hairs of the pileus about 300 p long, 5-6 broad, more or less hyaline, pseudoamyloid
(deep red in Melzer'sreagent) incrustedwith light red granuleswhich dissolve in 10%
KOH solution to give a bluish purple tint, distantly septate, undulating,ratherthin-
walled: hypotrichium formed by polygonal thick-walledcells, walls red in Melzer's
reagent.
On the bark of living trees, gregariouson base of Vitexsp. trunk.
MATERIALSTUDIED. Known only from the type locality. TRINIDAD. Path to
Maracaswaterfall,25 Nov 1949, Dennis 376 (K, type), not studied.
This species appearsto belong in subsectionInsignes, and on account of the rel-
atively small spores, it is different from all other species includingC. rubida (sect.
Grisentinae),with which it was originally- i e by Dennis 1951a p 430 - thought to
52 Flora Neotropica
be identical, but after the latter species was revised by Singer(1955), it became clear
that "C. rbidus sensu Dennis (1951a) is obviously a different species" which was then
redescribedas C. trinitatis by Dennis (1970). Since I have not restudiedDennlis376,
the description given above is an adaptation of Dennis's (1951a) description(data
derived from the illustrationin brackets).
The cheilocystidia are obviously different not only from those of C. rubida but
from those of C. siparuiae and C. perniciosa as well.
ILLUSTRATION. Dennis (1970), pl 5, fig 4.
briate, dry, often with projecting sterile margin,convex, attached with some central
to sublateralareaof the sterile surfaceand appearingcup-shaped,circularto sub-
elliptical, sometimes with a short pseudostipe, 1-8 mm broad. Lamellaewhite, dry pale
buff-tan (to umber when dried), rather narrowto medium broad, intermixed with
lamellulae when mature, moderately close to distant, concurrent to the stipe rudiment
to which they are adnexed or adnate, sporeprint white. Stipe only rudimentaryand
function-less,columelliform conical to knob-shaped,often obsolete, tomentose, white,
up to 1 mm long. Context extremely thin, white. Odor and taste none.
Spores 7-10.2(10.5) X 4.5-6.8 u, ellipsoid, smooth, thin-walled,becoming thick-
walled and often also with one cross septum when over-aged,inamyloid or pseudoamy-
loid (at first inamyloid in fresh material),hyaline in NH40H. Hymenium: basidia 26-
26.5 X 8-10 , 4-spored;cheilocystidia long, almost hair- or seta-like, occupying the
edges of the lamellae, but (as pleuroscystidia)also here and there appearingon the
sides of the lamellae, but may be absent in some sections, 31-90, long, entire or forked,
especially near the apical region, narrowlyfusoid to needle-shapedor cylindric, but
often with a ventricose swelling below, tip and branch-tipsacute or subacute, hyaline,
thick-walled,wall 1.5-1.7 p thick and psuedoamyloid, diameter 2-4.7 p but in basal
swellings where these are present reachingup to 7.5 p across, often deep-rootingwith-
out a septum and then reachingup to more than 100 p in length. Hyphae of the reg-
ular hymenophore hyaline, inamyloid, with some inflated cells intermixed, with clamp
connections. Coveringof the pileus consisting of usually simple, rarely forked thick-
walled pseudoamyloid hairs with a diameter of 2.7-7 p, either slightly incrusted or non-
incrusted, thick-walled,but wall somewhat thinner near apex, if the latter is attenuate
or acute, but some hairs with obtuse tip, always dense and long. Tomentum of the
stipe rudiment similar,hairs intermediatebetween those of the pileus and the hymenial
cystidia.
On palm fragmentsand frondose twigs fallen to the ground, gregarious,fruiting
in the rainy seasons in the lowland forests.
MATERIALSTUDIED. PANAMA. BarroColoradoIsland, 30 Jun 23 Aug 1952, Martin
& Welden7279, 8562, 8580, 8682 (F, IA). BOLIVIA. Beni, Vaca Diez, Ivon, 3 Apr 1956, Singer
B 2481 (LIL). INDONESIA. Java;south coast, Perigi,Donk (L, 11554). BORNEO. (K), type.
often yellowish white, narrowto broad (mostly about 1-2 mm broad), subdistantor
distant, intermixedwhen mature, concurrent to the rudiment of the stipe which
is either central or sublateral,and then often merged with the pseudostipe, the lamel-
lae attenuated and narrowlyadnexed or free. Spore print white. Stipe rudimentary
unless lateral and confluent with a pseudostipe, sometimes so poorly developed that
it is practicallyabsent but more frequently knob-like, low, white pubescent, snow-
white when seen under a lens. Context extremely thin to relativelythick, white;
odor none.
Spores 7.5-11.5(13) X 5-8 p, ratherbroadly ellipsoid, without a distinct sup-
rahilarapplanationor depression,with thin, later slightly thickened wall which is
inamyloid or in some specimens, especially when over-agedmore or less strongly
pseudoamyloid, smooth, hyaline in NH40H, often with small internal oil droplets,
cyanophilous in cresyl blue mounts. Hymenium:basidia 17-30 X 6-9 p, (2-)4-spored;
cystidia on edge and sides metuloid, 18-51 X 6-12.5 p, fusoid to ventricose, all simple
and entire or few with cristate apex or appendageslike the cheilocystidia, with pseu-
doamyloid wall 0.4-4.5 j which is in the upper portion or entirely encrusted by
coarse hyaline crystals, with mostly obtuse to subacute tip, but also often acute or
rounded;cheilocystidia similarbut more often with 1-4 apical appendagesor more
rarely effilate into an up to 100 p long, thin rostrate apex, some with a constriction
or vaguely ampullaceous,very rarely with a lateral appendagesor burge, but usually
the main body simple, ratherpolymorphous, the wall mostly inamyloid, more rarely
distinctly pseudoamyloid, thinner-walled,(wall 0.2-0.5 p thick) making the edges
heteromorphous. Hyphae: in the context of the pileus in a lower layer interwoven
and irregularlyshaped but mostly filamentous, in an upper layer more radiallyarranged
and more persistently filamentous, not gelatinized, inamyloid, hyaline, with clamp
connections. Coveringlayer of the pileus consisting of dense hairs 100-400 long
which have thick (3-5.5 p), hyaline walls which are pseudoamyloidand incrustedby
a thin, finely granularincrustation,or without it, with acute to obtuse tips; hypotri-
chium consisting of prolongationsof the hyphae of the trama where the last member
below the hair is slightly swollen and more or less cheilocystidium-likebut, instead
of the appendagebearingthe hair, divided from it by a clamped septum, inamyloid.
On fallen branchesand decaying sticks, also leaves, of mono- and dicotyledonous
plants in and near the forest and in plantations, from the coastal to the cloud forest
region. Fruiting the year around. Known hosts: Richeria grandis;Clibadiumerosum;
Chusquea sp.
MATERIAL STUDIED. USA. Florida: Dade Co., Matheson Hammock 17 Nov 1942, Singer
F 741 (F). CUBA. Wright 78 (paratype of Al. nidulus) (FH); Kotlaba (PR). GUADELOUPE.
Bains Jaunes, Duss 193, 445 (type of Crinipellis calosponrs). MEXICO. Veracruz: Estaci6n Biolo-
gica de las Tuxtlas, 7 km south of Monte Pio, 20 Jun 1969, Singer M 8066 (F); Oaxaca Huautla de
Jimenez, 12 Jul 1957, Singer M 1544 (MICH); Chiapas: above Soluchuchiapas, 1 Aug 1969, Singer
M 8793 (F), Between Ixtacomitan and Soluchuchiapas, 400 m alt; 1 Aug 1969, Singer M 8806 (F).
COLOMBIA. Valle: Mun. Cali, Horqueta, 2 May 1968, Singer B 6927 (F); Dagua Road, N above
km 25, 1900 m alt in the Cordillera Occidental, 28 Apr 1968, Singer B 6463 (F); Camino Cali-
Buenaventura, Cordillera Occidental, west slope at 1950 m alt, GuznaLn4524 (F); Saladito 15 Apr
1968, Singer B 6166 (F); Anchicaya, 26 Apr 1968, Singer B 6398 (F); Boyaca, Los Robles, 22 Jun
1960, Singer B 3489 (F); Cundinamarca, Salto, 24 Jul 1960, Singer B 3506 (F). VENEZUELA.
Dpto. Miranda between Caracas and Cumana, 5 Jul 1972, Dumont-VE 3903 (NY). Parque Nacional
El Avila, El Eden, 19 Jun 1971, Dumont-VE 234 (NY); 30 Jun 1972, Dumont etal VE 3859 (NY);
Dpto. Federal: El Junquito, 10 Jun 1958, 1870 m alt, Dennis 1165 (F). ECUADOR. Tungurahua,
Rio Topo, 30 Apr 1973, Singer B 7224 (F); Rio Verde 28 Apr 1973, Singer B 7175 (F); Pichincha,
Quito: Rev. L. Mlille 14 (Lloyd Herb. BPI); Lagerheim, Jul 1892 (Patouillard Herb., FH); Napo, Lago
Agrio, 8 May 1973, Singer B 7301 (F); B 7293 (F). BOLIVIA. La Paz: Nor-Yungas, Rio Yariza 23
Feb 1956, Singer B 1378 (LIL). ARGENTINA. Misiones: Iguazu 26 Nov 1949, Singer M 98 (LIL).
56 FloraNeotropica
6. ChaetocalathusaurantiacusSinger, sp nov
TYPE.Singer B 2201, from Bolivia.
Pileo obtuse aurantiaco;ceterum C. carneliorubersimilis. Typus in F conser-
vatur.
Pileus dull orange, finely pilose, finely long-pilosewhen seen under the dissecting
microscope dry, with hyaline or white hairs, sulcate, convex, attached to the substratum
by part of the sterile surface, up to 6.8 mm broad. Lamellaeorange-pallid,distant,
narrow,concurrent at the rear incision, not intervenose. Stipe none, but an indistinct
stipe rudiment in the rear sinus present as a tomentose patch.
Spores (8-)11-13.3 X (5.5-)6-7.5 u, ellipsoid to ovoid without suprahilardepress-
ion or applanation,smooth, hyaline, inamyloid. Basidia25-29 X 11-13 u; cystidia 32-
89 X 7-18 , of the Geopetaluhm-type, with 4-6 p thick-wall, mostly fusoid and acute to
subacute, rarely once forked, with a thick cystalline incrustation,strongly pseudoamy-
loid (exceptionally some inamyloid);cheilocystidia 30-32 X 4-6 u, narrowlyventricose
to subfusoid, often constricted in the middle, often appendiculateor with bi- to tri-
lobed apex, wall thin to slightly thickened (to 1 ,), inamyloid. Hyphae hyaline, not
gelatinized, inamyloid, with clamp connections. Corticallayer: hairs of the surface of
the pileus dense, remarkablythin (3-4 p broad) and long, wall 1-1.2 p thick, pseudoamy-
loid, tips obtuse or subacute.
On dead dicotyledonous branchesin hylaea.
MATERIALSTUDIED. BOLIVIA. Pando, Manuripi,Conquista,25 Mar1956, SingerB
2201 (F), types.
58 Flora Neotropica
LachnellasubgenusLachnella
Charactersgiven in the key above.
TYPESPECIES.LachnellaalboviolascensAlbertini & Schweinitz ex Fries.
consistingof hairs densely coveringthe sterile surfaceof the cups, up to 150 X 2.5-3 Iu,
densely covered by detersilerod-shapedcrystals, divergent,with slightly thickened wall,
thinningout towardsthe stronglytaperingwhip-likeapex and also towardsthe often
somewhatnarrowedbase, long, narrow,hyaline, continuous (non-septate).
On gall of Poll'lepis sericea.
MATERIALSTUDIED. VENEZUELA.Merida:Sierrade Santo Domingo,LagunaNegra,
3440 m alt 27 Jul 1958, Detnnis1809 (K), type (seen not sectioned;descriptionaboveaccordingto
type descriptionby Reid).
ILLUSTRATION.Reid (1961), fig 3, 1.
lamellae, regularlyinserted with lamellulae or with only a few shorter ones, with entire
margin,rather broad, collariate or broadly adnate or adnexed directly to the apex of
the stipe. Stipe gray with white apex when fresh, tending to become chestnut-fuligin-
ous or fuscous when dried, glabrous,shiny-setose, equal, insititious, about 12-13 mm
long, 0.1-0.2 mm broad. No rhizomorphs. Context thin, almost pallid, inodorous.
Spores (5.5-)7-8.8(-9.5) X 2.5-3.2(3.5) ,, oblong-subfusiformto subcylindrical,
sometimes up to three times as long as broad, smooth, hyaline, inamyloid. Hymenium:
basidia 16-24 X 5-7.5 p, 4-spored, often with a few 2-spored ones intermixed;cystidia
none; cheilocystidia numerousbut discontinuous, 16-24 p long ventricose or hyphous
but at times forked and the majority diverticulate,with irregularlydistributeddiver-
ticula 1-3.5 1 long, or with apical appendagesup to 5 1 or more in length, the whole
up to 8.5 p,in width; where cheilocystidia discontinuous, the edge either of modified
cheilocystidia, hot entire, or by basidioles and basidia, or both. Hyphae hyaline, more
rarely pale melleous in part inamyloid, or with a few vaguely pseudoamyloidstrands
in the pileus, paralleland colored in the lower portion of the stipe and there weakly
pseudoamyloid or nearly inamyloid, all with clamp connections and not gelatinized.
Corticallayer: epicutis of pileus formed by branched-diverticulatehyphae as in M.
androsaceuswith the diverticula 1-5.5 ulong; these elements not hymeniformly arranged
and not or scarcely incrusted by pigment, but hyaline to melleous (by an intraparietal
pigment) both here and in the hypodermium;some of the epicuticularelements cystidi-
form, elongated and diverticulateon the upper side.
On dicotyledonous sticks and wood particles fallen to the ground in Alnetum,
solitary in the montane forests.
MATERIAL STUDIED.ARGENTINA. Jujuy: Lagunasde Yala, 1900-2300 m alt,
13 Feb 1966, Singer T 5131 (F), type, 11 Feb 1966, Singer T 5059 (F).
MATERIAL STUDIED. JAMAICA. Castleton Gardens, 28 Oct 1902, Earle 269 (NY), type.
Spores 5-9.3 X 2-4 u, mostly 6.5-8 X 2.5-3.2 /, oblong, some ellipsoid, smooth,
hyaline, inamyloid. Hymenium: basidia 17-22 X (4-)5.5-7.5(-8) ,1, 4-spored;cystidia
none; cheilocystidia with a main body 13.5-19.5 X 4-6.5 p, versiformbut often clavate,
with diverticula(which are sometimes fasciculate on a finger-likeappendage)mostly
severalto numerous, lateral and apical, 1.2-5 , long, wall hyaline, thin to firm; some
occasionally smooth cheilocystidia with an apical mucro, some basidioles and basidia
occasionally intermixed with the numerouscheilocystidia. Hyphae of the pileus-trama
and the regualrhymenophoraltramahyaline, thin-walled,ratherloosely arrangedbut
not gelatinized, inamyloid or at times vaguely pseudoamyloid,hyphae of the stipe
parallel,here many thick-walledand melleous in KOH,pseudoamyloid or merely vag-
uely pseudoamyloid, all with clamp connections (but often numerous secondary septa
present). Corticallayers: epicutis of the pileus not hymeniform, its elements cylindric
to vesiculose, hyaline or somewhat fuscous in their lower portion laterally and/or ap-
ically diverticulatewith diverticula2-6.5 X 0.8-1.8 1, many of them pigmented by an
incrustingpigment but this incrustationthin, resinous-sheet-like,fuscous or fuscidu-
lous; the same slight to ratherstrong pigment incrustationalso on many elements of
the hypodermium,elements elongated but often branched;covering of the stipe char-
acteristic but somewhat variable,according to the state of development of the stipe,
of three types: (1) hairs 20-100 X 2.7-3.7,, with thin to thick + pseudoamyloidwall
and acute apex, entire and hyaline, fragile but rigid, incrustedby fine crystals, (2) hairs
12-33 X 5-15(-25) , cylindric, ventricose, clavate or vesiculose, thick-walled(wall up
to 1.2 j thick), hyaline to pale brownish, pseudoamyloid or not, (3) hairs(12-)50-170
X 4.2-9.5 /, long-filamentousor graduallytaperingto an obtuse tip, thick-walled,hy-
aline, the inner wall-layerpseudoamyloid,the outer wall layer and the interior inamy-
loid, sometimes incrusted by hyaline, amorphousmaterial,generally of the appearance
and structureof Crinipellishairs, wall 1-4 thick but sometimes much thinner, and
less pseudoamyloid in the apical portion, more numerousthan type (2) and (1) in
the lowest portion of the stipe.
On dead, dicotyledonous leaves, fallen to the ground in tropical forest as well
as the tropical-montaneforests, from Mexico to Brazil, also in tropical Africa and Asia.
MATERIALSTUDIED. MEXICO. Morelos:NWof Tepoztlanat 2,200 m alt, 27 Jun 1969,
SingerM 8208 (F). BOLIVIA. Beni: Vaca Diez: Guayaramerfn, 9 Mar1956, SingerB 1 776/III
(LIL); 9 Mar1956, SingerB 1779 (LIL), type of M. hyalotrichus;17 Mar1956, SingerB 2094
(LIL). BRAZIL. Pernambuco:Camaragibe,6 Jul 1960, SingerB 3117 (BAFC);B3111 (BAFC);
Paraibo:Joao Pessoa, 11 Mar1960, SingerB 3342 (BAFC). CEYLON. Hakgala,Jan 1914, Petch
3920 (FH), isotype. ZAIRE. Bas-Congo,Kisantu,1901, J. Gillet (BR).
When this species is growing on oak leaves, it is sometimesdifficult to disting-
uish from the precedingspecies unless the hairs of the stipe are studied. These are
always entire, smooth in M. hakgalensis;also, the spores are slightly narrowerin the
latter.
ILLUSTRATIONS. Singer(1965) pl 6, fig 8-9; Petch (1947), fig 12.
lamellae, some lamellulae), with entire edge and not intervenose,adnexed, not collar-
iate. Stipe chestnut to deep chestnut below, with white apex, insititious, pilose at apex,
less or scarcely so below (hairs not visible in fresh condition without a lens), subequal
to slightly taperingupwards,insititious, about 40 X 0.3 mm; dark rhizomorphsaccomp-
anying the carpophores. Context thin, inodorous.
Spores 5.8-6.5 X 2.5-3.5 g, ellipsoid to oblong, hyaline, smooth, inamyloid. Hy-
menium: basidia 19-21 X (4-)5.5-6 i, 4-spored;cystidia none; cheilocystidia like the
elements of the epicutis. Hyphae in pileus-tramaand the regularhymenophoraltrama
mostly thin-walled,not gelatinized, hyaline, inamyloid, with clamp connections, in the
stipe parallel,some with thin, some thick (1.3 p) or intermediatewall, multiseptate,
varyingbetween hyaline and brown, inamyloid, the brown ones very indistinctly pseu-
doamyloid. Corticallayers: epicutis of the pileus consisting of not hymeniformly ar-
arrangedversiformelements, these mostly erect and often cylindric to clavate, inamy-
loid, 7-10 X 4-7 p (main body), sometimes branching,some hyphous, all or most divert-
iculate, the hyphous ones often arisingfrom the clavate ones, hyaline, setulae or divert-
icular equal or more often taperingupwards,to a narrowbut obtuse tips, 3-5.2 X 1-(1.2)
L;hypodermiumcontrastingwith the epicutis in its pigmented elements which are
rather irregularin shape, mainly radially-elongatedwith hyaline to brownish wall and
thinly incrusted by an inconspicuous sheet-like spadiceouspigment-incrustation;cover-
ing of the stipe consisting of at apex denser and more conspicuous, towards base less
conspicuous hairs of two types (1) long (100-180 X (2-)5-10l), the broadenedbase-
portion up to 25 , long, the apex thin and piliform, 2-3 Abroad, with obtuse tip, at
times with secondary septa here and there, all walls inamyloid, hyaline and thin; (2)
short elements about (9-)24-26 X 8-10 , with thin to moderately thickened wall which
is hyaline to melleous hyaline, correspondingto the basal portion of the long hairs,
rounded obtuse above and not effilate, also inamyloid, all entire.
On dead leaves of Quercus,solitary. Mexico.
MATERIALSTUDIED. MEXICO. Morelos:NWof Tepoztlan,27 Jun 1969. SingerM
8250 (F) type.
This differs from other species growing on oak leaves in the characteristicsof the
hairs of the stipe which are not thick-wallednor acute (as in M hakgalensis)nor long
and diverticulate(as inM. splachnoides)but at least many of them characterizedby
the thin-walled,effilate apex. The spores are rather small in comparisonwith those of
M splachnoidesand relatively broaderthan those of M. hakgalensis.
Marasmiuscryptotrichus differs from the following species (which is closest) in
the presence of effilate hairs on the stipe, the more fuscous-grayrather than tan, och-
raceous brown color of the pileus and the less diversifiedcheilocystidia as well as by
the host.
ium; basidia 15-17 X 4.5-6 A, 4-spored, few 2-sporedat times; basidioles fusoid; cystidia
none; cheilocystidia versiform, many diverticulatelike the diverticulateelements of the
epicutis. Hyphae of the pileus-tramathin-walled,with clamp connections, hyaline, fila-
mentous, inanyloid to very slightly pseudoamyloid in places; hyphae of the stipe all
parallelwith each other, some dark spadiceous, some hyaline, some thick-walled,some
thin-walled,some pseudoamyloid, some inamyloid (even in the apex of the stipe), 1-6
, broad, with scattereddepth and some secondary septa. Corticallayers: epicutis of
the pileus consisting of diverticulatehyphae and broom cells of various shapes, with
diverticula2-10 mostly about 3 ,uhigh,hyaline; hypodermiumat least in the middle
of the pileus distinctly spadiceous to sepia incrustedby sheets of pigment, its elements
irregularlyinterwoven;covering of the stipe consisting of numerousbut mostly small
(4-29 X 5-11 I, rarely longer) obtuse hairs, those of the upper portion of the stipe
all thin-walledand hyaline to pale stramineous(wall 0.3-0.5 j thick), those of the
lower portion of the stipe as above, or, fewer, hyaline to brown and thick-walledbut
otherwise similar, wall here 1-3 p thick.
On dead roots and woody rootlets of trees (Ficus?), gregarious. Brazil.
MATERIALSTUDIED. BRAZIL. Pernambuco,Dois IrmSos,14 Apr 1964, SingerB 4078
(F), type.
This differs from the type variety mainly in inodorous carpophoresand perhaps
slightly less elongated spores (6.5-7.5 X 3.5-4 j) and somewhat paler marginand stripes
along the ridges of the sulci of the pileus.
On oak leaves in Mexico.
MATERIAL STUDIED.MEXICO.Chiapas:15 kmWof SanCristobal de lasCasas,2400
m alt 2 Aug1969,SingerM 8816 (F), type of variety.
18. Marasmiustomentellus Berkeley & Curtis, Jour. Linn. Soc 10: 298. 1868.
Fig 39.
TYPE. Berkeley Herbarium(K), from Jamaica(not studied).
Pileus fulvous to vinaceous brown, subrugulose,not sulcate, later more or less
sulcate, glabrous,convex, obtuse or with a small and obtuse umbo, 1-3 mm broad.
Lamellae becoming concolorous with the pileus in the herbarium,few (? 10) distant,
more rarely subdistant, with few lamellulae(1-5), not intervenose, not collariate, ad-
nexed. Stipe badious to blackish, pubescent, curved, rising from a common black
rhizomorphs,solid, subequal with slightly to distinctly narrowedapex, 1-4 X 0.2-0.6
mm; rhizomorphsblack, long, as thick or thicker than the diameter of the stipe.
Spores 11-11.5 X 4-5 ,, cylindric to oblong with mostly recurvedbasal portion
and flattened inner outline when seen in profile, smooth, hyaline, inamyloid. Hymeni-
um: basidia 27 X 6.5-8.5 ,, 4-spored;cystidia scattered, 60 X 6.5-7.5 1, ampullaceous
or fusoid, thin-walled,hyaline. Hyphae filamentous, inamyloid except for the pseu-
doamyloid rind-zoneof the stipe, without clamp connections, hyaline except for the
hypodermialzone of the pileus and the rind-zone of the stipe, which are pigment-in-
crusted, not gelatinized, those of the pileus radiallyarrangedand those of the stipe
parallel. Corticallayers: epicutis of the pileus consisting of irregularlyarranged,ir-
regularlydiverticulatedelements which are hyaline to subhyaline, forming a very thin
layer; liypodermiumvery broad, consisting of interwoven hyphae with conspicuous
brown pigment incrustation, 2.5-8 , broad, with walls up to 0.8 , thick; covering of
the stipe consisting of few erect hairs (which are merely continuations of the hyphae
of the rind of the stipe, about 25 X 5 ,', entire, with the wall about 0.7 ,uthick) and
seta-like hairs 50-125 X 5-7 ,, with hyaline, thick (1 ,) pseudoamyloidwall.
On heaps of dead twigs in accumulatedlitter. Southern U.S.A. and West Indies.
MATERIALSTUDIED. U.S.A. Louisiana:La Salle parish,May 1936, FrankSmith, comm.
F. A. Wolf (FH).
Marasmius sect Hygrometrici Ktlhner, Le Botaniste 25: 60. 1938 (as Hygrometricae).
TYPE SPECIES. Marasmius hygrometricus (Briganti) Saccardo.
Pileus usually small and well pigmented with an epicutis formed by broom cells
(usually of the Rotalis-type) which, in portions of the pileus form a hymeniform lay-
80 Flora Neotropica
er, often some of the epicuticularelements in chains, often some of them smooth,
(but never all of them); spores usually medium-sized;lamellae free to adnate, not or
incompletely collariate, rarelyabsent; cystidia on the sides of the lamellae usually
present or cheilocystidia dimorphic. Hyphae with clamp connections, inamyloid.
Stipe central, always insititious. On dead vegetable matter.
the upper three quartersor two thirds covered with rod-like setulae (Rotalis-type).
Hyphae of pileus and hymenophoraltrama hyaline, filamentous, interwoven, with
clamp connections, moderately thick-walled to thin-walled,not gelatinized, inamyloid.
Hyphae of the stipe parallel, in cortex brown and moderately to rather strongly thick-
walled, the paler to subhyalinehyphae of the core thin-walledto moderately thick-
walled, inamyloid. Cortical layers: epicutis of the pileus hymeniform, consisting of
broom cells much like cheilocystidia, some hyaline, some chestnut brown to stram-
ineous, thick-walledabove if pigmented;there are also some smooth, thick-walled,
strongly pigmented bodies, similarto the broom cells in shape but rising from the
hypodermium;they do not reach the level of the epicutis and remain somewhat im-
bedded; covering of the stipe: in the middle of the stipe of an adult specimen con-
sisting merely of some external hyphae which are nodulose-verruculose,some incon-
spicuous brown excrescences which are smooth, very scattered;some dermatocystidia
like the hymenial cystidia, also smooth.
On dead, fallen, dicotyledonous leaves and fruits, known hosts: Crescentia,
Quercus.
MATERIAL STUDIED.MEXICO: Morelos,NWof Tepoztlan,2200 m alt, 27 Jun 1969,
SingerM 8251 (F). CUBA. 5 milesE of Santiagode lasVegas,1 Sep 1904,Earle184 (NY)type.
Another narrow-sporedspecies is Marasmtius capillipes Saccardo(= Miarasmtius
pyrinus Ellis) which differs in size, color and having fewer lamellae. All collections
seen by me come from the temperate regions of Europe and North America,always
on deciduous (not coriaceous) leaves (of rosaceous trees, Sryringa,Acer, Popuhlts,etc.):
neither the fungus nor the hosts have been found in the native vegetation of the
neotropical regions but this species is very close to AMarasnnius crescentiae and might
yet show up in the subtropicalbelts, introduced into gardensor parks. It was indicated
for Chile by Spegazzinibut his descriptivedata and illustration refer to Marasmnius ilicis.
the broom cells, the dermatocystidiamuch like the hymenial cystidia; covering of the
stipe consisting of scattered dermatocystidia,much like those of the hymenium but
much smaller,graduallydisappearingin old specimens.
On dead leaves and twigs. Known hosts: Salix humrboldtiana, Spathodea campa-
nulata
MATERIAL STUDIED. HAWAII. Manoa Valley, Oahu, Jan 1969, R. D. Goos (F). AR-
GENTINA.BuenosAires:La PlataRiver(no morecarpophores presentin the type envelope),
(LPS),type.Tierradel Fuego:Ushuaia,authenticmaterialbut only somestipesleft (LPS).
This species is evidently of wide distributionbut has been rarely collected, prob-
ably because of its small size. However, Spegazzini'soriginal description(aside from
the spore measurementswhich are obviously wrong) fits the Hawaiianmaterial so well
and the lack of lamellae is so characteristicfor this one species in section Hygrometrici
that I have no doubt but that they are identical with each other.
23. Marasmius echinosphaerus Singer, Bull. Jard. Bot. 34: 325. 1964.
TYPE.Schmitz-Levecq169, from Zaire.
84 FloraNeotropica
Pileus blackish spinulose on yellowish ground, dried dark bay, reddish chestnut
(e g 15 E 12, M&P),strongly sulcate over half the radius, echinulate-spinuloseall
over, subglobose to hemispheric,often higher than broad, 1.5-3 mm broad, 1.5-5 mm
high. Lamellae white, with the edges concolorous with the pileus but when young
only pale ocher, subdistant(about 10-17 through-lamellae),about one fifth of the
diameter of the pileus broad, attenuate-subfreeto free. Stipe concolorous with the
pileus below but at first with whitish apex, later and dried almost uniformly chestnut
brown, macroscopicallyglabrousbut finely echinulate-flocculose-roughwhen seen under
a lens, insititious, equal, 10-15 X 0.2-0.5 mm. Context rather fleshy and relatively
thick in the middle of the pileus, thin at margin, inodorous.
Spores 6-8.5 X 3.2-4 p, ellipsoid to oblong, smooth, hyaline, inamyloid. Hymen-
ium: basidia 18.5-26 X 6-6.5 u; basidioles fusoid; cystidia both on the edges and in
the lower part of the sides of the lamellae of two types: (1) 14-34 X 2.5-5.5 , ampull-
aceous, lanceolate or acicular,subacute, acute, or somewhat obtusate, hyaline, thin-
walled, often or rarely covered by a finely granularhyaline mass, (2) broom cells of
the Rotalis-type, 10-34 X 4-9 p, vesiculose to subvesiculose,beset with divergentset-
ulae 0.5-3 p projecting, melleous-hyalineto brown above, mostly hyaline below, wall
0.2-0.5 p thick. Hyphae of the pileus-tramahyaline, filamentous, thin-to firm-walled,
not gelatinized, with clamp connections, inamyloid, few swollen to 6.7 p broad; those
of the stipe strictly different abruptly changingat the apex to parallel,hyaline in the
core, but incrusted by a coarse succineous brown pigment in the rind, all firm-to
thick-walled,inamyloid. Cortical layers: epicutis of the pileus consisting of a hymen-
iform layer composed of four kinds of elements (a) broom cells like the cystidia of
type (2), above these reachinga wall diameterat their apex of up to 5 P, often broadly
clavate ventricose of subvesiculoseand more pigmented (these dark, thick-walled,even-
ly distributedamong the others; (b) similarelements agglutinatedin parallel rows con-
sisting of seriate catenulate broom cells and making up the ornamentationof the pil-
eus, in the marginalzone often replaced by large, one-celled broom cells 17-81 X 10-
16 p; the catenulate cells are about 19-36 X 7-20 p, becoming shorter towards the
terminalcells, many of them without setulae or merely rough;(c) occasional dermato-
cystidia correspondingto the cystidia of type (1) or slightly smaller:(d) broom cells
of the Siccus-type or entire, 12-20 X 5 p, often ramose-forkedinto branchesabout 2 P
broad, or simple, with obtuse tips, main body cylindric to slightly ventricose, chestnut
colored, scarce and sometimes wanting;hypodermiuma palisade where the terminal
cells constitute the epicutis, but here less or not pigmented, less or not thick-walled
and occasionally widened into endocystidia of the shape and sixe of the hymenial
cystidia of type (1); covering of the stipe consisting of elements similarto the epicutis
elements of type (a) and (d), the latter scarce and of some versiform rough to sub-
smooth brown, thick-walledelements 18-23 X 3-5 M.
On fallen branches,sticks, leaves and petioles of dicotyledonous trees.
MATERIALSTUDIED. ARGENTINA. Jujuy: Posta de Lozano, 1500 m alt, 21 Feb 1966,
Singer T 5429 (F). ZAIRE. Haut-Katanga,Kipushi,Kipopo, 1250 m alt, Schmitz-Levecq169 (BR)
type.
Singer (1964b), fig 14, E-H.
ILLUSTRATION.
The African Marasmius kroumirensis (Patouillard) Saccardo & Sydow has less
ornamentedpileus, adnate and fewer lamellae, broadly umbonate pileus and shorter
stipe, according to Pegler's(1966) type analysis.
However, I have found a fungus in Ecuadorwhich is somewhat intermediatebe-
tween M. kroumirensisand M. echinosphaerus. The materialis too scanty to determine
Marasmius 85
obtuse, about 1.5 X 1.5 p, both main body and diverticulahyaline, the upper portion
often forked. Hyphae: hymenophoral trama regular,consisting of filamentous(1.3-2,u
diam.) hyphae which are not gelatinized, inamyloid, those of the pileus radiallyarranged,
otherwise similar,also inamyloid but having pseudoamyloidcontents at times, opaque
and long, often irregularlybranched;all hyphae with clamp connections, those of the
stipe black incrusted, also inamyloid. Cortical layer: epicutis of pileus consisting of
globose, pedicellate cells which are either hyaline or with a porphyry sap, aside from
that, wall colored deep sepia by an intraparietalpigment, globose part (without ped-
icel) 6-14 I in diameter,pedicel up to 13 , long, these elements in hymeniform arrange-
ment with the contents partly pseudoamyloid;hypodermiumconsisting of filamentous
hyphae with membranapigment.
On dead dicotyledonous leaf in cloud forest (2200 m).
MATERIAL STUDIED.BOLIVIA. La Paz: Nor-Yungas, Cerro Uchumachi, 25 Feb 1956,
Singer B 1444 (LIL), type.
This species differs from the Hygrometriciin the smooth epicuticularelements,
from the Epiphylli in the pigment and the absence of pleurocystidia,from Pararotulae
in inamyloid hyphae and in non-collariatelamellae, and finally from the genus Gloio-
cephala in the absence of long hairs or oleocystidia and from all these groups in the
characteristic,almost unique cheilocystidia. It seems thereforejustified to erect a
special section for this species even though the section has thus far only a single rep-
resentative.
ILLUSTRATION.Singer (1965), pl 6, fig 11.
Marasmiussect Marasmius
RotulaeFries,Epicrisisp 384. 1838.
CollariatiBataille,FloreMonographiqueMarasmiusd'Europep 26. 1919.
TYPESPECIES. M. rotula (Scopoli ex Fries) Fries.
Pileus generally small, white or pigmented and often with a knob or papilla in
the middle which in mature specimensis often located in an umbilicus and surrounded
by a pallid ring zone, with hymeniform epicutis which eventually may become slightly
disorganized,consisting of broom cells which at their apex or the entire upper portion
are covered by minute to ratherlong erect or divergentsetulae without long conspic-
uous hairs;spores small to large, ellipsoid to oblong, mostly (or always) acyanophilic;
lamellae subcollariateto mostly distinctly collariate;hyphae with clamp connections,
mostly pseudoamyloid at least at the apex of the stipe, more rarely all inamyloid;
stipe centrally attached to leaves or wood, sometimes to black rhizomorphswhich are
92 FloraNeotropica
attached, on leaves or wood, always distinctly insititious; cystidia none, rarely present
but inconspicuous;stipe and rhizomorphsmostly glabrous,in a few species pilose.
15. Pileus white or whitish when fresh or mostly so, but never
flesh pink ever;sporesdifferent.
16. Pileuswith a ratherpale or pallidumbo;spores12.5-14.2
X 7.5-9,u;lamellaeabout seven. 44. M. psychotriophilus.
16. Pileus with a dark umbo; spores 5.5-7.8 X 2-3(-4)u;
lamellae14-20 (if fewer compare"6" above).
17. Pileusabout 6 mm broad;lamellaeabout 20; on oak
leaves. 45. M. cundinamarcae.
17. Pileus up to 5.5 mm broad;lamellae11-17; on dead
leavesother than oak. SouthernSouth America(if in
NorthernNorth America.compare"11" above).
46. M. leucorotalis.
I. Pileus in mature specimensnot white in the largermarginalregion (if only old, dried
pale avellaneous,pale ocher or pale argillaceouspilei seen, compare also "2" above).
18. Pileus sepia or mummybrown, 2-3 mm broad;lamellaeon edges discolorous(con-
colorouswith the pileus);spores11-14 X 3.5-5 ; on bark. 47. M. arimanus.
18. Not combiningthese characters.
19. Pileusgray to beige (betweenblackishgrayto pale cinereouswhen fresh).
20. Sporeslargerthan 9,ulongand 4.5g broad,(see also subsectionPenicillati).
48. M. griseofuscescens.
20. Sporessmaller.
21. Lamellaewith cinereousto darkgray edge; spores7-9 X 3.5-4.7p; cheilo-
cystidiawith mostly sepiacoloredsetulaeand hyalinemainbody. Mexico.
49. M. scototephrodes.
21. Lamellae with white edge which may become brownish after drying;
spores 5.5-8 X 2.5-4ui;cheilocystidiaeither entirely hyaline or if setulae
arebrown, mainbody with hyalinewall but fuscous cell sap.Colombia.
50. M. leucozonitiformis.
19. Pileusnot so colored.
22. Pileus when fresh purplishred or red or pale flesh pink with dark colored
centraldot on papilla;lamellaeeitherwith discolorousedge and collariumor
evenly flesh-pinkall over.
23. Spores 8-11 X 4.5-5u;edge of lamellaeand collariumred;stipe 15-35 mm
long. 54. M. carminis.
23. Sporesnarrower;edge of lamellaenot discolorous;stipe sometimeslonger.
24. Pileuspale flesh pink;lamellaeconcolorouswith pileus. 53. M. carneotinctus.
24. Pileusdifferently colored; lamellaewhite with white edges but collar-
ium + discolorous(see "25" below).
22. Pileus neither red nor pale flesh pink but either brown or ferruginous or
orange or violet-lilac (if gilvous or pale argillaceous, see "2" above).
25. Spores over 10p long and up to 3.5g broad; pileus white when young but
soon becoming brown with a paler zone around the central dark dot and
on the margin. 51. 1M.tanyspermus.
25. Spores less narrow.
26. Pileus violet-lilac; lamellae ? 12; spores 6.2-9.5 X 2.5-3w; on dicotyle-
donous leaves in tropical-montane forest. 52. M. violeorotalis.
26. Pileus differently colored. Key II.
Key II
1. Spores voluminous: 8.5-13 X (5-)5.7-9(-10.3),(; pileus orange ferruginous, small, with a
pale, avellaneous-cinnamon zone around a central, black dot, conical-truncate when
fresh, barrel-shaped when dried; collarium ochraceous. Cloud forest species. 55. M. buzae.
1. Spores less voluminous or pileus and collarium different.
2. On dicotyledonous wood (if on woody stems of vines compare M. variabiliceps var
separatus) in tropical forest and montane woods.
3. Stipe rising from a common rhizomorphic strand, yellowish; pileus brown, less
than 1 mm broad (if broader and stipe black, see subsect Penicillati); spores not
reaching 4 p broad. 56. M. populiformis.
3. Stipe often accompanied by rhizomorphs; although not attached to them but
insititious on the substratum.
4. On slender twigs, thin vines(or occasionally on small pieces of Aextoxicum).
5. Pileus Brussels Brown (R) 1.5-2 mm broad; spores 6-7 X 2-2.6p; lamellae
8-10. 57. M. minimus.
94 FloraNeotropica
5. Pileus not Brussels brown, larger; spores broader; lamellac more numerous
(see "9" below).
4. On rotten logs and their fragments, on fallen branches or twigs, never on
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 (com-
pare M. rotuloides and related species, also - Key I, "5"). Tropical
montane and subtropical zone. 46. M. letlcorotalis var leucozonites.
12. Pileus deep orange-ferrugineous (see "19" below).
10. Spores generally somewhat larger; pileus with or without a white or pallid
zone around the central dot, of variable size.
13. Lamellae (17-)20(-22); collarium often brown; pileus broader than
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 collar-
ium concolorous with the sides of the lamellae; hyphae of
pileus not or scarcely pseudoamyloid; spores 8.2-11.2 X
3.5-4p1;pileus 2-2.5 X 2 mm with vertical sides. Rain forest
species. 68. M. tereticeps.
19. Lamellae about 9-14 and the collarium and often also edges
of lamellae discolorous, ochraceous, orange or brown or else
lamellae more numerous (16-21); hyphae of pileus weakly
to distinctly pseudoamyloid; spores and pileus as above or
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. Marasmiusmulticeps Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 298. 1869.
Pol)ymarasmlius
muLlticeps(Berkeley& Curtis)Murrill,North AmericanFlora 9: 286. 1915.
Polymarasmiussubnulticeps Murrill,North AmericanFlora 9: 286. 1915.
Marasmiussubimulticeps(Murrill)Saccardo& Trotter, Sylloge Fungorum23: 155. 1925.
TYPE. Wright160 (K), from Cuba.
Pileus white but in age and on drying sometimes tending to become brownish ful-
vous, with chestnut-fuliginousto black umbo both fresh and dried, sulcate, but the
umbo and papilla smooth, hemisphericor convex, umbilicate, mostly with a distinct
papilla or umbo in the umbilicus or at least a somewhat elevated, dark dot present,
1.5-6 mm broad. Lamellaewhite, distant, with (9-)11-14(-16) through-lamellae,all
entire or 1-3 lamellulaepresent, ratherbroad, shallowly but distinctly collariate. Stipe
black (at first with pallid or brown apex), glabrous,insititious with a ring-jointand
risingfrom black rhizomorphs,equal or taperingupwards, 5-25 X 0.2-0.5 mm . Black
rhizomorphsglabrous,creepingor arbuscular-ascendant, their main stem 0.6 mm thick.
Context of the pileus white, thin.
Spores 6.2-8.5 X 2.2-3.7,, ellipsoid to ellipsoid-oblong,smooth, hyaline, inamy-
loid. Hymenium: basidia4-spored, basidioles fusoid, some remainingsterile (cystidioles);
cystidia none; cheilocystidia with main body hyaline, 12-13 X 6.8-7 p, beset with div-
ergent setulae 4-5 X 0.8-1 ,u,also hyaline, attenuated to a subacute tip. Hyphae hyaline
in the pileus-trama,chestnut-fuliginousin the stipe, there paralleland thin-to thick-walled,
many distinctly pseudoamyloid but in the pileus-tramaand the hymenophoraltrama
(which is regular)mostly thin-walledand only slowly and vaguely pseudoamyloid,those
of the rind of stipe and rhizomorphsstrongly forking-interlaced,those of the core of
the stipe and rhizomorphsoften granular-incrusted by a hyaline incrustation,all with
clamp connections and filamentous but some swollen to 10 p. Cortical layers: epicutis
of the pileus hymeniform or subhymeniform,consisting of broom cells 12-17 X 9-12.5
It, of the Rotalis-type, hyaline, with numeroussetulae which are either rod-shapedor
conical or even rounded-verruculose,0.8-4 X 0.6-1.9 u, the main body often laterally
extended or else vesiculose, inamyloid.
On dead logs and trunks. Cuba, Belize, Guatemala.
MATERIALSTUDIED. CUBA. Wright160 (FH, K), type; Fungi CubensesWrightiani132.
(FH). GUATEMALA. PlateauaboveChiatla,18 Nov 1959,Degener,comm.Petrak,II (BAFC).
Dennis (1951c) data on the type of M. submulticepsshow that it is conspecific
with A. multiceps.
ILLUSTRATIONS. Dennis (1951 b) fig 11; (1951 c) fig 28.
not or vaguely pseudoamyloid;those of the stipe and the region above its insertion
distinctly pseudoamyloid,here parallel, all hyphae with clamp connections. Cortical
layer: epicutis of the pileus nearly hymeniform, consisting of two types of epicuti-
cular broom cells, (1) of the Rotalis-type, with the main body 11-14 X 7-11 U, cla-
vate to vesiculose, sometimes 2-3-forked, setulae 1-2 X 0.5-0.7 p, divergentor obliquely
erect in the subapicularregion, hyaline, (2) of the Siccus-type, 20-34 X 6-13 p,, the
latter type dense and numerous in the center, widely dispersedto rare in the margin
al region of the pileus, pigmented, rarely subhyaline, with walls 1-4.5 u thick, setulae
apical and erect or sometimes in form of side-branches,finger-likeor obtusely coni-
cal, 2-15 X 1.5-4,, some of the "broom cells" without appendagesor setulae,
Over dead wood and leafmold in tropical rain forest. Pantropical(?), at least
in the Amazonas and the Congo regions.
MATERIALSTUDIED. BRAZIL. Amazonas:"Panure"(=Sao Jer6nimo),Spruce 75 (K),
type. ZAIRE. J. Ghesquiere1355 (BR).
ILLUSTRATION.
Singer (1964b), fig 15, D, E, F.; Berkeley (1856) pl 5. fig 3.
34. Marasmiusrotuloides Dennis, Trans. Brit. Mycol. Soc. 34: 415. 1951.
TYPE.Dennis 232, from Trinidad.
Pileus white or whitish in tile marginalzone, towards the center mostly flushed
flesh-ocher to yellow-beige or light ochraceous to pallid, with a white or paler ring
zone around the central dot which varies from almost flat to low-umbonate, dried
more beige to pale grayishbeige, the central dot or umbo concolorous with the zone
around the white zone, not chestnut either fresh or dried (the flesh-ocherzone fresh
about 12 C 6/7 or "yellow beige" M&P),long and deeply sulcate, glabrous,strongly
convex with an umbilicus and with or without a low umbo in the umbilicus, 9-17 mm
broad. Lamellaewhite, distant (13-16 through-lamellae,no lamellulae), broad, distinct-
ly collariate. Spore print white. Stipe dark red-brownto usually black, at first with
white apex, glabrous, smooth, insititious or widened into a very narrow, insititious
disc, equal, not attached to rhizomorphs,20-37 X 0.2-0.9 mm. Context in pileus
white, thin, inodorous.
Spores 5-7 X 2-3.5 ,, ellipsoid to fusoid-oblong, smooth, hyaline, inamyloid.
Hymenium: basidia 17-19 X 4-5,u, basidioles fusoid; cystidia none; cheilocystidia hy-
aline thin to firm-walled,otherwise like the epicuticularbroom-cellsin shape and size.
Hyphae with thin to often thick walls, filamentous, pseudoamyloid, with clamp con-
nections, hyaline in pileus-tramaand the regularhymenophoraltrama, not gelatinized.
Cortical layers: epicutis of pileus hymeniform, with broom-cellsof the Rotalis-type,
main body 12-27 X 8-26 ,, obpiriform to vesiculose, some laterally extended and even
hyphous at times, hyaline, inamyloid, with a wall thin- to 1 p thick, over most of the
surface beset with divergentsetulae which are rod-shaped,hyaline to light stramineous,
1-2 X 0.88.
On dead, fallen, dicotyledonous leaves in tropical forest.
MATERIALSTUDIED. TRINIDAD.Cerrode Aripo, 23 Oct 1949, Deinnis232 (K), type.
BRAZIL. Para,EstanciaPirelli,7 Jun 1966, SingerB 4231/I (BAFC), B 4224 (BAFC);Belem,
10 Jun 1966, SingerB 4252 (BAFC);B 4269 (BAFC).
Dennis (1951a) pl. 19, fig 14.
ILLUSTRATION.
38. Marasmiusapatelius Singer, Bull. Jard. Bot. Bruxelles 24: 332. 1964. Fig 44A.
TYPE.Vanderyst695, from Zaire.
Pileus at first white (?), dried beige to dull pale brownish or beige-ochraceouswith
paler margin,long sulcate, glabrous,strongly convex, umbilicate, with a small umbo or
papilla in the umbilicus, 1-3 mm broad. Lamellaewhite, distant (9-12 through-lamellae,
no lamellulae), broad, some forked, distinctly collariate. Stipe bister to blackish with
white apex, glabrous,setiform, smooth, insititious, some not developingpilei (telepodia),
some associated with very thin, black rhizomorphsbut not attached to them, 10-20 X
0.1-0.2 mm. Context of the pileus white, very thin.
Spores 6.8-10 X 2.8-4.8,, ellipsoid, fusoid or oblong, some cylindrical, smooth,
hyaline, inamyloid. Hymenium: basidia 18 X 5.7 u; cystidia none; cheilocystidia like
the epicuticularbroom cells. Hyphae of the pileus trama densely packed and inter-
woven, basically runningradially,parallelin the stipe, pseudoamyloid, with clamp con-
nections. Cortical layer: epicutis of the pileus hymeniform, consisting of hyaline or
subhyaline broom cells of the Rotalis-type, main body 12-24(-35) X 10-19(-25) , ves-
Marasmius 99
iculose, often pedicellate to 10 u deep, setulae hyaline to pale melleous 1-2.5 X 0.5-1.5 p.
On coriaceous, fallen, dicotyledonous leaves, in tropical forest. South America
and tropical Africa.
MATERIALSTUDIED. VENEZUELA. Sucre:alongRio Media,15 Jul 1972,Dumont
et al VE 5145 (NY). ZAIRE. Bas-Congo,Kisantu,Vanderiyst695 (BR), type.
Singer (1964b) fig 16, H- I.
ILLUSTRATION.
39. Marasmiusrotalis Berkeley & Broome, Jour. Linn. Soc. Bot. 14: 40. 1873.
TYPE. Twaites 810, from Ceylon.
Pileus whitish to pale cream, soon (even before being dried) turning dirty beige or
brownish(e g "oakwood," "wigwam"),center "coffee" M&P,grayishbeige or concol-
orous with the margin, sulcate, glabrousor sometimes rivulose-subscabrousin a zone
around the disc (and this zone consequently more pallid even in dried materialbut not
constantly so) the umbo itself not discolorous-darkbut pallid to light avellaneous-argill-
aceous, convex, umbilicate, with a small, low umbo or an obtuse papilla in the umbilicus,
1.5-5 mm broad. Lamellaewhite to pale cream, with white to (dried) slightlydiscolorous
edge (concolorous with the marginof the pileus), distant (13-17 through-lamellae,no
lamellulae or more rarely 1-2 lamellulaepresent) broad, collariate, the collariumwhen
dried also concolorous with the gill edges. Stipe black, blackish bister, or very dark
chestnut, with white apex, glabrous,smooth, insititious, equal, usually accompaniedby
thin, dark rhizomorphsbut never attached to them. Context very thin, inodorous.
Spores 6.8-8.8 X 3-4 p, oblong, smooth, hyaline, inamyloid. Hymenium:basidia
19 X 5.5 u; cystidia none; cheilocystidia like the epicuticularbroom cells. Hyphae of
pileus-tramainamyloid, with clamp connections. Cortical layers: epicutis of pileus
hymeniform, broom cells 10-23 X 5-15 s,, of the Rotalis-type,hyaline, with thin wall
and brown-fuscoussetulae which are divergent, 1.3-3.5 X 0.6-2.5 1, mostly 1.8-2.8 X
1.2-1.4 g, rod-shaped,obtuse.
On dead coriaceous leaves of dicotyledonous trees, pantropical. Known host:
Quercuscolumbiana.
MATERIAL STUDIED.COLOMBIA. Boyaca.:Tunja,Arcabuco,28 Jul 1960,SingerB
3560 (BAFC).CEYLON.Peradiniya, Thwaites810 (K),type. ZAIRE.Yangambi, ReserveIsa-
lowe,470 m alt Louis14934 (BR).
This species is close to Marasmiuscundinamrarcae, M. apatelius, and M. leuco-
rotalii In the Colombianoak woods it may most easily be confused with M. apatelius
which has fewer lamellae and M. cundinanarcae with more numerouslamellae and
smallerspores.
ILLUSTRATION. Singer (1964b), fig 16, E-G.
blackish at first with white apex, glabrous, equal, insititious, about 11 X 0.1-0.2 mm.
Context of pileus white, thin, inodorous.
Spores 5-6 X 2.5 p, ellipsoid, smooth, inamyloid. Hymenium: basidia 5-5.5 p
broad; cystidia none; cheilocystidia like the epicuticularbroom cells. Hyphae of the
hymenophoraltrama and the pileus-tramahyaline, many of these distinctly pseudoamy-
loid, with clamp connections. Cortical layer: epicutis of the pileus hymeniform, con-
sisting of broom cells of the Rotalis-type, main body 17-20 X 12 u, hyaline, setulae
short and divergent.
On dead dicotyledonous leaf in tropical rain forest.
MATERIALSTUDIED. BRAZIL:Para, EstanciaPirelli,7 Jun 1966, SingerB 4199 (BAFC),
type.
hymeniform, consisting of broom cells which are 12-35 X 10-33 ,, of the Rotalis-type,
vesiculose and sometimes broaderthan long, sometimes longer than broad, sometimes
pedicellate, setulae dow to the middle (or lower) of the cell, rod-shaped1-2 X 0.3-1 p,
all hyaline.
On fallen dicotyledonous leaves in tropical-montaneforest. Bolivia.
MATERIALSTUDIED. BOLIVIA. La Paz; Nor-Yungas,Rio Yariza,23 Feb 1956, Singer
B 1414 (LIL) type.
Singer(1965) pi 7, fig 17.
ILLUSTRATION.
form, broom cells of the Rotalis-type e g 28 X 16 p,, beset with hyaline to (in herbar-
ium materialmostly) pale melleous brownish setulae 202.7 plong, divergent.
On leaves of dicotyledonous trees, frequently (but not exclusively) on Phoebe
porphyria.
MATERIALSTUDIED. ARGENTINA. Tucuman,Sierrade San Javier,CiudadUniversit-
aria, 1000 m alt, 15 Jan 1955, Singer T 2018 (LIL), type.
The precedingvariety differs from this in being less pigmented, smallerand with
fewer lamellae and the central dark dot as well as the stipe becoming paler on drying,
also in slightly smallerand relatively broaderspores. Both differ from the type variety
by a few rather secondary characters;the type variety appearsto be intermediatebe-
tween the other two varieties.
The binomialMarasmiuspluvialis was intended for a species I have now redes-
cribed as Marasmiushvlaeicola Singer (no. 63). Because of an unfortunate printer's
error, the description of Marasmiusleucozoniiteswith its type designation(T 2018) has
been repeated under the headingMarasmiuspluvialis Singer (1965, p 163) so that
formally it becomes a synonym of Marasmiusleucozonites.
Young specimens of Marasmiusplatvspermusmight be looked for here or under
M cundinamarcae. It differs however in habitat and broaderspores.
47. MarasmiusarimanusDennis, Trans. Brit. Mycol. Soc. 34: 418. 1951 (as Xi. ari-
mana).
TYPE. Dennis 391, from Trinidad.
Pileus sepia or mummy brown, striate-sulcateto the center, convex with a
shallow umbilicus, 2-3 mm broad. Lamellaepale buff, distant (7), moderately broad,
attenuated to each end, attached to a shallow collarium forming little more than a
ridgeround the apex of the stipe, with sepia colored edges. Stipe almost black, slen-
der, wiry, smooth and polished throughout. Context thin, white, membranous.
Spores 11-14 X 3.5-4 ,, tear-shaped,inamyloid. Hymenium: cheilocystidia like
the epicuticularbroom cells, 8 lpbroad. Coveringlayers: epicutis of pileus formed by
pear-shapedelements with reddish brown processes.
On bark of dead logs, Trinidad:Arima, MissDora Kong, 28 Nov 1949, comm.
Dennis 391 (K), type.
I have studied the material superficially;it consists of a fragmentwhich I did
not dare section. It is not fully clear whether the epicuticularbroom cells and cheil-
ocystidia are of the Rotalis-type or the Siccus-type. The species seems to be close
to MarasmiusfuligineorotulaSinger. The description is adopted from Dennis' original
account. It would suggest that even if the broom cells were considered as being of
the Siccus-type, the spores would be more elongated and the carpophoresmuch
smaller in Dennis' species.
ILLUSTRATION. Dennis (1951a) pl 23, fig 5.
umbo correspondingin extension to the blackish dot inside the umbilicus, convex
otherwise, 1.5-3.2 mm broad. Lamellaepale cream, 10-11, ratherbroad, with white
edge, collariate. Stipe umber with white apex, glabrous,insititious, 20 X 0.1 mm. No
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 epicuticularelements. Hyphae
of pileus, hymenophoraltrama hyaline, with clamp connections, in stipe (apex and
hyaline interior of lower portion) moderately strongly pseudoamyloid,somewhat thick-
walled, otherwise inamyloid to very weakly pseudoamyloid. Coveringlayers: epicutis
of pileus hymeniform, consisting of cells of the Rotalis-type, hyaline, 9-37 s,broad,
beset with hyaline or (in places) pale melleous setulae, these small (1.2-1.8 X 0.8-1 a),
divergent.
On dead dicotyledonous leaves in tropical rain forest.
MATERIALSTUDIED. BOLIVIA. Beni: Vaca Diez, Guayaramerin,16 Mar1956, Singer
B 2052 (LIL), type.
When seen in the white stage, the carpophoresmay be mistaken for those of
Marasmiuspanldoalus or Marasmiusoaxacanus which do not become brown when
mature and have broaderspores.
visible (under a lens) surface, showing the trama, glabrous,sulcate, excepting the cen-
ter, strongly convex, umbilicate, with a low papilla in the umbilicus, about 5 mm broad.
Lamellaelilac-pallidwith pale lilac edge and pale lilac ground, distant (+ 12 through-
lamellae, no lamellulae), broad, collariate. Stipe black, glabrous, flexuous, insititious,
55-58 X 0.2 mm. Context inodorous.
Spores 6.2-9.5 X 2.5-3 p, mostly 7.5-8.5 X 2.5 j,, fusoid to subcylindrical,smooth,
hyaline, inamyloid. Hymenium: basidia 14-15 X 4.5-5.5 p; basidioles fusoid; cystidia
none; cheilocystidia like the epicuticularbroom cells but more often subhyaline. Hy-
phae in the pileus hyaline (KOH), inamyloid, with clamp connections. Cortical layer:
epicutis of the pileus hymeniform, broom cells of the Rotalis-type, 13-21 X 6-14 ,,
vesiculose, more rarely clavate, setulae mostly pale melleous, short, rod-like, 1-1.5-(2)
x 0.4-0.8-(1) 1,
On dead dicotyledonous leaves in tropical-montaneforest, CordilleraOccidental,
Colombia.
MATERIALSTUDIED. COLOMBIA.Valle, near Daguaroad above km 25, at 1900-2000
n alt, 28 Apr 1968, SingerB 6450 (F) type.
wall in the main body, with hyaline pedicel, main body about 14-26 X 8.5-16.5m.
Growing on the bark of Duranta serratifolia,living tree, about 3 m above the
ground, gregariousbut not cespitose.
MATERIALSTUDIED. ARGENTINA. Tucuman:DepartanentoTaff, CiudadUniversit-
aria, Sierrade San Javierat 1200 malt. 24 Mar1960, Singer T 3591 (LIL), type.
This seems to be a habitat form with somewhat paler pileus and very slightly
smallerspores, rare. It should not be confused with AL.leucorotalis which differs in
having fewer lamellae and narrowerspores.
This commonspecies of the ArgentineNorthwest is confined to the lower range
of the mountains. The precedingspecies (no. 58) differs in less numerousand there-
fore more distant lamellae, slightly different colors, generallyshorter and thinner stipe
and more consistently acanthophysoid(equally and always divergentlyshort setuloid)
epicuticularelements. The rhizomorphsare either less or not developed in M. dode-
caphyllus.
Among the holarctic species, MI.wettsteinii differs in growing on conifer needles,
slightly smaller,especially narrowerspores and a slight differencein colors. M bul-
liardiidiffers in growing on frondose leaves, fewer lamellae and even narrowerspores.
entire, collariate. Stipe black, glabrous,setose, insititious, 7-17 X 0.2-0.3 mm. Context
very thin, white in the pileus, inodorous.
Spores 5.5-6.5 X 4-4.5 p, ellipsoid, smooth, hyaline, inamyloid. Hymenium:
basidia 17-23 X 5-6.7 1, 4-spored;cystidia none; cheilocystidia in mature specimens
mostly smooth, vesiculose, 21-30 X 8.5-18 j, thin-walled,hyaline, some, a minority
broom cells, these with a main body as above and apical diverginghyaline setulae,
1.5-2.5 s long. Hyphae of the pileus trama hyaline, with clamp connections, inamy-
loid, often very broad (3-19 p diameter in mature specimens);thin- to firm-walled;hy-
phac of the stipe dark pigmented, parallelwith each other, vaguely pseudoamyloid.
Cortical layer: epicutis of the pileus hymeniform, consisting of broom cells of the
Rotalis-type, main body 10-35 X 8-19 , often (the longest) long-pedicellate,thin-
walled. hlyaline,setulae succineous-brown,rodshaped, 1-2.2 p long, divergingand
coveringa wide area of the cell, smooth cells very scattered or absent.
On dead standing branchesor roots of leguminousshrubsand trees in the Andean
re,ionl, gregarious.
MATERIALSTUDIED. ECUADOR.Pichincha,Rio Machaingaro,
2600 m alt, 20 May 1973,
Singer B 7538 (1), type.
This differs fronmM. icnbularumt in the not uniformly colored pileus, broaderyet
shorter spores and, as from mlost species in this group, in the fact that the fully mature
specimenllshows a large majority of smooth cheilocystidia.
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.JuanFernandez: pathto El Camote,about400
Masatierra,
m alt, 18 Mar1955, SparreF 56 (S) type; Coquimbo:Fray Jorge, 16 Apr 1967, SingerM 6515
(SGO). As for CentralandSouthChileancollectionsstudiedsee Singer(1969, p. 103).
This species is somewhat marginalas far as the neotropics are concerned since
the only locality known in the subtropicalbelt is a fog forest in North Chile.
(saturated deep ferruginous)or brown, dried between "tortoise shell" and "cognac" M&P
(brown without a reddish tinge) glabrous, sulcate to a narrowwhite or paler often indis-
tinct zone in umbilicus, the white zone surroundinga black spot which is flat and becomes
dark gray on drying, barrel-shapedand practicallyas high as broad, 2-4 mm broad. Lam-
ellae white with white to slightly browned (dried) edges and brown-marginedcollarium,
narrowly collariate, subclose to subdistant,(10-13), broad. Stipe black with at first
white apex, glabrous, insititious setose, often long, 15-60 X 0.1-0.3 mm; no rhizomorphs.
Spores 9-10.7 X 2.7-4,u, rarely as short as 8.5, and as broad as 4.2,. Hymenium:
cheilocystidia like epicuticularelements. Hyphae of pileus pseudoamyloid. Corticallay-
ers: epicutis of pileus with Rotalis type elements, main body 17-26 X 15-22A, vesiculose;
setulae rod-shaped,2-3.5 X Il,.
On dead dicotyledonous leaves fallen to the ground in tropical forest.
MATERIAL STUDIED. ECUADOR. Napo. Shushufindi, 15 May 1973, Singer B 7460 (F).
BOLIVIA. La Paz: Nor-Yungas, Rio Yariza, 23 Feb 1956, Singer B 1395 (LIL) type.
Key II
1. Pileusred to brown-redor red-brownor purplewhen fresh, either so coloredwhen wet
or becomingso when dryingout in situ.
2. Sporesthree timesas long as broador narrower(Q73).
3. Sporessmallerthan 7.7 X 3,; risingfrom rhizomorphs. 85. M. polycladus.
3. Spores largeror stipe not rising from rhizomorphsbut directly from the sub-
stratum.
4. Sporesin theirgreatmajorityover 14p long.
5. On leaves and litter of all kinds, tropical; spores (9-)14-18.5 X
(1.5-)2.5-3.3/; lamellaedistant (11-15); edge of the lamellaepink or con-
colorous with the pileus, the latter tendingto be a rich deep brownwhen
dried. 86. M. rubromarginatus.
5. On wood in subtropicalvegetationof South America;spores still more
voluminousor lamellaesubdistant(16-17).
6. Lamellae >14. 87. M. marthae.
6. Lamellae<14. 88. M. sanguirotalis.
4. Sporesnot reaching14/uin length(see "10" below).
2. Sporesless than three times longerthan broad(Q < 3).
7. Stipes risingfrom rhizomorphs;both stipe and rhizomorphwith seta-likehairs(if
stipeglabroussee no. 80, p 131).
8. Pileus pink without a white zone around the dark purplelow umbo; spores
7.5-8 X 2.8-3U. 89. M. microdendron.
8. Pileusred with a white zone arounda white to blackishcentralpapilla;spores
7-9 X 3.5-4.2,u. 90. M. hippiochaetes.
7. Stipe either rising directly from the substratumor from rhizomorpns;in either
caseboth stipes and rhizomorphs(if present)glabrous.
9. On wood; stipe risingdirectlyfrom the substratum;pileuspurplered, 3-10 mm
broad;spores4-7,ubroad;context of pileus and hymenophoraltramapseudo-
amyloid, collariumoften discolorous(brown). 91. M. xerampelinus.
9. On fallen leavesor smalltwigs or stems; stipe risingdirectly from substratum
or from dark rhizomorphs;pileus as above or smaller;hymenophoraltrama
124 Flora Ncotropica
Key II1
1. Stipes rising from rhizomorphs (some may rise directly from the substratum).
2. On wood.
3. Spores 7.3-11 X 3.5-5.3p or somewhat larger.
4. Pileus 1.5-2 mm broad; stipe 3-4 mm long; all or most stipes rising from
rhizomorphs (see "6" below).
4. Pileus 1-8 mm broad; stipe 12-40 mm long; most stipes rising directly from the
substratum (see "11" below).
3. Spores 8.2-9 X 2.2-3.5(-5)u or 7-8 X 4-5 . 101. M. trichorhizus.
Marasmius 125
2. On leavesand monocotyledonousculms.
5. Spores9.5-13.2 X 4.5-6.2u; edge of lamellaediscolorous(see Key I above, "18").
5. Sporesless broad.
6. Pileus orange-ferruginous,drying brown ("cocoa" M&P);edge of lamellae
white and cheilocystidiawith hyaline setulae; spores 8.5-11 X 3.5-4.1M.On
dicotyledonousleavesin the Amazonasregion. 103. M. robertsonii.
6. Pileus orange with deeper colored papilla,sometimessomewhatbleachingin
the marginalregion; edge of the lamellae intermittentlyor indistinglydis-
colorous, some cheilocystidia with pigmented setulae; spores 7.5-11(-13) X
3.5-5 p. Mostlyon monocotyledonousleavesand culms. 102. M. crinisequi.
1. All stipesrisingfrom the substratum.
7. On wood, generallyof Dicotyledones.
8. Edgeof lamellaenot discolorous.
9. Carpophoresaccompaniedby black rhizomorphs.pileusoften ratherpale dirty
orange to pale cinnamon buff when wet, without a white zone around the
deep browncentralpapilla.
10. Spores9.5-12.3u long. 104. M. dicotyledoneus.
10. Sporesusuallyshorter(see "11" and "12" below).
9. Carpophoresnot accompaniedby black rhizomorphs.pileus orange with or
without a white zone around a darkerpapilla or umbilicus;spores 7.5-10l
long, about twice as long as broador slightlybroader(if sporesmoreelongated
compareno. 105 (M.rhizomorphogeton).
106. M. graminumvarschiniand varsubalpinus.
8. Edgeof lamellaediscolorous.
11. Spores4.7-5.3, broad.SouthernSouth America. 107. M. praecox.
11. Spores3.5-4.3,ubroad.Floridato Amazonas. 108. M. ruforotula.
7. On dicotyledonousand monocotyledonousleaves,on culmsof Bambuseaeand twigs
of other Monocotyledones,not on woody materialof dicotyledonous trees and
shrubs,rarelyon stemsand frondsof Pteridophyta.
12. Edgeof lamellaenot discolorous;on monocotyledonoushosts.
13. Spores relatively elongated, 10-13.2 X 3.8-4.2p; pileus ratherpale and dull
orangeocher, dried deeper colored (rufescent).Amazonasregion (see key II
above "18").
13. Sporeseither smalleror broader;pileuswell pigmentedorangeto ferruginous
orange or orangered, sometimespallescentin age. Subtropical,tropicaland
subtropical-montane zone of both hemispheres.
106. M. graminumn vargraminumand varculmisedus.
(Note: If on Bromeliaceae, compare A bromeliacearum, Key II, "11" no.
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 "ferrugin-
ous" (M&P), becoming brown ("Arab" or "hazel" to 15 J 10, M&P)with
a pallid zone around a low umbo; hyphae of the trama of the pileus
inamyloid;rhizomorphseither none or scarceand inconspicuous;lamellae
12-13; collariumwhite. 110. M. pallidocinctus.
15. Spores 6.8-8.5 X 3.5-42; pileus deep rufous to rufescent-orange, on drying
becoming brown ("kis kilim" M&P) with a light brown zone around a
deep brown central dot, exumbonate; hyphae of the trama of the pileus
pseudoamyloid; inside of collarium tending to be brown; lamellae about
10; rhizomorphs accompanying the carpophores (if papillate and without
rhizomorphs, see "16" below). 109. M. brunneocinctus.
14. Edge of the lamellae discolorous at least at full maturity.
16. Spores 9.5-1 3.2 X 4.5-6.2/p (see key I above, "18").
16. Sporessmaller(cf also M. praecoxno. 107, see "11" above).
17. Spores 6.5-10(-10.2) X (2.5-)3-4.5 ; at least the dry pileus with a low
umbo or papilla;lamellae7-11. On dicotyledonousleaves.
111. M. rufomarginatus.
17. Spores 7.3-10 X 4-5u, i e relatively broader; central dark dot of the
pileus generally not elevated; lamellae 10-13. More frequently on
monocotyledonous than on dicotyledonous leaves. 112. M. foliicola.
126 Flora Neotropica
72. Marasmiuschrysochaetes(Berkeley & Curtis),Jour. Linn. Soc. Bot. 10: 297. 1868.
Crinipellischrysochaetes(Berkeley& Curtis)Singer,Lilloa8: 461. 1942.
TYPE.Wright162, from Cuba.
Pileuswhite, dried tending to "pinkishbuff' Ridgway,and developingsometimes
a brown umbo, sulcate, glabrous,convex, umbilicate,with a small umbo in the um-
bilious, 1.2-2 mm broad. Lamellaewhite, distant (about ten throughlamellae,no
lamellulae),ratherbroad, distinctly collariate. Stipe fulvous or light brown, shining,
insititious,glabrous,smooth, 22-25 X 0.2 mm (dried), some telepodes. Context ex-
tremely thin.
Spores 8-9 X 3.5-4.2u, ellipsoid-oblong,smooth, hyaline, inamyloid. Hymenium:
basidia4-spored,basidiolesfusoid; cystidia none; cheilocystidialike the epicuticular
broom cells but generallyhyaline and inamyloid. Hyphae of pileus-tramaand hymen-
ophoral tramahyaline, thin, ratherthin-walled,inamyloidwith clamp connections; those
of the stipe parallel,pseudoamyloid. Corticallayers:epicutis hymeniformor subhymen-
iform, broom cells of the Siccus type, 14-28 X 7-9u, with apical obtuse appendages
which are often finger-likeand 2-3/ubroad, entire wall firm to thick (to 3.5 ,)- walled,
pseudoamyloid.
Subcespitoseon dead leaves of the evergreenkind. Cuba.
MATERIALSTUDIED.CUBA. Wright162 (FH) type.
I have not studied part of the type deposited at Kew but Dennis indicates
similarcharacteristicsas given above althoughhe finds the sporesslightly narrower
(9-10 X 3-3.5g).
ILLUSTRATIONS: Singer(1942)fig 1,g. Dennis(195lb) fig3.
shaped, in other cases erect and cylindric, blackish fuscous, thick-walled,setose, but
obtuse, e g 11-14 X 5.5,.
On dead leaves and culms of monocotyledonous plants, especially Bambuseae
(Banibusa,Chusquea),but also frequently on Bromeliaceae(Aechmea) and palms.;
smaller carpophoresoften on grasses.
MATERIALSTUDIED. TRINIDAD. St. Joseph, 30 Nov 1949, Deinnis409 (K). BOLIVIA.
La Paz: Nor-YungasCarmenPampa,2000 m alt, 26 Feb 1956, SingerB 1544 (LIL);B 1545 (LIL).
ARGENTINA. Tucuman:Capital,22 Jan 1955, Singer T 2077 (LIL); 10 Feb 1955, T 2149 (LIL);
T 2150 (LIL); Tafi del Valle, 17 Feb 1956, Singer T 4081 (F); Jujuy: Lagunasde Yala, 14 Feb
1966, Singer T 5127 (F). VIETNAM. Hanoi, Kes6, 31 May 1890, Bon 4397 (FH), type; Demange
6 7 (H). INDIA. Gurdespur,JaggatPur, 15 Aug 1941, SultanAhrmad509 (FH). CEYLON. Bo-
tanic Gardens,Peradiniya,1 Nov 1914, Petch 4238 (K, FH). typus of M. griseoviolaceus.
ILLUSTRATION.Singer (1958a), fig 48 [as AI. griseoviolaceus]; Dennis (1951a) 19,
fig 18 [as MI.griseoviolaceus/.
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.
MATERIALSTUDIED. ARGENTINA. Misiones:Iguazfu,Refugio Yaguarete,16 Apr 1957,
Singer M 1047 (LIL) type.
ILLUSTRATION.Singer (1965) p1 10, fig 4.
and at times few, numberingfrom one forked setular up to 12 setulae, 4-12 X 1.3-1.7
,, rarely reachingup to 2.8s diameter at base, either taperingupwardsto a subacute
or obtuse tip, or more frequently cylindric with broadly rounded tip.
On the thick central nerve of dicotyledonous leaves (Inga ? edulis).
MATERIALSTUDIED. BOLIVIA. La Paz: Nor-Yungas,Coroico, 12 Feb 1956, SingerB
1098 (LIL), type.
This species, in dried condition, has the appearanceof Marasmiusdicotyledoneus,
I. gramfiumtvar schinii,etc. Only few spores were found in the type collection and
it may be expected that the maximum-minimumrange is still wider than indicated
liere.
Marasmiiusboliviae differs from the Argentine forms mentioned above, in the
more voluminous spores, the discolored edge of the lamellae and the sometimes scanty,
sometimes very long setulae of the broom cells, also by the habitat on dicotyledoneous
leaves, and most easily, by the color of the fresh pileus.
ILLUSTRATION. Singer(1965), pl 10, fig 46.
83. Marasmiuspallipes Spegazzini, Anal. Soc. Ci. Argentina 16: 272. 1883.
TYPE. Balansa3379, from Paraguay.
Pileus cinnamon buff, ochraceous buff, light brown or clay color, at times slightly
zonate in the declivous portion, or graduallypaler towards the extreme margin,dried
between "gold br." and "Bombay M&P,"occasionally with somewhat more ochraceous
extreme margin("chipmunk") and deeper regions in center ("coffee"), without a white
or pallid zone around a black central dot or papilla, the central dot well differentiatedwit]
an extraordinarilysmall papilla in its center, in younger stages papilla often prominent
and deep chestnut to blackish chestnut, later slightly convex to flattened, sulcate except
in the central umbilicus containing the black dot, otherwise convex and glabrous,
2-8 mm broad. Lamellaewhite, edges and sides white, eventually or dried becoming
somewhat paler than the pileus, broad, distant, equal or with 1-2 lamellulaeor 1-2
forked ones when mature, 9-13 through-lamellae,collariate. Stipe paler than the
pileus, often reachinglight chestnut below and pale stramineousabove, dried usually
almost uniformly pale cinnamon or pale reddish fuscous, or sordid stramineous,
smooth and glabrous,shining, insititious to the substratumwith a very minute basal
disc, at apex often white in young specimens, setose-wiry,equal 18-40 X 0.1-0.7 mm;
rhizomorphsconcolorous with stipe or black, not regularlyobserved, accompaniedby
"telepodes," all glabrous,often branched,mostly scanty. Context thin, white, in-
odorous.
Spores (7.5-) 8.2-14 X (4-)4.5-6.2p, in some collections relatively small with the
most common size being 9.7 X 4.5k or 10-11 X 4-4.8 uellipsoid to ellipsoid-oblongor
even slightly subfusoid, smooth, hyaline, inamyloid, tending to germinateon the lam-
ellae. Hymenium: basidia(19-)23-32 X 6.8-8.10.5, , 4-spored;basidioles fusoid; cyst-
idia none: cheilocystidia like epicuticularbroom cells but predominantlyhyaline to
pale melleous with melleous to melleous golden setulae. Hyphae: trama of pileus and
hymenophoral tramaweakly pseudoamyloid;that of stipe somewhat more strongly but
not deeply or rapidly pseudoamyloid;all hyphae with clamp connections. Cortical
layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type,
main body 11-22 X 6.5-12,L, often hand-shapedor Clavariella-shaped, some slightly
thick-walled(0.6gi), brownish hyaline to pale melleous, some quite hyaline, setulae
apical and erect but sometimes slightly spreading-divergent-subapicular, 2-5.5 X ? 1.3j
134 Flora Neotropica
(at base), deep golden melleous, melleous, or brown, sometimes coarse, spinulose to
subcylindric,more often obtuse than subacute.
On bark,roots, woody monocotyledonous and dicotyledonous materialin forest
humus, dead twigs, mossy logs in tropical and subtropicalforest, densely gregariousto
cespitose.
MATERIALSTUDIED. BRAZIL.Santa Catharina:Sao Canisiode Porto Novo, 1928, J. Rick
642 (FH). PARAGUAY. Guarapf,Mar 1880, Balansa3379 (LPS), type. ARGENTINA. Salta: Orin,
3 km west from Rio Blanco bridge, 17 Mar 1955, Singer T 2302 (LIL);Misiones:Iguazu, Punto Canoa
29 Feb 1960, R. T. Guerrero118 (LIL). BOLIVIA. Beni: Vaca Diez, Ivon, 3 Apr 1956, SingerB
2480 (LIL).
This occurs also in Trinidadon Ficus benjaminasince materialdescribed and ill-
ustrated by Dennis (195 la) under the name of M. aciculaefbrimisB & C. appears to be
this.
Spegazzini apparentlydescribed the materialexclusively from dried material,and
it is obvious that he misdescribedthe colors.
The Salta and Bolivian materialdiffers from the other materialsin having smaller
spores; these are in the former only 9.7-10 X 4.8-5.4s, but some spores reach 11.2 X
5.8,u, in the latter 9.7 X 4.5 more rarely 8.2-1 1.7 X 4-4.8,u. It is this material that
has a distinct papilla when young, becomes graduallypaler colored (from chestnut to
ochraceous buff and light brown and eventually pallescent to pallid), maintaininga
deep chestnut central dot (umbo), and becoming between "gold br." and "Bombay"
M&Pwhen dried. The Brazilianmaterialbecame more gray-brownwhen dried and
showed less of a darkercentral umbo. It was also less pseudoamyloid. The mater-
ial from Misiones has very short and broad basidia(19-20 X 8-10.5,).
Whetherthese small differences are significant can be determined only after
the recollection of more material from more regions.
ILLUSTRATION. Singer(1965) pi 10, fig 43; Dennis (195 a) pl 20, fig 5.
84. MarasmiusaciculaeformisBerkeley & Curtis, Jour. Linn. Soc. Bot. 10: 297. 1868.
85. Marasmiuspolycladus Montagne,Ann. Sci. Nat. Bot. IV. 1: 113. 1854. Fig 54.
TYPE.Leprieur, from French Guyana.
Pileus brownish purple to blood red, brown when dried, glabrous,sulcate, hemispher-
ical to convex, with a strong papilla later in an umbilicus, 4-8 mm broad. Lamellaefuscou
(dried), with not discolorous edge, distant, broad, collariate. Stipe spadiceous-black,
short, about 0.25 mm broad, inserted in a sheet- to volva-likenode of the broad black,
shining, glabrousrhizomorphs.
Spores 6.3-7.7 X 2-2.5(-3) u, ellipsoid-oblong,smooth, hyaline, inamyloid. Hymen-
ium: basidioles fusoid; cystidia none; cheilocystidia like epicuticularbroom cells but
more often hyaline. Hyphae weakly pseudoamyloid, almost inamyloid in hymenophoral
trama and the interior of the rhizomorphs. Cortical layer: epicutis of pileus hymeniform
broom cells of the Siccus-type, main body 11-26 X 7-8c, clavate, with hyaline to mell-
eous thin to thick (1.5 ) wall, with melleous to reddishbrown setulae, these acicularto
rod-like,acute to obtuse, 4.5-6.5 X 1-2.
On and around dead branchesof trees, French Guyana.
MATERIAL STUDIED. FRENCH GUYANA.Cayenne,Leprieur(K),partof type.
1949, Dennis 339 (K) type. ECUADOR. Napo, Lago Agrio, 16 May 1973, SingerB 7512 (F).
BOLIVIA. Beni: Vaca Diez, 22 km east of Riberalta,near Arroyo Florida, 7 Apr 1956, SingerB
2565 (LIL). BRAZIL. Pernambuco:Camaragibe,7 Jul 1960, SinlgerB 3282 (BAFC);B 3250
(BAFC);6 Jul 1960, SingerB 3125 (BAFC).
There is a puzzling inconsistency in the reaction of the hyphae with the Melzer's
reagent. Collections B 3250 and B 7512 showed distinctly pseudoamyloid pileus-trama,
B 3282 only in the zone underneaththe epicutis. Marasmiuns martlae has slightly broad-
er spores and more lamellae;it is lignicolous.
ILLUSTRATION. Singer (1965) pl 9, fig 35; Dennis (1951a) pl 19, fig 4.
pigment as the setulae; setulae 1-3.2 X 0.5-1.2 , golden melleous to succineous, brown
in the Melzer'sreagent, not initially red; obtuse, sometimes forking;on the umbo of the
pileus there are some irregularlyclavate broom cells with thick walls turningvinaceous
in Melzer's,with few setulae or these replaced by rounded, lobulate protuberancesor
simply uneven above.
On dead tough leaves of dicotyledonous trees and shrubs. Puerto Rico.
comm. Johnston
MATERIALSTUDIED. PUERTORICO. San Juan, BrotherHioram7,
Nov 1912 (NY), type.
ILLUSTRATION. Dennis (1951c), fig 8.
The color of the pileus was annotated as red by Johnston who says that he has
collected the same (?) fungus also on Polypodialesand sugarcane, dicotyledonous
branches,etc. The following variety agrees with the type in almost all characters,
certainly in all important charactersseparatingthis species from the following related
species exceptingthe spore length.
like and mostly obtuse, more rarely attenuated upwards,sometimes forked, all or most
(with a few hyaline) pigmented melleous to rather deep warm brown but not red (not
even partly and initially); among these, particularlyin the center of the pileus some
thick-walledand more strongly pigmented(but not red) broom cells with variableshape
and fewer setulae, even no setulae at all, at the apex.
On dead stems and mostly leaves of dicotyledonous trees fallen to the ground but
also on monocotyledonous (e g musaceous) leaves. South America.
MATERIALSTUDIED. BRAZIL. Guapore,Guajaramirim, 10 Mar1956, SingerB 1820
(LIL). BOLIVIA. Pando: Madrede Dios. Las Piedras,5 Apr 1956, SingerB 2499 (LIL), type;
B 24995 (LTL);Beni: Vaca Diez, Guayaramer'n,8 Mar1956, SingerB 1704 (LIL);La Paz: Nor-
Yungas,CarmenPampa,26 Feb 1956, SingerB 1483 (LIL).
ILLUSTRATION.Singer(1965) pl 10, fig 40 (as AI.aripoensis).
ing insititiously directly from the substratum,in the first case attached to a disc-like
node or ring, 15-50 X 0.1-0.2 mm. Context white, thin, inodorous.
Spores 6.5-8.5 X 3.3-4.5 s, mostly 7-7.7 X 3.5-(4) ,, ellipsoid to ellipsoid-oblong,
smooth, hyaline, inamyloid. Hymenium:basidia 17-25.5 X 5.5-6 u, 4-spored;cystidia
none; cheilocystidia like the epicuticularbroom cells but mostly hyaline. Hyphae of
the pileus-tramahyaline, with clamp connections, with thin or ratherthin wall, dis-
tinctly pseudoamyloid;hyphae of the stipe parallel,pseudoamyloid. Corticallayer:
epicutis of the pileus hymeniform, broom cells of the Siccus-type, main body clavate
and sometimes above broadenedlaterally, more rarelyventricoseof ampullaceous9-16
X 3.5-7.5-(14) , with often slightly thickened wall (0.5-1 A), hyaline to cinnamon and
often red at the apex (but becoming melleous after a while in alkali medium), with
setulae 1.8-5.5 X 0.7-1.8 ,u, equal or attenuated to an obtuse tip, cinnamon to purple
or brownish purple (but becoming melleous in alkali), in some cells yellowish to hyaline.
On dead leaves of dicotyledonous trees (e g "Pau Ferro"). Brazil.
MATERIALSTUDIED. BOLIVIA. Beni, Vaca Diez, Riberalta,29 Mar1956, SingerB
2330 (LIL), type. BRAZIL. Pernambuco:Camaragibe,7 Jun 1960, SingerB 3251 (BAFC).
In both varieties(which differ mainly in the breadth of the spores), the number
of stipes attached to a rhizomorphsratherthan the substratumvaries from zero to
the majority but there are always many stipes directly attached to the substratum;in
some collections only the latter can be found. In view of the inconstancy of this
characterin this group, I now combine var subtrichorhizuswith var puttenansii.
ILLUSTRATION. Singer (1965), pl 10, fig 42.
specimens) and the small central spot or umbo which is brown, the umbo being red when
young and black or blackish when mature in fresh condition, sulcate over the convex
portion, otherwise smooth, glabrous,4-5.6 mm broad. Lamellaewhite or whitish, dis-
tant (9-11 through lamellae), ratherbroad with white or whitish edge and collarium.
Stipe sordid ochraceous brown (13 K 10 to "tortoise shell") to sordid umber, with pal-
lid apex when young, and with deeper colored base when old, in dried condition whole
stipe light brownish, pale cinnamon, or cinnamon, shining, smooth, and glabrous, 26-60
X 0.2-0.3 mm, directly on substratuminsititious; no rhizomorphs. Context extremely
thin, white, inodorous.
Spores 9-11.8 X 4.8-6.2 p, ellipsoid, more rarely ellipsoid oblong, or somewhat
clavate-ellipsoid,smooth, hyaline, inamyloid. Hymenium:basidia 19-19.5 X 6-6.5 ,:
cystidia none; cheilocystidia similarto epicuticularbroom cells, but often few and scat-
tered. Hyphae of pileus filamentous, hyaline, varying from inamyloid to distinctly
pseudoamyloid,with clamp connections. Corticallayers: epicutis of pileus consisting
of elements of the Siccus-type, main body hyaline, subhyalineor golden melleous, some-
times forked at tip and often hand-shaped,clavateto subcylindricif entire. 12-20 X 6-7.
5 P; setulae about 2.5-5.7 u long and 1.5 u broad at base, hyaline to golden, mostly golden
spinulose, sometimes forked at tip; intermixed with these broom cells there are sometimes
some smooth hyaline or golden yellow to melleous vesiculose bodies.
On fallen epiphytic Bromeliaceaein subtropicalforest.
MATERIAL STUDIED. ARGENTINA. Tucuman: Sierra de San Javier. Parque Aconquija at
800 m, Singer T 1457 (LIL), type.
ILLUSTRATION.Singer (1965) pi 10, fig 45.
00a. Marasmiusguyanensisvarguyanensis
TYPE. Leprieur,from French Guyana.
Pileus yellow buff, yellow ocher, chrome yellow or cadmium yellow, bronze yellow,
("nugget, bronze y.," "daffodil," Spanish y.," "deep chrome," "cadmiumy." M&P)when
dry (in situ) turning orange-goldbrown to ferruginous("Punljab,""Arabianbr.," "ferrug-
inous," "feuille morte" M&P),often collected in this shade when not quite freshly grown,
dried a color between "Punjab"and "feuille morte" or near "Arabianbr." M&Por yellow-
ish brown and sometimes if poorly preparedreachinga sordid brown color in the herb-
arium, fresh with a central black dot which in dried materialappearsas a deep brown to
blackish papilla or small umbo prominent in young materialbut later hidden in a small
umbilicus,without white zone around black dot, but sometimes in fresh condition with
a paler yellow ring around it, glabrous,sulcate outside umbilicus when mature, 1.2-6 mm,
mostly 2-3.5 mm broad. Lamellaewhite, in age whitish to cream, with the edges con-
colorous with sides, in dried materialeventually sometimes entirely subconcolorouswith
pileus, medium broad to broad, mostly broad, distant, 8-11, collariate. Stipe succineous
brown to umber, later black, at first entire stipe white (in buttons when quite fresh),
later only apex whitish to concolorous with pileus, and eventually apex concolorous
with lower portion of stipe, smooth and glabrous,rarelyminutely pruinate(?) at base,
insititious directly on the substratum, 10-40 X 0.1 mm, black rhizomorphsnot accom-
panying the carpophores.
Spores 8.5-14 X 2.5-4 ,, clavate or comma-shaped,smooth, thin-walled,eventually
often collapsing, hyaline, inamyloid. Hymenium: basidia 19-21 X 6-8 , 4-spored, cla-
vate, hyaline: basidioles fusoid; cystidia none; cheilocystidia, broom cells of the Siccus-
type, hyaline to yellowish hyaline, otherwise like the epicuticularbroom cells. Hyphae
of subhymenium,hymenophoraltramaand trama of pileus hyaline in ammonia, with
clamp connections, inamyloid in subhymenium,inamyloid to weakly pseudoamyloidin
hymenophoraltrama and more distinctly (though weakly) pseudoamyloidin pileus trama
accompaniedby some oleiferous hyphae; tramaof pileus a very thin layer, pseudoamy-
loidity noticeable only in accumulationsof hyphae; hyphae of stipe in strictly parallel
arrangement,with numerousclampedsepta, weakly pseudoamyloid. Corticallayers:epic-
utis of pileus hymeniform, consisting of broom cells generally of the Siccus-type but
some with slightly divergentsetulae which descend somewhat from the extreme tip of
the cell and thus vaguely remindone of some epicuticularelements of the Rotalis-
type, clavate, or vesiculose-ventricoseand always broadly rounded, even applanate,often
forked or cauliflower-likedivided in upper portion, 10-25 X 4.8-17 p, either entirely
hyaline or hyaline in lower portion or entirely brownishyellow or melleous to deep
orange, generally thin-walledbut often some thick-walled,setulae 1.5-4 X 0.7-1.5 s,
i e ratheithin at base, rathercoarse,acute, subacuteor obtuse, simple,varyingfrom
hyaline to light yellow to deep golden yellow or at times even brownish yellow. Cells
of the black-dot-regionof the center similarbut either without setulae or these replaced
by broadenedrounded conidia-likeappendages,chestnut brown and thick-walled,form-
ing a palisaderatherthan a hymeniform layer. Stipe surface without a coveringlayer.
Marasmius 147
concolorous, pliciform, few, ?collariate. Stipe dark, somewhat shining, erect, glabrous,
10-20 X 0.15-0.18 mm risingfrom thicker rhizomorphswhich are creeping and form a
very extended (200-500 mm), very loose "rhizomorphoidmycelium," which forms
annularjoints where the stipes are inserted (description adapted from Spagazzini, 1883).
Microscopicalcharacterssee below.
On bark of rotting logs in virgin forest Caa-Guazfi,Paraguay.
Spegazzinihimself says that his materialconsisted of old, somewhat obliterated
specimens which should be recollected and restudied. He indicated later that this species
was recollected in Paraguay(Fiebrig, February 1920 on trunks of Pithecollobium scalare)
but reduced his species to synonymy with Marasmiusequicrinis(M. crinis-equiMueller
ex Kalchbrenner);this species is strongly papillate. Paraguayanmaterial(Piragu,July
1879) is conservedat LPS and does indeeed show a papilla but no sporescould be re-
covered by me or by Dennis (1951a); but Spegazzinifound them to be 7-8 X 4-5 p. It
is indeed probable that the Paraguayancollections refer to Marasmiuscrinisequi. There
is however a further collection determinedMarasmiustrichorhizusby Spegazzini. AR-
GENTINA. Formosa: Dec 1900, Kermes 723 (LPS). This materialhas spores 8.2-9
X 2.2-3.5,, thus narrowerthan indicated by Spegazzini. It has epicuticularbroom cells
of the Siccus-type, 12.3-19.2 X 8.2-11 p, setulae 2.5-3.7 p long, main body pale yellow-
ish, setulae brown. Hyphae of the pileus inamyloid,with clamp connections. The color
of the pileus is now brown but its originalcolor is not indicated.
There are two possibilities. Either the original description is taken at face value
and the later Paraguayancollection only is taken as identical with the type; in this
caseMarasmiustrichorhizuswould become a (lignicolous?)form of Marasmiuscrinis-
equi Or else the Formosanmaterialis taken as authentic and Spegazzini'sfailureto
indicate the presence of a prominent papilla and normal(not fold-like) lamellae is con-
sidered erroneous or exceptional. In this case Marasmiustrichorhizuswould be a good
species, not fully and partially wrongly described. A decision as to which approachto
take, should, I believe, be postponed until an attempt has been made to recollect the
species at the type locality, inasmuchas Spegazzini'sspore measurementssuggest that
the type might have been close to our no. 80 (M. schultesii) if the hairinessof the
rhizomorphshas disappearedor was overlooked.
walled cells, all with apical setulae 2-5.5 X 1-1.5 i, these spinulose-obtuse,often some-
what curved, hyaline to more frequently golden melleous.
On grassculms, both living and dead, also on grass roots and leaves, solitary or in
small groups. Known hosts: Triticuni,Agroprymum, Cynodon, Bambuseae,possibly also
on small twigs of other Monocotyledones.
MATERIAL STUDIED. TRINIDAD. St. Joseph, 30 Nov 1949, Dennis 411 (K), type of var
brevisporus; St. Augustin, 16 Jun 1949, Dennis 101 B (K); 101 A (K). MEXICO. Chiapas, between
Finca Sospiro and El Pozo, 4 Aug 1969, Singer M 8956 (F). HONDURAS. Atlantida: Telas, 14 Dec
1927, 15 Mar 1928, P. Standley, det. V. K. Charles (BPI). ARGENTINA. Catamarca: Suncho, Santa
Rosa, 20 Jan 1952, Singer T 1787 (LIL); Rio Potreros at 1400 m alt, 10 Feb 1952, Singer T 1816
(LIL). U.S.A. New England, Sprague (FH), type of M. pniinatus; Florida: Highlands County, High-
lands Hammock State Park, 25 Aug 1942, Singer F 395 (F); Illinois: Prairie W of Highlands Park,
9 Jul 1973, Singer N 4670 (F); Abington, 12 Jul 1924, Young (BPI) type of M. tritici, Minnesota:
Ramsay County, St. Paul, 20 Jul 1928, Christensen (BPI). U.S.S.R. Leningrad, 19-26 Aug 1938,
Singer (LE). FRANCE. Mesnil-Val, Aug 1931, Madamiefi FPraniois (LIL). BELGIUM. Ardennes,
Libert 119 (BPI, LE), type.
The spore size alone cannot serve to- distinguishvarietiesof taxonomic significance
since in the majority of the collections both types of spores are present, even in the
type of var brevisporus Dennis we find a small number of spores up to 11.5 , long.
ILLUSTRATION. Singer (1958a), fig 50.
This variety differs from the type variety in less or not pseudoamyloidpileus-
trama, slightly largerspores and the more rusty color.
ILLUSTRATION. Singer (1958a), fig 51.
the stipe parallel, pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform,
broom cells of the Siccus-type, main body 8-29 X 4-12 p, clavate or cylindrical, with
ratherthin wall, golden melleous to (few) subhyaline, with apical coarse setulae 3-8
X 1-2 u, deeper colored than main body, acute or obtuse.
On both mono-cotyledonous and dicotyledonous hosts (mostly dead dicotyledon-
ous wood) but also on living roots etc, sometimes among mosses, gregariousin subtrop-
ical and tropical forests, from Florida to Ecuador. Known hosts: Coccolobis, Carya,
Quercus,Ficus, Sabal.
MATERIAL STUDIED. U.S.A. Florida: Dade County, Matheson Hammock, 3 Nov 1942.
Singer F 1456 (FH) type, and various other collections from Florida (F, FH). MEXICO. Veracruz:
Estacion Biologica de las Tuxtlas, 19 Jun 1969, Singer M 8020 (F); M 8043 (F). ECUADOR. Napo:
Lago Agrio, 16 May 1973, Singer B 7506 (F).
11Oa.Marasmiuspallidocinctusvar pallidocinctus
TYPE.Dumont et al VE 701, from Venezuela.
Pileo ferrugineo-aurantiaco,zona pallida circum maculum centralem fumoso-ful-
igineum ornato, 3-6 mm lato; lamellis albis, subdistantibus,collariatis, stipite atrocast-
aneo; rhizomorphisnullis. Sporis 9-10.5 X 3.5-4.3 1; hyphis pilei debiliterlentoque
pseudoamyloideis,fibulatis;elementis epicuticularibustypi Marasmiisicci. Ad folia
dicotyledonea delapsa in Venezuela. Typus in NY conservatusest.
Pileus (dried) rusty orange with a pallid zone around a central fumosous-fuliginous
spot, glabrous,sulcate-plicate,convex, deeply depressedin the center with a low papilla
becoming visible in dried material, 3-6 mm broad. Lamellaewhite with white edge and
collarium, subdistant(about 12-23 through-lamellae),broad, collariate. Stipe deep chest-
nut, glabrous,equal, insititious on the substratum,not rising from nor accompaniedby
black rhizomorphs,long: 75-80 X 0.3 mm; "telepodes" often formed. Context whitish,
thin, inodorous.
Spores 9-10.5 X 3.5-4.3 p, ellipsoid-oblong,smooth, hyaline, inamyloid. Hymenium:
basidia4-spored;basidioles fusoid; cystidia none; cheilocystidia like the epicuticularbroom
cells but all hyaline. Hyphae of the pileus-tramayellowish-hyalineto hyaline, weakly and
slowly but undoubtedly pseudoamyloid,with clamp connections, filamentous, 4-5 u broad,
thin-walled,not gelatinized;hyphae of the stipe parallelpigmented, pseudoamyloid. Cor-
tical layer: epicutis of pileus hymeniform, somewhat mottled because of alternatinglypaler
and darkerbroom cells, these of the Siccus-type, main body 11-15 X 4.5-6 u, thin-walled,
yellowish hyaline, with setulae 2-5 X 0.5-1.5 ,, acute or obtuse, simple or forked, yellow-
ish hyaline to golden ochraceous.
On dicotyledonous leaves fallen to the ground. Venezuela.
MATERIAL STUDIED.VENEZUELA. Dto Federal, Mt. Naiguata, above La Planta, 25 Jun
1971. Dumont et al VE 701 (NY).
The different size of the spores seems to be the only important difference between
this and the type variety.
smius inaequalis) strongly pseudoamyloidin pileus and stipe; stipe centrally attached,
growingfrom leaves or wood etc, always with basal mycelium; black rhizomorphsabsent.
113. MarasmiusinaequalisBerkeley & Curtis, Jour. Linn. Soc. Bot. 10: 297. 1869.
TYPE. WVright, from Cuba.
Pileus white, center brown ("cookie" M&P),dried entirely like illarasmiutshelvolus
in color, glabrous,sulcate, campanulateto convex, 2-5 mm broad. Lamellaewhite with
white edge, distant or subdistant(10-12 through-lamellae),rathernarrow,subconcolorous
with the pileus when dried, narrowlyadnexed to free, definitely not collariate. Stipe
chestnut brown with pallid or white apex, dried chestnut-amber,smooth, glabrous,shin-
ing, 4-32 X 0.1-0.2 mm; basal mycelium tomentose or strigose, white, accompaniedby
whitish mycelial patches and rhizomorphsin many populations. Context very thin, in-
odorous.
Spores 11-15 X(2-)3-3.3 ,, guttiform-subclavateto cylindric-subfusoid,often miore
or less curved or at least the hilar appendagestrongly curved inwards,smooth, hyaline,
inamyloid. Hymenium: basidia 20-21 X 5.5-6.5 p, 4-spored;cystidia none; cheilocystidia
like the epicuticularbroom cells but here all thin-walled,hyaline to melleous. Hyphae
of the trama of the pileus with clamp connections, inamyloid. Cortical layers: epicutis
of the pileus hymeniform, broom cells of the Siccus-type, of two kinds, (1) thin-walled,
hyaline to melleous, sometimes all hyaline, main body 8.5-17 X 5.5-10u, setulae acute
or subacute, 3.3-4.5 X + 1 ,, (2) as above but thick-walled,wall up to 1.7 p thick.
On rotting dicotyledonous wood (sticks, chips, roots, etc.), gregarious.
MATERIALSTUDIED.CUBA. Santiagode Cuba, 1857. Wright114.4 (FH), authentic.
BOLIVIA. La Paz, Nor-Yungas,Coroico, 30 Jan 1956, SingerB 736 (LIL).
ILLUSTRATION.
Singer(1958a) fig 18.
Key II
1. Spores 11-16.2(-17) X 2.7-6.3u.
2. Lamellae subdistant to distant.
3. Dried pileus "cocoa," "leather or" M&P often radially striped in different shades.
of brown; fresh pileus not orange or orange-ferrugineous.
4. Spores 11-15.2 X 2.8-4u; pileus 9-31 mm broad; cn woody twigs and vines and
on dicotyledonous leaves; cystidia usually present but often few or in-
conspicuous. 174. Mi.helholus.
4. Spores 12-16 X 2.5-3, (i e relatively narrower); pileus 27-42 mm broad; on
wood in Pacific rain forest. 130. M. helioloides.
3. Dried pileus differently colored, e g "Ginger," "persimmon," tending to "Alamo,"
"terracotta," "ferruginous," "caldera" in the center, young pilei often entirely
"raw sienna" i e more brightly ochraceous brown, brown-red or orange-rust when
dried, fresh pileus mostly orange or ferruginous or bay to ferruginous.
5. Lamellae with discolorous edges; spores 12-13.8 X 3.3-3.7,u; pileus 4-9 mmi
broad, orange, rusty orange; mostly growing on grass leaves, more rarely passing
over unto Populus or Taxodtium leaves. Southern subtropics. 131. Al. graminizcola.
5. Lamellae discolorous only in a narrow intermittent line (under a lens. \when
dried) or concolorous with the sides of the lamellae; pileus as small as above or
larger; spores as above or smaller or larger; not with the above habitat and
more widely distributed; not combining the characters indicated above.
6. Spores only up to 13 long; lamellae only medium distant to subdistant;
pileus small to medium (up to 20 mm broad); species of the northern part
of South America and reaching Mexico and the whole Caribbean area (see
"19" below).
6. Spores, at least many of them, reaching more than 12.5 p in length, often
up to 16(-17)p long. Widely distributed all over the tropics and subtropics
of the Americas. 126. M. berteroi.
2. Lamellae close or subclose.
7. Edge of lamellae distinctly and constantly discolorous; spores 11-15 X 3-4 g pileus
6-9 mm broad; lamellae subclose. On leaves of Dicotyledones. 129. M. neglectus.
7. Edge of lamellae concolorous with the sides or, if slightly discolorous, with dif-
ferent spores or not combining the above characters.
8. Spores 12.5-16 X (2.5-)3-4u; pileus on margin subsmooth or sulcate, in the
center smooth or convoluted; stipe covered with subhyaline, clavate, entire
dermatocystidia, or without any covering.
Marasmius 163
mycelium 33 X 1.5 mm, base reaching0.5 mm, basal mycelium tomentose, brown,
with alternatinghyaline fibers even in fresh material. Context very thin, in cortex of
stipe almost purple, then blackening,fragile, inodorous.
Spores 10.3-15 X 3.3-4.8A, mostly about 11.5-13.7 X 3.5-4,, with strongly
attenuatedhilar portion which is either straightor curved towards the inside or out-
side and then assuminga sickle or tilda shape, otherwise ventricose, hyaline, smooth,
inamyloid. Hymenium:basidia 25-30 X 5.5-7,, clavate, 4-spored;basidioles fusoid;
cystidia none or few and then not differentiated(cystidioles), 30-44 X 7.5-8.5jA,
neither metachromaticnor opaque. Hyphae pseudoamyloidwith clamp connections.
Corticallayers: epicutis of the pileus hymeniform, consisting of broom cells which are
hyaline to reddishbrown or melleous, 17.5-20.5 X 7-7.5., mottled in scalp view, set-
ulae 5.5-7.5Mlong obtuse.
On dicotyledonous, dead, fallen twigs and branchletsof the humid forest in the
subtropicalzone, solitary or in small groups in the shade.
MATERIALSTUDIED. ARGENTINA.Tucuman,Taff: ParqueAconquija,13 Mar 1951,
Singer T 1460 (LIL) type; T 1477 (LIL) paratype(cf Singer1965, p 258, footnote!).
narrow, free. Stipe deep chestnut below and slightly paler above, glabrous,smooth,
shining, equal, setose, 30 X 0.5 mm; basal mycelium white, not extensive. Context
very thin, inodorous.
Spores 13.8-18 X 3.5-4.8p, hyaline, smooth. Hymenium: cystidia, none seen;
cheilocystidia like epicuticularbroom cells; hyphae hyaline, psuedoamyloid. Cortical
layer: epicutis of pileus hymeniform, consisting of broom cells, 15-16.5 X 7-8.5 u,
sometimes spreadingbecause of divergenceof setulae to a diameterof 9-11 p, setulae
5.5-8.5 X 1.2-1.8u (at base), golden succineous, few hyaline, obtuse orsubacute, spin-
ulose or rod-shaped.
On dead dicotyledonous wood in subxerophytic woods.
MATERIAL STUDIED. ARGENTINA: Tucumnan:Burruyacuf, 5 kim south of town, 19 Feb
1957, Singer T 2995 (LIL) holotype.
It is possible that an additional collection (Argentina:Santiago del Estero, El Sal-
vador, July 1962, D. Foguelman, BAFC) also belongs here but this is only 15 mm high
and 1.5 mm broad and may be a nanism of M. berteroi.
Illustration. Singer (1965) pl 14, fig 76.
type of elements evenly distributedamong the other paler broom cells and giving
the scalp preparationsa mottled appearance),setulae hyaline to golden melleous,
3.5-3.8 X 0.7-1.5 (at base),, subacute to acute.
On forest litter adheringto mixtures of sticks and fallen leaves of various tree
species rotting in the woods and forests, singly or in closely joined pairs.
MATERIALSTUDIED. PUERTORICO. "Agaricusfuillus Bertero,"Bertero
(HerbariumPersoon (L) Type. BOLIVIA. La Paz: Nor-Yungas,Charobambaat 1200-1300 m
alt, 30 Jan 1956, SingerB 725 (LIL). ARGENTINA. Salta: Oran, 3.5 km west of Rib Blanco
bridge, 17 Mar 1955, Singer T 2310 (LIL);Tucuman:CiudadUniversitaria.1000 m alt, 26 May
1957, Singer T 3080 (LIL);Taficillo, 1500 m alt, 23 Feb 1955, Singer T 2219 (LIL);Yerba
Buena,600 m alt, 12 Feb 1955, Singer T 2159 (BAFC).
This variety differs from the following variety in habit and size. Mlarasnmius
flavofuscus Berkeley & Curtis from the Bonin Islandsis close. The type (FH) differs
mainly in the color of the pileus as described.
ILLUSTRATION. Singer (1965) p 14, fig 74.
126b.Marasmiusberteroi var major Singer, Sydowia 18: 334. 1965 (ex errorep 249
"var macrus").
TYPE. Singer T 2886, from Argentina.
Pileus bright orange fulvous, orange, orange red, rufous ferruginousor intensely
ferruginous(between "feuille morte" and "Eldorado"or between "feuille morte" and
"Peruvianbr.", "Agate," "Kobe", "henna",4-A-i2, towards marginsometimes "sun-
burst";"Ponce de Leon," ll-L-10, or "Saratoga"(M&P)on center and ribs, sometimes
reaching"Maracaibo"or deeperand richer than "caldera";color often reachingtones
deeper and richer than those of Maerz& Paul (such as Villalobos SO, 12 , 8), dried
similarto the fresh color, or "gold pheasant"or 12-F-9, glabrous,not viscid, at first
over half, soon over 4/5 to 5/6 of radius,sulcate to deeply furrowed, otherwise (on
disc) either smooth or rugose-venose,campanulateor conic-truncatewith flattened
disc, later convex and often umbonate, eventually convex-applanateor with uplifted
marginand tending to become obtuse, frequently even with depressedcenter, 10-56
mm broad. Lamellaecream, pale cream, cream whitish, yellow white, milk white,
often becoming pale orangeyellow when dried, usually more yellow if older, the
edges usually concolorous, rarely few lamellaemostly not intervenose,narrowlydis-
colorous, more rarely sparselyto densely intervenose,all entire or with few lamell-
ulae (through-lamellae9-19; lamellulae0-18), ventricose or not, narrowto broad,
3-12 mm broad, sometimes not quite reachingthe margin,narrowlyadnexed to
free, not collariate, spore print pure white. - Stipe reddishbrown to chestnut be-
low shading deeper and deeper towards the base, less deep and sometimes almost
umber further up (below "mohawk," "Mandalay,""Hindu," above "broncho"),
apex generally concolorous with lamellae (whitish to cream) but sometimes yellow,
especially when young, ("lemon yellow," 10-I/L-2/3 M&P),base often almost black,
glabrous,smooth, shining or opaque, at first taperingupwardslater sometimes equal,
narrowlyhollow (with a thin, rigid cortex and a tubulose cavity) 30-80 C 1-3 mm,
basal mycelium well developed, tomentose, white, more rarely buffy white.
Context white, in youth often lemon yellow in the apex of the stipe, rather
tough, even hard in stipe when old, inodorous.
Spores (8-)9.5-15.3 X 2.7-4 p, fusoid-ellipsoid, hyaline, smooth, inamyloid.
Hymenium: basidia 20-30 X 5.5-7 ,, clavate, 4-spored, hyaline, some opaque;
basidioles fusoid; cystidia (if present) not morphologicallyor chemically differen-
tiated from the basidioles;cheilocystidia like the epicuticularbroom cells but some-
Marasmius 173
times rather scattered and rare, with obtuse, sometimes long, erect setulae, more
rarely rathernumerousbut not making the edge heteromerous(intermixed with
basidia and basidioles). Hyphae, all thin-walledin pileus, everywherepseudoamy-
loid and with clamp connections, in old specimensmany hyphae, especially in stipe,
with thick walls. Cortical layers: epicutis of the pileus hymeniform, consisting of
broom cells, these 23-26 X 6-8.2 1,, of two types, (1) with hyaline and thin-walled
main body and hyaline to melleous setulae, (2) with the main body melleous to
bright ochraceousferruginousat least in upper portion which also may be thick-
walled; setulae apical and erect, often ratherlong, especially in type 2 some rather
coarse, e g about 1.7 , thick below, but in type 1 usually much more slender,pro-
jecting 5-11 .; the two types regularlydistributedand scalp appearancemottled.
On branchesand trunks fallen to the ground(dicotyledonous trees) and on leaf-
mold of dicotyledons, monocotyledons, conifers, in montane, subtropical,tropical-mon-
tane forest, solitary or gregariousbut scarcelycespitose (Podocarpus,Gramineaespp,
Alnus, Phoebe, etc).
MATERIALSTUDIED.BELIZE. El Cayo, Valentin, 28 Jul 1936, E. B. Mains3660 (MICH).
COLOMBIA.Valle: Buenaventura,Calima,22 Apr 1968, SingerB 6295 (F), a large-sporedform
see annotationbelow. ECUADOR. Napo: Lago Agrio, 17 May 1973, SingerB 7518 (F); Tungurah-
ua, Rio Topo, 30 Apr 1973, SingerB 7225 (F). BOLIVIA. La Paz: Nor-Yungas,Rio suapi, 1250-
1300 in alt, 9 Feb 1956, SingerB 1043 (LIL);Rio Yariza,23 Feb 1956, SingerB 1394 (LIL);Char-
obamba,30 Jan 1956, Singer B 693 (LIL). PERU. Cusco, Paucartambo,Atalaya,Carbon,700 m
alt, Vargas13519 (LIL). BRAZIL. SantaCatarina:Porto Novo, 1928, J. Rick (FH). ARGENTINA.
Tucuman:Anta Muerta,1100 m alt, 10 Apr 1949, Singer& Digilio T 418 (LIL);La Angostura,1800
m alt, 1 May 1949, Singer T 540 (LIL);Taficillo, 1700 m alt, 4 Mar 1951, Singer T 1368 (LIL);
Rio de los Sosas, 900 m, 4 Feb 1955, Singer T 2126 (LIL);CiudadUniversitaria,1100 m alt, 14
Jan 1957, Singer T 2886 (LIL) type; T 2887 (LIL);23 Jan 1957, T 2879 (LIL);26 Jan 1957, T
2913 (LIL); 1 May 1957, P. Wygodzinsky,det. R. SingerT 3073 (LIL);Misiones:Iguazu,Ref. "Yag-
uarete ," 29 Feb 1960, R. T. Guerrero133 (LIL).
The collection from Colombia (B 6295) agreesmacroscopicallywith var major but
may key out with Marasmiusrhabarberinusrather than M. berteroi because the spores
were 12-17 X 3 p, thus largerthan usual in M. berteroi var major. Since this collection
differs in severalaspects includingthe habitat on wood ratherthan dead leaves, from M.
rhabarberinus,it has been considered a large-sporedform of M. berteroi but it may re-
quire a special subspecific taxon.
The collection from Ecuador(B 7518) agreeswith var major except for the dis-
colorous or partly discolorous(concolorous with pileus) edge of the lamellae and may
also be a variety in its own right or subvarietyof var major.
ILLUSTRATION. Singer(1965) pl 14, fig 73.
to subfree. Stipe dark brown, subglabrous,under a strong lens very finely pubescent
or subpruinose,smooth, setose-equal,up to 25 mm long, 0.3-0.6 mm broad; basal
mycelium strigose, tending to be or to become fulvous. Context thin, white, inodor-
ous.
Spores 12.5-16 X (2.5-)3-4 u, fusoid, on apex and base sometimes slightly re-
curved towards the same side, smooth, hyaline, thin-walled,with severalsmall oil
droplets,inamyloid. Hymenium: basidia4-spored;basidioles fusoid; cystidia none;
cheilocystidia like the epicuticularbroom cells but subhyaline throughout. Hyphae
of pileus tramahyaline, of stipe brown, parallel, thick-walled. Corticallayers: epic-
utis of pileus consisting of broom cells with a main body 10-12 X 5.5-7 p, thin-walled,
brownish to hyaline, claviform;setulae 4.5-12 X 1-1.8 p, pale golden-ferruginous,acute
or subacute, spinose-conical;covering of the stipe consisting of widely spaced (long
stretches of surface layer without any covering), dermatocystidioidsmall hairs which
are 9-27 X 3.8-7.2 j, hyaline or subhyaline,claviform,more rarelycylindrical,with
thin to more rarely thick wall, entire or occasionally with one short inconspicuous
apical apiculus.
On dead wood (wet rotting logs).
MATERIAL STUDIED. BRAZIL. Amazonas: Municipalidad Manaus, Itacoatiara road in
virgin "terra firme forest," 30 Jan 1963, G. & L. T. Eiten & Gil M. Felipe 5254 (F), type.
This is close to M. matrisdeiwhich differs, especially with regardto the type of
coveringon the stipe.
few elongated to 18 plength, total measurementsof broom cells of pileus 24-31 X 7.5-9
p; surface of stipe with numerousbroom cells but the main body often strongly reduced
and often nothing but setulae (sometimes elongated up to 23 p and then setiform!) visible.
On fallen stems and branchletsof both mono- and dicotyledonous plants in trop-
ical rain forest.
MATERIAL STUDIED. BOLIVIA. Pando: Madre de Dios, Las Piedras, 5 Apr 1956, Singer
B 2523 (LIL), type.
ILLUSTRATION. Singer (1965) p 14, fig 80.
133. MarasmiusproletariusBerkeley & Curtis, Jour. Linn. Soc. Bot. 10: 296. 1869.
Fig 64.
TYPE. Wright, from Cuba.
Pileus white when fresh but soon becoming cream white to pale cream, on drying
gilvous to pale ochraceouscinnamon in the center and gilvous-whiteon the margin,gla-
brous, with sulcate margin,dull convex, eventually applanate,2-5.5 mm broad. Lam-
ellae white, eventually cream, dried gilvous-whitish,with edges concolorous with the
sides, distant to medium distant, tridymous, with about 12 through-lamellae and many
lamellulae,not intervenose, rathernarrow,to broad, adnate, more rarely adnexed.
Stipe pallid-white,mostly avellaneousto chestnut from the base upwards,pallescent
on drying if chestnut and becoming pale stramineous or pale bister in the herbarium,
glabrous to very finely pruinate (under a lens) smooth, equal, hollow-tubulose, 6-18 X
0.2-0.3 mm; basal mycelium radiating,white, cottony-tomentose and sometimes some-
what ascendant on the stipe; white rhizomorphsoften present, tomentose. Context
white, very thin, inodorous.
178 Flora Neotropica
135. Marasmiustrinitatis Dennis, Trans. Brit. Mycol. Soc. 34: 425. 1951.
Marasmius 179
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.
25-54 X 1.5-2 mm; basal mycelium white. Context inodorous, extremely thin in pil-
eus.
Spores 7.5-8.2 X 3.5 p, ellipsoid-oblong,hyaline, smooth. Hymenium: basidia4-
spored;cystidia none: cheilocystidia like epicuticularbroom cells, but all hyaline. Hy-
phae of pileus and hymenophoraltrama strongly pseudoamyloid. Cortical layers: epic-
utis of pileus hymeniform, consisting of broom cells of the Siccus-type, main body
hyaline, rarely concolorous with the setulae, 10-14 X 7p, setulae golden yellow, 4.8-
5.5 X 1 p.
On dead (mainly dicotyledonous) leaves in rain forest.
MATERIAL STUDIED.BOLIVIA.Beni:VacaDiez,ArroyoFlorida,22 kmwestof Rib-
eralta,7 Apr 1956, SingerB 3047 (LIL), type.
This is intermediatebetween M. trinitatis var trinitatis (from which it differs in
the smallerspores and subdistantlamellae) and M. digilioi (which differs in its smaller
carpophoresand closer lamellae).
ILLUSTRATION. Singer (1965) pl 15, fig 87.
138. Marasmiusdigilioi Singer in Singer & Digilio, Lilloa 25: 201. 1952.
TYPE. Digilio & Singer T 488, from Argentina.
Pileus olive gray, olive brown, olivaceous-spadiceous("buffalo" "olive brown,"
14-J-6, 15-J-7/8, 15-L-8/9 M&P)sometimes with "cattail" or "biskra"in center), gla-
brous, smooth to distantly furrowed-subsmooth,eventually often becoming long-sulcate,
otherwise smooth, or more rarely rugulose in the center, often undulate, with a slight
depressionor/and a papilla in the center, otherwise convex, but eventually applanate
to irregular,5-16 mm broad. Lamellae(olive-brown)whitish or grayishpallid when
young, later between pure white and "rubber,"with the edges concolorouswith sides or
more rarely with pileus, subclose to medium close, narrow to broad, ventricose or not,
rounded-freeto rounded adnexed, Spore print pure white. Stipe rusty ochraceousbrown
to deep chestnut below and graduallypaler above with white apex ("Java," "alamo,"
"chutney"), glabrous, smooth, shining, filiform hollow, 15-40 X 0.4-1.5 mm; basal
mycelium tomentose or strigose, fresh usually white, dried often pale fulvous. Context
thin, white in pileus, inodorous.
Spores 6-9 X (3-)3.5-4.5(-5),, ellipsoid with suprahilardepression,hyaline, smooth.
Hymenium: basidia 20-28 X 6.8-9,u, clavate, 4-spored;basidioles fusoid; cystidia none or
cystidiole-like near edge; cheilocystidia either all like the epicuticularbroom cells or dif-
fering in being all hyaline, or else these broom cells intermixed with ampullaceousto ven-
tricose elements which have sometimes a slight knoblike swelling at the tip, at times
branchedbut not setulae-bearing,20-27.5 X 5.8-6.5 p, i e cystidiole-likeand in this
form also occasionally occurringnear the edge. Hyphae with clamp connections, pseu-
doamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of
the Siccus-type, usually mottled in scalp view (hyaline-melleous)broom cells 14-30 X
5.8-11.7 u, main body hyaline, cylindric, clavate or vesiculous setulae erect, acute, some-
times relatively long, most melleous to bright golden melleous but on some cells all
hyaline e g 5-8 p long.
On rotting wood, more rarely on dead foliage mixed with other detritus in mon-
tane and subtropicalmontane, also subxerophyticpremontaneand tropical montane
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-
182 FloraNeotropica
cylindricalor ventricose, 9-19 X 4-12 g, setulae 2-16 X 0.7-2 u, conical, acute or sub-
acute, palest golden to golden hyaline or quite hyaline, or with a more conspicuous in-
traparietalmelleous golden pigment.
On dicotyledonous leaves fallen to the ground as well as on nearby dicotyledonous
wood, gregarious.
MATERIAL STUDIED. MEXICO. Veracruz, 3 km west of Fortin de las Flores, 24 Jan 1969,
Singer M 8188 (F), type. ECUADOR. Napo, Shushufindi, 13 May 1973, Singer B 7414 (F).
This variety approachesM1.iiapoesis because of its broader spores but differs in
havingthe colors of M. leoninus, glabrousstipe and non-interwovenlamellae.
146. Marasmiusfloriceps Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 298. 1869.
Fig 69.
TYPE. Wright31, from Cuba.
Pileus bright orange-rusty,bright orange or bright red-brown,sometimes orange
with some inconspicuousbrown radial stripes and/or orange brown or orange yellow
marginalband (e g "terracotta"with "Nasturtium"center or "toltec" with center 11
C 10 to 12 B 10 or "doubloon" with "Punjab"center or "golden wheat" with "rose
amber" center, dried e g 3 A 12, the orangebrown becoming "copper br" M&P),
glabrous,sulcate excepting the disc, rarely short-sulcatewhen mature with, more rarely
without rugose or venose center, campanulateto convex or subhemispherical,later
often with either more declivous or upturned margin,the center often papillate but
also with umbilicate center, 9-20 mm broad. Lamellaewhite to orange-cream,subclose
to subdistant, the edges not discolorous or only near margin pale orange when dried,
narrowto ratherbroad, at first somewhat ascendant, then horizontal or almost so,
free to rounded-adnexed,intermixed with many lamellulae. Stipe brown to deep chest-
nut with white to pale cream apex at least at first, glabrous,equal or subequal, hollow,
23-58 X 0.5-1 mm; basal mycelium scanty to more often abundantand strigose. white
or fulvous to tawny when fresh and dried. Context of the pileus white, very thin,
inodorous.
Spores(8-)8.5-13 X (2.7-)3-4.2(-4.8),, fusoid to fusoid-oblongbut some some-
times relatively broad and often curved in lower portion if narrow, smooth, hyaline, in-
amyloid. Hymenium: basidia 16-24 X 6-8 ,, 4-spored, basidioles fusoid; cystidia none
except for a few inconspicuoussubulate and hyaline ones at the gill edges among the
cheilocystidia which are broom cells like the epicuticularbroom cells but usually all
hyaline or with golden hyaline or orange-hyalinesetulae. Hyphae of the trama of the
pileus hyaline with firm but not distinctly thickened wall although often slightly swell-
ing in alkali but not dimorphic, filamentous, not gelatinized, with clamp connections,
but often many secondarysepta present, pseudoamyloid,those of the stipe parallel,+
thickwalled, deep yellow. Subhymeniumcellular. Hymenophoraltrama regular,con-
sisting of filamentous but sometimes relatively broad, thin-walledelements. Cortical
layers: epicutis of the pileus hymeniform, consisting of broom cells, their main body
9-13 X 8-11 u, hyaline to ochraceous, scalp preparationsappearingmottled because the
setulae are either hyaline or golden to deep golden melleous, acute to subacute, (2-)3-10
M,long; covering of the stipe none, except for scarce, minute excrescencieslooking like
setulae without main body.
On fallen dicotyledonous leaves, stems and twigs of Leguminosae,rotten wood
of all kinds, but also on small monocotyledonous stems.
MATERIAL STUDIED. CUBA. Wright 31 (K, FH), type. MEXICO. Veracruz: Estaci6n
Biologica de las Tuxtlas, 7 km S of Montepfo, 21 Jun 1969, Singer M 8116 (F); 29 Jul 1969,
Marasmius 191
SingerM 8741 (F); 20 Jun 1969, SingerM 8088 (F). COLOMBIA.Valle, MunicipalidadCali,
Rio Prance,1 May 1968, SingerB 6511 (F); BuenaVentura,Calima,18 Apr 1968, SingerB 6206
(F).
This species is often difficult to distinguishfrom M. berteroi var berteroi and
I. corrugatusvar auranltiacusbut the charactersemphasizedin the key should make
it possible to come to an identification in all cases if the distributionof the three
species is taken into considerationand the cortical layers of M. corrugatusare duly
investigated.
ILLUSTRATION. Dennis (1951b), fig 7 (type).
tively small number of broom cells among the setula-lesscells in the epicutis and gill
edge, in the strongly dimorphic characterof the hyphae of the hymenophoraltrama
and in somewhat smallerspores. It differs from M1.aurantiacuswhich it resembles
in size, by the scarcity of broom cells on edges of the lamellae and surface of the
stipe and likewise by the absence of setulae on the majority of the epicuticularele-
ments of the pileus, and in the color of the pileus.
ILLUSTRATION. Singer (1958a) fig 25.
Stirps Cladophyllus
This stirps is intermediatebetween seriesLeonini and Actinopodes; it is charac-
terized by the type of stipe-covering,crowded to close and often anastomosinglamellae
and often glabrescentstipe; there are two subgroups,one with, one without distinct
pleurocystidia.
151. Marasmiuspusio Berkeley & Curtis,Ann. Mag. Nat. Hist. 12: 427. 1853.
lMarasmius guatopoensis Dennis, Kew Bull 15: 95. 1961 (a variety).
On dead dicot wood in tropical rain forest and tropical montane forest, arising
from rhizomorphsor directly from the substratum.
MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, Guayaramer'n, 18 Mar 1956, Singer B
2122 (LIL); La Paz: Nor-Yungas, Charobamba, 30 Jan 1956, Singer B 722 (LIL) type.
ILLUSTRATION. Singer (1965) pl 16, fig 92.
Marasmius sect Sicci subsect Siccini ser Haematocephali Singer, ser nov
Cystidiis ad latera lamellarum bene evolutis. - Typus: Marasmius haematocephalus
(Montagne) Fries.
Pleurocystidia present.
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 13 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 sub-
distant.
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
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.
204 FloraNeotropica
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 Valnrhloiblurgiasilvestris 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).
206 Flora Neotropica
very long (up to 20 X22), the hyaline cells evenly distributedand the surface appearing
mottled scalp preparations.
On dead fallen dicotyledonous leaves in rain forest.
in virginforest,
MATERIALSTUDIED. BOLIVIA.Beni:Vaca Diez, 3 km below Guayaramerin
12 Mar1956, SingerB 1879 (LIL) paratype,B 1920 (LIL), type.
ILLUSTRATION: Singer (1965) pl 21, fig 111.
StirpsOleiger(no. 154-156)
ronate, 20-55 X 8-12.5 p with conspicuousoily contents (not so strongas inMA.oleiger), hy-
aline or melleous, opaque, refractive;cheilocystidiaexactly like epicuticularbroom cells.
Hyphaestronglypseudoamyloid. Corticallayers: 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-acuteor obtuse,
spinulose.
On fallen dicotyledonous leaves and moncotyledonous herbaceousstems, perhapsalso
on other debris.
MATERIAL STUDIED. VENEZUELA. Monagas, S. of Quebrada Seca, 16 Jul 1972, Dumont
et al V'E5173 (NY). BOLIVIA. Beni: Vaca Diez, Guayaramerin, 3 km below village in virgin forest,
12 Mar 1956. Singer B 1875 (LIL) type.
Stirps Megistosporus
cate, glabrous,broadly and shallowly conical with broad obtuse tip, almost convex,
about 35 mm broad. Lamellaesordid white, rather broad (up to 4 mm broad), sub-
distant, not intervenose, rather narrowlyadnexed. Stipe chestnut black, glabrousor
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-acicularto subclavate,in profile almost
Fusarium(conidia)-shapedbut only weakly curved, thin-walled,hyaline, smooth, in-
amyloid. Hymenium:basidia33-39(-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 variouslyappendiculateeven with oblique protractedapiculi or
forked or with short irregularexcrescences,thick-walled,opaque, without conspicu-
ous contents, often distinctly deeper rooting than the basidia and some strongly pro-
jecting beyond the level of the sterigmata,others either not deep-rootingor not pro-
jecting; cheilocystidia much like the epicuticularbroom cells. Hyphae of hymenoph-
oral trama (which is regular)hyaline, strongly vinous pseudoamyloid,with numerous
clamp connections. Corticallayers: 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.
MATERIALSTUDIED. BOLIVIA. Beni: Vaca Diez, Guayaranerin,14 Mar1956, Singer
B 1971 (LIL), type.
This species is among all Marasmiiremarkablebecause of the enormously long
spores. These it shareswith Marasmiusmegistus 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 perishablespores
would be a handicap to any but a rain forest species where, at least duringthe larger
part of the year, a constant high degree of humidity is maintainedon the substratum
as well as in the lowest air-level.
ILLUSTRATION: Singer(1965) pl 16, fig 93.
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
marblingthe surface in scalp view, some with pale melleous fuscous, some with hy-
aline setulae, exuding a very short-livedpink 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.
Singer (1965), pl 18, fig 100 M.
ILLUSTRATION.
but entirely hyaline. Corticallayers: epicutis of pileus with broom cells which have a
mottled (in the grayishlilac forms) or uniform(in the deep violaceous specimens)appear-
ance and appearin mass deep brown-redto spadiceous,the individualsetulae porphyry
brown to fuscous.
On monocotyledonous leavesand on dicotyledonous sticks and branchletsfallen
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.
Sospiro,4 Aug 1969, SingerM 8976 (F), nearChontalpa31 Jul 1969, SingerM8776 (F); Veracruz:
7 km S of Montepio,Estaci6nBiologicade los Tuxtlas,20 Jun 1969, SingerM 8064 (F); Puertode
Veracruz,2 Oct 1962, Yolanlda Ayala (HerbariumENCB). CUBA. Sanvalle1868, "misitWright"
(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. ArenaForest2 Aug 1947,
Baker1488 (K). COLOMBIA.Garz6n,29 Nov 1939, Garces0., det. Charles(BPI);Buenaventura,
Singer(COL). VENEZUELA:Sucre,Rio Media,15 Jul 1972, Dumont et al VE5127 (NY). BRAZIL.
Amazonas,roadfrom Humaitato Labrea,29 Nov 1966, Prance,Pena&Ramos3461 (F).MinasGerais,
AgriculturalCollegelands,LocalidadVicosa, 5 Mar1930, YnesMexia4429 (UC). ECUADOR.Napo,
LagoAgrio,9 May 1973, SingerB 7328 (F). BOLIVIA.La Paz:Nor-Yungas,La Cienaga,E. R. de la
Sota, comm. & det. SingerB 1067 (LIL);Suapi, 1250 m alt, 9 Feb 1956, SingerB 1045 (LIL);Charo-
bamba,1200 m, 30 Jan 1956, SingerB 740 (LIL). ARGENTINA.Misiones:Frontera,GeneralMan-
uel Belgrano,19 Apr 1957 SingerM 1090 (LIL);Salta:Guemes,Quebradade
Yaquiasme,800 m alt,
7 I eb 1957, SingerT 2938 (LIL),T 2956 (LIL);Tucuman:Capital,Parque
Avellaneda,22 Mar1949,
SingerT241 (LIL);Jardindel InstitutoMiguelLillo, 14 Jan 1952, 20 Feb 1955, T 2191 (LIL);Buru-
yacii in subxerophyticwoods, 19 Feb 1957, SingerT 2992 (LIL);Tafi, ParqueAconquija,800 m alt,
12 Mar1949. SingerT222 (LIL);Diquede Escaba,31 Mar1957, Singer30671
(LIL). Also material
from tropicalAfrica(see Singer 1964b p 377) and Asia.
The materialfrom New York is probablyadventitious.
StirpsHaedinus(no. 161-162)
161b.Marasmiushaedinusvar minor Dennis, Trans. Brit. Mycol. Soc. 34: 421. 1951.
TYPE.Dennis 57 A, from Trinidad.
Pileus only 5-10 mm, lamellae slightly less distant. Otherwiselike the type var-
iety (see also Dennis 1951a where a full descriptionand a colored figure, 1, 19 fig.
10, is given). Trinidad.
Dennis (1951a) describesalso a var minrlor forma brevisporaDennis 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,Marasmiiusmicrohaedin,usSinger (no. 133). I have not studied
it.
163. Marasmiushypophaeus Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 298. 1869.
Fig 74.
TYPE. Wright,from Cuba.
Pileus dried rusty brown or rusty orange, not pallid-stripedbut at first unicolor-
ously "leafmold"(M&P), later "henna" in the center and along the ridges of the sul-
cate region, between ridges reaching"Mandalay"(M&P),broadly and rathershallowly
sulcate over 2/3 to 5/6 of the radius,glabrous,shallowly conical and subumbonatebut
at first often conical and papillate, eventually more obtuse and more repand,even with
umbilicus 7-17 mm broad. Lamellaepale grayishcream or cream, with at least part-
ially 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, narrowlyadnexed to rounded-free. Stipe rusty brown to chestnut
brown, becoming graduallydeep chestnut below, with at first cream colored or whit-
ish pallid apex, the whole stipe tending to become light to deep umber in the herb-
arium, glabrous,equal or taperingupwards,sometimes with slightly broadenedbase,
8-43 X 0.8-1.5 mm: basal mycelium scanty to ratherabundant, whitish to yellowish,
sometimes developing a himantium-likepellicula 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, rathernumerous, sometimes very numerousand crowded, versiform,often cla-
vate or subulate, fusoid-ventricose,cylindrical or broadly ventricose, acute or more fre-
quentlv broadly rounded at the apex, but occasionally or frequently with a subapicalor
apical appendageor mucro, well differentiatedandrefringent-opalescent,withoutdiffer-
entiated contents, hyaline: cheilocystidia like the epicuticularbroom cells. Hyphae of
the pileus-tramahyaline, 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 versiformbut often clavate,
with the wall often thickened in the upper portion, entirely subhyalineor with the upper
portion golden melleous to ochraceous brown, setulae pale brownish ochraceousor deep
ferruginous,1-14(-20) X 0.7-2.2 p, epicutis often mottled in scalp view.
On leafmold and rotten wood, also on accumulatedherbaceousstems, 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, SingerB 7454 (F).
This species is by no means the same as Marasmiushaemzatocephalus as Murrill(1915)
thought, even though the color of the fresh pileus is not known with certainty. Dennis
218 FloraNeotropica
167. Marasmius anomalus Lasch in Rabenhorst, Klotzchii Herbarium Vivum no. 1806.
1854 (non Peck, 1872).
Androsaceusepodius var imicrospornsMaire,Bull Soc. Mycol. France24: Ivi. 1908.
Marasmiusepodius var microsporus(Maire)Kuhner,Botaniste25: 101. 1933.
MarasmiuscoprophlilusSpegazzini,Bol. Acad. Nac. Ci. C6rdoba29: 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,campanulateto convex, then
Marasmius 221
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, roun-
ded to subacute at tip, rarely apex incrusted with a rusty granular-resinousincrustation,
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 epicuticularelements
although never thick-walled. Hyphae thin-walledand hyaline in the pileus and the hy-
menophoraltrama, 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 hy-
aline or main body more or less golden, the latter thin-walled,rarely thick-walled,
9.5-21 X (3-)5-8 1, setulae thin-rodshapedor spinulose, obtuse to subacute, 2-7 X 1-
1.5 M,few occasional epicuticularelements smooth (without setulae).
On fallen dead twigs of dicotyledonous tree in subtropicalforest, perhapsalso on
leaves, gregarious.
MATERIAL STUDIED. ARGENTINA. Misiones: Arroyo Piray Guazui, 24 Feb 1960, R. T.
Guerrero 61 (LIL), type.
This species was misinterpretedby 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 underMarasmiusferrugineusno. 169).
ILLUSTRATION. Singer (1965) pl 17, fig 98.
169. Marasmiusferrugineus(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).
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 narr-
owly conical, 4-5.5 1long.
On dead pieces of bark and dead branchesof dicotyledonous trees and shrubsin
tropical forest.
MATERIAL STUDIED.BRAZIL.MinasGerais.(Berkeleyset at K), holotype;Amazonas:
Panure(=Ipanure,Sao Jeronimo),Spruce 139 p p (K). BOLIVIA. La Paz: Nor-Yungas,Carmen
Pampa,19 Feb 1956, SingerB 1270 (LIL).
apart from the Berkeley Herbariumset which Dennis, to whom I am obliged for assist-
ance in this matter, considers the set from which the holotype of M. ferruginleus varfer-
rugineus 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 collec-
tions 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 char-
acteristic for M. ferrugineus sensu Dennis (195 la, non 1970 which refers to our IM.fer-
rugineus) and our M. hypophaeus (see there no. 163).
ILLUSTRATION. Singer(1958a) fig 29.
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, SinLgerB 12255 (LIL).
My intcrpretationof Marasmiusbainbusinusis based on the originaldescription.
We htavenot studied the type and do not know whether it is preservedand if so, in what
state it is. The original diagnosisdescribesit as having "lamellispaucis aequalibusad-
natis" whicli is quite correct for our species. In Epicrisis,probably because of a lapsus
plumae, Fries transcribes"lamellisadnatis paucis aequalibusvenosis albis." Whetherthe
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 narrowlamellae which, observed on a specimen that had been
pressed flat according to the methods of preparationat the time in use, will easily appear
venose and we know other cases where mycologists even more experienced in tropical
agaricsthantFries in 1830, have committed the same error in their diagnoses. Having
studied a large number of marasmioidagaricsin and around Brazilon bamboo (and
otlhermonocotyledonous leaves), I cannot think of any other Marasmiuswhich would
fit this descriptionand only in Marasiniellusdo we have similarlycolored (although less
brightly so) small carpophoresoccurringon that host.
Fries'species was redescribedand illustratedonly by Bresadola(1929), as far as I am
aware,and his materialcame from China rather than the neotropics. The color of this
species is much less bright than Beyrich's("luride fulvellusvel subochraceus"ratherthan
rufescens), has small spores and apparentlyinsititious stipe. This specimen (WU) is difficult to
determine but it seems to be a Marasmiellusratherthan a Marasmius,in fact perhapsthe
same as a smallMarasmiellussometimes occurringon bamboo in the neotropics. It can-
not be proved at the moment which of the two interpretationsis correct but if the orig-
inal Beyrich materialdoes not exist or is not in good condition, it seems that the inter-
pretation based on neotropical materialought to have preferenceinasmuchas it agrees
with the entire description of 1830 including the color of the pileus.
ILLUSTRATION. Singer (1965), pi 17, fig 96.
Stirps Helvolus
tant, adnexed to adnate, sometimes to a widened apical disc of the stipe. Stipe some-
what variablein color, fresh chestnut with ochraceouspallid to pale ochraceous brown
apex, then umber to liver brown, dried light tan to light fuscous, glabrous,shining,
smooth, mostly equal but sometimes with enlargedbase or with discoid enlargedapex,
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 characteristicfor some species of Cortinarius(alboviolaceus, etc).
Spores 11-15.2 X 2.8-4 ,, fusoid-clavateto 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 projectingnor deep-rooting,most frequently
clavate to fusoid-ventricose,often variouslyappendiculate;cheilocystidia in dried mat-
erial like epicuticularbroom 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-walledbut eventually
some with slightly thickened walls, in cresyl blue mounts a heterogeneoushyphal sys-
tem becoming evident which is metachromatic(pinkish) while the other hyphae become
pale blue, often irregularin 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-shapedand obtuse to spinulose and acute, offer-
ing a mottled surface in scalp preparations.
On dead leaves, leaf petioles, small fallen twigs and woody parts of small vines,
apparentlygenerally on dicotyledonous hosts, solitary or gregarious.
MATERIALSTUDIED. BRAZIL. Amazonas:Panure(SSo Jeronimo)Spruce 139 (K), type.
ECUADOR. Napo, Shushufindi,13 May 1973. SingerB 7412 (F). BOLIVIA. Pando: Manuripi,
Conquista,25 Mar 1956, SingerB 2235 (LIL), 26 Mar1956, SingerB 2282 (LIL).
Marasmiushelvolus is often similarin colors to M. helvoloides (see no. 130) which
should be compared. AMarasmnius helvohls has somewhat inconstant cystidial characters
and may be looked for in seriesLeonini where it is also keyed out.
ILLUSTRATION. Singer (1965), pi 16, fig 94.
Stirps Confertus
175. Marasmiusconfertus 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, glabroussulcate over half of
the radius or more, with smooth center, with a small depressionin the center and/or
subumbonate, 8-25 mm broad, mostly about 11-15 mm. Lamellaewhite, then becoming
cream, often with edges concolorous with pileus from marginpart way to the stipe but
not always so, rather narrowor narrow,about 22, intermixed with lamellulae,close or
subclose, free to adnate. Stipe ochraceous brown to chestnut, base often umber to al-
most 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 taperingupwards,hollow, 30-80 X 0.5-2 mm; basal mycelium strigose and tomen-
tose, wet watery pallid to ochraceous whitish, dried white with fulvous tawny tips or all
brownish fulvous; whitish to ochraceous rhizomorphsvery frequently present. Context
of pileus thin, white, inodorous.
Spores 10.5-13.7 X 3-4 p, fusoid-oblong,applanateinside when seen in profile, hy-
aline and smooth, inamyloid. Hymenium: basidia e g 21 X 5.5-7.5 p, 4-spored, some-
times 2-spored;cystidia 24-35 X 4-9 y, hyaline to melleous, moderately numerousto
numerous, opaque, smooth, versiform,e g ventricose-fusoid,cylindric or clavate, ampull-
aceous, often mucronate or appendiculatein some collections, well differentiated;cheil-
ocystidia golden hyaline to hyaline with more often hyaline than melleous setulae, other-
wise like the epicuticular broom cells, in some collections rather scattered, in
others crowded. Hyphae of trama of pileus and hymenophoraltrama hyaline, some with
distinctly thickened walls, others thin-walled,broad or narrowlyfilamentous, strongly
pseudoamyloid, with clamp connections, subhymeniumconsisting of small elongated and
small subisodiametricelements, hyaline. Corticallayers: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 numerousper cell, 3.5-6.5 X 1-2.2 (at base)j , alternatingsubhyalineto
golden hyaline and melleous to tawny, acute to obtuse of subacute, rod-shapedto spin-
ulose, surface mottled in scalp view.
On termite nests (woody materialwith termite holes) on rotting wood in forest,
on monocotyledonous remaindersand generally forest debris with woody particles,
rarely solitary, usually characteristicallycespitose to fasciculate, in tropical-montaneand
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 precedingspecies in smallerspores, distinctly sulcate pileus,
smalleraveragesize and often cespitose habit.
The Venezuelan collection was compared with the type by Dennis. Petch redes-
cribedAI1.conifertusexactly the way we observe this species in South America. He in-
dicates as synonyms M. chondripes Berkeley & Broome and M. hemibaphusBerkeley &
Broome.
ILLUSTRATION.Singer (1965) pi 19, fig 106; Dennis (1961) fig 44.
turity but narrownear the margin, distant to very distant, 10-12 through-lamellaebut
lamellulaealso present, adnate to the fleshy disc or merely adnexed. Stipe "cocoa
brown" to chestnut with white to pallid apex when mature, in young stages concolor-
ous with the pileus with white apex, glabrousor slightly pruinate, glabrescent,equal,
30-63 X 0.5-1.5 mm, basal mycelium sometimes arising from a basal socle, fluffy-to-
mentose to strigose, white, strigosity partly especially at tips tending to become ful-
vous, usually forming gilvous-whiterhizomorphs. Context white in pileus: odor none.
Spores 9.5-13.2 X 3.5-4.5 p, hyaline, smooth, tending to germinateon the lamel-
lae. 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,
differentiatedclearly from the basidia by being opaque and/or rooting deeper than the
basidiaand basidioles, often subcapitateor appendiculate,hyaline, thin- to thick-walled:
cheilocystidia: Siccus-type broom cells, much like the epicuticularbroom cells, main body
hyaline, 18-25 X 4.5-10.7 jp, often cylindric or cylindric clavate, beset with slightly diverg-
ing 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 sub-
acute, narrow(1.2-1.5 y diameterat base). In the hymenium sometimes hyphal pegs pres-
ent, very scattered, hyaline, of interwoven filaments variablein size and shape. On edge
occasional transition between cystidia and cheilocystidia (smooth cheilocystidia) observ-
ed. Hyphae in trama of pileus irregularlyshaped arld somewhat interwoven, of different
diameterand some very short, some pale stramineous,some with moderately thickened
walls, but most filamentous, hyaline and thin-walled,all pseudoamyloid, with clamp con-
nections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of
the Siccus-type, main body 13.5-17 X 7.5-10U, sometimes shorter,hyaline, mostly some
golden ones intermixed, versiform, often vesiculose, setulae either all erect or some sligh-
tly 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.
MATERIALSTUDIED. BOLIVIA. La Paz: Nor-Yungas.Rio Yariza, 16 Feb 1956, SingerB
1194 (LIL), 23 Feb 1956, Singer B 1369 (LIL) type; Carmen Pampa 2000 m alt, 26 Feb 1956, Singer
B 1489 (LIL).
Singer (1965) pl 19, fig 107.
ILLUSTRATION.
25-30 mm broad. Lamellaenot pure white but a very pale sordid grayishwith inter-
mittently narrowly discolorous (rusty spadiceous) edge when seen dried under a strong
lens, broad, regularlyintermixed with lamellulae(didymous, but lamellulaeof variable
length), not intervenose, distant (14 through-lamellae)rounded-subfreejust reachinga
brownishdisc around the extreme apex of the stipe. Stipe brown, at first with pallid
apex, glabrousbut under a strong lens appearingslightly 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 rathernumerousto very
numerous, 18-35 X 5.5-7 A,most opalescent, fusoid-subventricoseor ampullaceous,some
appendiculateor mucronate, with mostly thickened wall, inamyloid;cheilocystidia scat-
tered and present only where the lamellae-edgesare discolored, like the epicuticular
broom cells. Hyphae of the pileus hyaline, thin-walled,with clamp connections, pseu-
doamyloid. Cortical layers: epicutis of the pileus hymeniform, consisting of broom
cells, main body 7-14 X 4.5-8.2,, mostly clavate, subhyalineto 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 rathernumerousbroom cells but many with
strongly reduced main body and long, often spreadingor 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).
179. Marasmiushinnuleus Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 297. 1869.
Fig 79.
TYPE. Wright155, from Cuba.
Pileus ferrugineous-cinnamonto fulvous, dried "terrapin"M&Por "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 dry-
ing, narrow to broad, not intervenose, distant, adnexed. Stipe dried umber to brown be-
low with white apex at least at first, setose, glabrous,equal, 25-35 X 0.2 mm; basal my-
celium 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 epicuticularbroom cells but often scarce
and entirely hyaline or few with bright melleous setulae. Hyphae of the pileus-trama
hyaline, with clamp connections, pseudoamyloid. Corticallayer: 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 narrowlyconical, erect, hyaline, golden-hyaline,
sometimes some brightermelleous ones on some cells.
On dead leaves in Cuba. Known hosts: Cecropiapalmata,Hibiscus sp.
MATERIALSTUDIED. CUBA. Wright155 (FH) type; on "Paritium"(Hibiscus,Malvaceae)
(FH); Sebaruco,Soledad,9 Jul 1941, White846 (FH, BPI).
In view of the ratherpale color of most of the epicuticularbroom cells the color
of the fresh fungus might be slightly paler than describedabove from dried material.
agreeperfectly with the type which is evidently not fully maturesince there are only few
spores in it. Yet, there are also, as often in Marasmius,individualswhich are actively and
abundantlysporulatingwithout havingreachedthe charactersof a fully matureor old car-
pophore. Specimenscharacterizedby largersize and sulcate pileus and studied by me are
those from Peru: B 1723 from the Beni, the one from Argentinaand the one from Florida.
Marasmiusspegazziniidiffers from most cystidiate species of this section in its small
spores.
floridalus Murrillwas originallycomparedwith M. berteroiby Murrilland
Mlarasmliuls
thereforethought to be acystidiate inasmuchas an acystidiatespecies (M. leoninus) has ac-
tually been found in Floridabut the type is clearlycystidiate and identicalwithM. spega-
zzinlii.
Marasmzius virginianusSingerfrom the temperatezone of North Americais similar
but differs in habitat, somewhat smallersporesand slightly largersize and distinctly inter-
venose lamellae,especially nearthe peripheryof the pileus. See Singer(1958a, p 112).
The following species,also very close to M. spegazziniiandM. virginianusdiffers in
habitat, somewhatlargerspores, distinctly though not extensively intervenoseand at the
edges discolorouslamellae.
ILLUSTRATION. Singer(1965) pl 20, fig 108.
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 stramine-
ous, 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 epicuticu-
lar broom cells or cystidia pseudoamyloid. 190. M. pseudoniveuts.
14. Pileus cinnamon when fresh; epicutis composed of broom cells only; cheil-
ocystidia 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.
16. 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. 1Mcohaerens.
19. Setoid cystidia hyaline (see "14" above).
ico dein convexo, 8-14 mm lato; lamellis albis vel aurantiaco-pallidis,ad aciem aurantiacis
ve] ferrugineo-aurantiacis praesertimin siccus, confertis vel paene confertissimis;stipite
ferrugineo-castaneo,dein obscuriore,pubescentevel velutino, apice albo vel aurantiaco-pal-
lido, 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 cystidiiformibusnumerosis9-80 X 4.11 /j, pleruinquecy-
lindraceisornato. Ad folia arborumdicotyledonearumin 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, even-
tually convex-applanatewith slightly depressedcenter, obtuse, more rarely somewhat um-
bonate, 8-14 mm broad. Lamellaewhite to orange-pallid,the edges discolorous (pale or-
ange to rusty orange) especially after drying, rathernarrow,close to almost crowded, roun-
ded-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 taperingupwards,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 broadenedin
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 apicalappendagewhich is often oblique, sometimes con-
stricted 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 largerthan the
basidioles, best seen in cresyl blue mounts; cheilocystidia like the epicuticularbroom
cells, making the edge hetermorphous. Hyphae of the pileus-tramahyaline, thin- to mod-
erately tlhick-walled,with clamp connections, pseudoamyloid. Hyphae of the stipe yellow,
sometimes tending to olive in the rind layer, parallelwith each other, with clamp connec-
tions. 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-hyalineand scalp preparationstherefore appear-
ing mottled; covering of the stipe consisting of cystidiform hairs, these numerousor even
dense, 9-80 X 4-11 , most cylindric with rounded-obtuseapex, 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, subhya-
line 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: Platainusoccidentalis.
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. MarasmiuspersonatusBerkeley & Curtis, Jour. Linn. Soc. Bot. 10: 297. 1869.
TYPE. Wright10, from Cuba.
Pileus white, drying yellow, closely striate towards the margin,otherwise smooth,
convex, eventually depressed,6-10 mm broad. Lamellaecream, edge concolorous, crowd-
ed, of three lengths, adnexed. Stipe fulvous or dark brown, minutely pubescent(glabrous
accordingto 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
protrudingabove the basidia(Dennis); cheilocystidia like epicuticularbroom cells. Hyphae
Marasmius 239
This group is characterizedby 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-setaldermatocystidiaor long hairspresenton the pileus in some species.
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, vary-
ing 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.Marasmiusatrorubensvar atrorubens
This agrees in most characterswith var cystidifer but Dennis (1951b) describes
the materialas having no pleurocystidia.8Acystidiate 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-ferruginouswhen 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 correspondto 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:
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.
?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.
244 FloraNeotropica
either entirely hyaline or with very pale stramineoussetulae. Hyphae of the trama of the
pileus hyaline, most filamentous but some (especially near the epicutis) swollen and short-
celled, thin- to firm-walledbut not distinctly thick-walled,with clamp connections, pseu-
doamyloid. Cortical layers: epicutis of the pileus hymeniform, consisting of broom cells,
these with a main body 8-20 X 6-9 A, cylindricalto clavate or subvesiculose,stramineous
or hyaline or melleous-ochraceousto (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,rodshapedto narrowlyconical, acute or obtuse. Aside from
these, particularlyin the marginalzone of the pileus but not constantly so, some thick-
walled hairs present like those of the surface of the pileus but reachingup to 450 p in
length, all inamyloid, entire, wall 1-2 p thick, hyaline to stramineous,about 6-7 p broad
and taperingto an acute tip; covering of the stipe consisting of numeroushairs, at least
27-170 p long, 6-11 Abroad, acute, the shorter ones often obtuse, thick-walledbelow (wall
there 1-1.3 Athick), most entire, few at base or at apex bifurcate, stramineousor (fewer)
hyaline, inamyloid.
On woody branchesand dead dicotyledonous leaves, Venezuela.
MATERIAL STUDIED.VENEZUELA. Sucre,TrailfromLosPocitosto Rio GrandeArriba,
13 Jul 1972, Dumont et al VE 4877 (NY) type.
the broom cells; covering of stipe consisting of numerousbut not crowded dermatocysti-
dioid hairs or setae which are 20-65(-100) X 4.5-9 u, entire or with one to three setuloid
or sterigmatoidapical erect excrescences, cylindric or ventricose, rarely clavate, apex ob-
tuse to rounded, rarely subacute, hyaline to stramineousor 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.
MATERIALSTUDIED. COLOMBIA.Valle: Buenaventura,San Joaquin,180 m alt, 22 Apr
1968, SingerB 6273 (F). BOLIVIA:Beni: Vaca Diez, 7 Mar1956, Singer1648 (LIL) type; 3 km
below Guayaranerfnin virginforest, 12 Mar1956, SingerB 1889 A (LIL), paratype.
ILLUSTRATION:
Singer (1965) p1 23, fig 122.
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 Marasmiuscoh-
aerens; in one species these bodies are all hyaline but then the lamellae are subdistant
and the pileus strongly pigmented (this species, Marasmiusflammans Berkeley, forms
the transition to stirps Actinopus, coming closest to Marasmiuspseudoniveus var amy-
locystis, which differs in close or subclose lamellaeand white pileus).
ored, inamyloid, much like the setae occurringon the surface of the pileus; sides of lam-
ellae 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 subhyalineand setulae short to medium long, moderately
thick at base, light golden melleous in some, subhyalinein other cells, spinulose;setae
like those of the surface of the stipe; stipe surface of a discontinuouslayer consisting
of numeroussetae which are actually a special kind of broom cells but have the setulae
so strongly pigmented that they appearquite well differentiated,usually simple and con-
sisting 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
rarelyforking, mostly brownishor melleous (at least in lower portion), attenuate to a
subacute to needle-sharpacute tip; wall of lower portion up to 1.3 uthick; there all
transitionsfrom 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.
MATERIALSTUDIED. BRAZIL. Guapore,Guajaramirim, 20 Mar1956, SingerB 1806
(LIL), paratype. BOLIVIA. La Paz: Nor-Yungas,Rio Yariza,16 Feb 1956, SingerB 1206 (LIL)
type.
This is extremely close to M. fulvovelutinus Beeli from Africa (Zaire) which differs
in somewhat largerpileus, creamwhite to pale buff lamellaeand broadercystidia.
ILLUSTRATION. Singer (1965) pl 21, fig 112.
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 marginalbroom cells also
occurring(but very sparsely) on the sides of the lamellae in the neighborhoodof the
edge. Hyphae hyaline, thin-walledin trama of pileus and hymenophore, strongly pseu-
doamyloid, with clamp connections; nongelatinous. Cortical layers: epicutis of pileus
hymeniform, consisting of broom cells of the Siccus-type and setae and also some trans-
itional formations (between broom cells and setae); broom cells hyaline to pale stramin-
eous, few brownish-ocher at times, about 14.5-16 X 5.3-6 ,, main body mostly hyaline
or subhyaline, thin-walledor more rarely somewhat thick-walled,setulae collectively
sometimes spreadingto as broad as 8 s, setulae 4-8 , long, strictly apical; setae scattered
to numerousamong 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 mid-
dle of seta 2.5-4.5 Mthick, wall 0.7-1.2 , thick, smooth, weakly to distinctly pseudoamy-
loid, in older specimens often scarcely pseudoamyloid,tip acute to needle-sharp;trans-
itional formations characterizedby deep forking pseudoamyloidsetulae, the swollen base
assumingthe 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 Pteridophytaand leaves of Monocotyledones (Bromeliaceae),Dicotyledones (e
g Inga, mostly mixed leaf mold) perhapsalso on conifers (Podocarpus),solitaryor more
often in groups, not cespitose.
MATERIALSTUDIED. COLOMBIA.Valle: MunicipalidadCali; Saladito, 15 Apr 1968. Siiiger
B 6154 (F). BRAZIL. Rio Grandedo Sul, Rick (asM. echinatus, ty pe of M. echinatiilusand M. echinatus
Theissenex SingerTheissen), (FH), type. BOLIVIA. La Paz: Nor-Yungas,Coroico, 26 Jan 1956
SingerB 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), par-
atype; Las Lenguas,27 Feb 1957, Singer T 3026 (LIL);Tafi, CiudadUniversitaria,15 Jan 1955, Singer
T 2029 (LIL), 30 Dec 1951.
ILLUSTRATION:
Singer (1965) pl 21, fig 113.
layer of the trama of the pileus, the terminalcells 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 diverticula2-7 X 0.5-2.5 A, finger-likeand
mostly erect and apical, some more subulate but the tips of the terminalcells and the
diverticulamostly obtuse; dermatocystidiaalso mostly present but rather rare, similar
to the hymenial cystidia but more irregularand with less incrustations,e g 30 X 6 u;
hypodermiumof filamentous hyphae which ascend towards the terminalhyphae; cov-
ering of the stipe consisting of terminalmembers of the cortical hyphae and a few
dermatocystidia,these ampullaceousor fusoid but smaller than the epicuticularand
hymenial ones, all hyaline and inamyloid.
On wood or bark of dead shrubsand trees. Known hosts: Phoebe porphyria,
Durantaserratifolia,Eugeniapungens, 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 palaeotropicalspecies which is representedby
MarasmliuscampanellaHoltermann,(1898) as interpretedby Petch (1947, p 35; speci-
men: Peradeniya,rufous form, 14 Oct 1914, Petch 4171 (FH), studied by me) which
differs from A. isabellinusin narrowerspores (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.
cortical layers: epicutis of the pileus hymeniform,consistingof broom cells, main body
11.5-17 X 7-10 , hyaline or brownish,ventricose,setulae either as in the cheilocystidia
or orangebrownishin their majority(in maturecaps) and often simple.
On bark of dead branchesof Dicotyledones. Yungas.
MATERIALSTUDIED. BOLIVIA. La Paz: Nor-Yungas,26 Feb 1956, SingerB 1499 (F),
type.
yellow," very distant, 4-7 through lamellae, one or two forked or anastomosingby
lower lamellae or veins, rather narrow,adnateor concurrent. Stipe absent or sublateral
and inconspicuous, later lateral, inserted in the sinus in rear portion of pileus, rudimen-
tary, 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-walledor 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 divergentin places while in the entire cells the
setulae are generally apical and erect, in the majority of the cases all erect (Siccus-type),
cells alternatingsuccineous melleous and hyaline (scalp view: mottled) 8-10 in diameter,
setulae up to 4 long.
On fallen dicotyledonous leaves in tropical forest, perhapsalways on Cecropia.
MATERIALSTUDIED. VENEZUELA. Miranda:Guatopo, 25 Jun 1958, Dennis 1427 (K)
type. BOLIVIA.La Paz: Nor-Yungas,CarmenPampa,2000 m alt, SingerB 1496 (LIL).
Singer (1965) p 11 1, fig 60; Dennis (1961) fig 42.
ILLUSTRATION.
Spores 7-10 X 3-6u,, mostly about 8-9.5 X 3.5-4.8A, variablein size and shape
from as elongated as 10 X 3, to as broad as 8.5 X 4.5p, i e subfusoid-oblongto ellip-
soid, 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-subcylindricali e equally variableas the spores; cystidia present as
cystidioles, these neither deep-rootingnor strongly projecting, numerous,especially
near the marginof the pileus, 15-18 X 5-7u, clavate to ventricose, obtuse, with firm to
slightly thickened wall (wall 0.5-0.6p thick), entire, stramineousto stramineous-hyaline,
or hyaline; cheilocystidia like the epicuticularbroom cells but mostly hyaline or else
Marasmius 261
pale golden melleous here and there, never with long flagella-likeappendages,diverticula
either indistinct or setulose and then only 1-3p high. Hyphae of the pileus-tramaand
the regularto irregularhymenophoraltramahyaline, 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-walledones (wall 0.3-0.6,6) not or extremely slowly and weakly
pseudoamyloid,whereas the thick-walledones (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 variablediameter(2-122) but elongated. Cortical
layers: epicutis of pileus hymeniform or subhymeniform,consisting of broom cells
and similarcells which are devoid of setulae but often with a finger-likeoutgrowth, 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 stramineousor pale
golden melleous, mostly thin-walledbut some with slightly thickened wall, not in-
crusted, setulae extremely variablebetween short and broad and needle- to flagella-like
and thin and long (3-8 X 0.2-0.4 ), the short ones coarsely verrucoseto 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 transitionsbetween
the Siccus- and the Rotalis-type. Coveringof the stipe: the outermost hyphae of the
stipe-rindon their outer side beset with setula-likebodies (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 branchesfallen off dicotyledonous trees in wet forests,
gregarious.
MATERIAL STUDIES.VENEZUELA. DtoFederal,MountLaNaiguata, 25 Jun1971,
Dumontet al VE 725 (NY). BOLIVIA.LaPaz,Nor-Yungas, gorgenearSanJeronimo,1900 m
alt, 7 Feb 1956,SingerB 980 (F), type.
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 pseudoamyloidhyphae
in the tramaof the pileus.
the pileus surfaceis "formed of radiatinghyphae 5-6 ,wide, their terminalcells often
swollen to 30 X 12-13 ,. Tramanonamyloid."
In Dennis's (195 la) interpretation,the species may not belong in Marasiniusbut prob-
ably in Collybia. In Patouillard'sinterpretationit is a Marasmiusof this group. I have not
been able to find the type in Montagne'scollection (PC).
Pileus fresh "yellow beige" to pale fuscous (13 G 7), centralportion very frequently
deeperfuscous in youth ("winter leaf," "Englishoak," at times reaching"Congo")but
bleachingto palerand dirtierthan "ivory"and often found in intermediatecolors, dried
assuminga color near "olive wood," "teak wood," "mummy,""cocoa," "seal"in the cen-
ter, the marginremainingpallid or dryingto "cookie," "burntumber"or "roe," i e gener-
ally reddishbrown to grayishfuscous with palermarginand the more rapidlydried and the
less pressedthe less reddishin tone, subhygrophanousor almost non-hygrophanous,short
transparentlystriatetending to become short coarsely sulcatewhen dried, opaque to weak-
ly opimous, not viscid, glabrous,naked, smooth in fresh condition, with sometimessome-
what irregularmargin,not shiningor very little so when dried,convex, then irregularlyflat-
tened, subumbonate,more rarelyquite obtuse or distinctly umbonate, 18-45 mm broad.
Lamellaesordidbeige ("Malacca"to "Arizona"),bleachingin age to palerand dirtierthan
"ivory," driedwhitishto subconcolorouswith marginof pileus, usuallywhitish, subdistant
to distant, broad to very broad,ventricose,adnexed to rounded-adnexed,not intervenose
except in large,old specimensoccasionallyvery minutely intervenose. Spore printwhite.
Stipe whitish to grayishpallid or pale creamgray (usually about "seaside,tea time" to
"cracker"below and paler to pallid above), finely pruinateto pruinatefibrilloseespecially
above, but glabrescent,hollow, sometimesbecomingcanaliculate,equal, subequal,or with
slightlyenlargedbase, 20-50 X 2-9 mm; basalmycelium typically tomentose nearbase and
at the base unless scanty when base immersedin stronglyadheringearth particles,sordid
white to grayishor dull cream. Context white, unchangingbut under the cuticle concolor-
ous 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 innerside, thin-walled,inamyloid. Hymenium:basidia27-29 X 4.8-6.3 p, 4-
spored;cystidia none; cheilocystidiavery inconspicuous,filamentous,clavateor clavate-
forked, mostly basidiomorphic,sometimesseptate, hyaline. Hyphae: Hymenophoral
tramaregular,consistingof hyaline, thin-walled,subparallelhyphae;all hyphae with
clamp connections and stronglypseudoamyloid. Corticallayers:epicutis of pileus sub-
hymeniform,consistingof thin-walledhyaline or (by a dissolvedintracellularpigment)
mielleousto light fuscous cells which are clavateto balloon-shaped.
On earth among leaves or grassesin open places, in North Americain low altitudes,
in South Americaonly in the drierportions of the alpine and puna zones but preferring
the marginof wet places, water puddlesand generallywet soil, gregariousto generally
cespitose or subcespitose. Fruitingin South Americafrom Januaryuntil 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 Quichuapopulation uses this species as food. There the
specieshas a certaineconomic importancesince in those altitudes few other edible species
can be gatheredin quantity.
The South Americanhigh altitude populationsare morphologicallyinseparablefrom
the Californiatype. The whitish gray pileus turns reddishbrown in both the North Amer-
ican and the South Americanforms. Some differencesin spore measurementsare due to
individualvariabilityof the averagespore sizes in differentpopulationsbut do not coincide
with areaor altitude differences. A certainweak intervenosityof the lamellaecan likewise
be observedhere and there. The largersporesof the type of C. fimicola are explainedby
276 FloraNeotropica
228. Marasmiusstrictipes (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 stramineouscenter 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-rugoseor reticulate-rugosecenter, more rarely
entirely corrugated,with entire or lobed marginwhich is incurvedat first, convex, later
more applanateand even depressedaround a low and obtuse umbo or subumbonate, 26-
60 mm broad. Lamellaewhite to whitish, not or weakly intervenoseor 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 subglabrousbut with mealy apex, equal or subequalor 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 neighboringspecimens. Context white or
whitish, unchanging,odor of crabs or crayfish mixed with the odor of Marasmiusoreades.
Spores 7.5-8.5 X 3.8-5,, mostly about 7.7-8.3 X 4-4.8p , ellipsoid to oblong or
narrowlyovoid, smooth, hyaline, inamyloid. Hymenium: basidia 23-28 X 5.5-8 5, 4-
spored;cystidia none; cheilocystidia 26-28 X 2-5 p, irregularin shape and size, often
filamentous to cylindric-flexuousor cylindric-subclavate,stramineousto subhyaline.
Hyphae hyaline, with clamp connections, with thin walls, some with thick walls (these
having the aspect of hairs of Crinipellisand are particularlystrongly pseudoamyloid),
all pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform, consisting of
erect hyaline or subhyalinecells which are subglobose, clavate vesiculose or piriform,
entire, thin-walled,not intermixedwith dermatocystidia.
On rotting leaves and rotten pieces of wood of dicotyledonous trees growing
gregariouslyor mostly cespitosely, fruiting in summerand fall. Known hosts: Quercus,
Carpinus,Inga, Coffea. Restricted to the temperate to tropical-montanezones 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-
tlibn948 (ENCB), 7 Jul 1969, Singer M 8331 (F); near Chiltepec, 30 Oct 1960, Guzman 2786 (ENCB).
Kauffman(1918) pl 161. Smith (1938) fig 1 b, d.
ILLUSTRATION:
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 occurringin southern Florida has less chestnut brown stipe, more hygro-
phanous pileus, and an odor of honey; it may representa variety and, again, be the same
as Marasmiustortipes, see p 284.
finely fibrillose all over, but some glabrescent,often eccentric, equal or sometimes slight-
ly 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
coveringsubstratumwith white arachnoidfilaments, forming thin membranousfilms,
but sometimes scanty. Context white, inodorous or with slight crustaceous(shrimp) or
marinealgae 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 epic-
utis of the pileus but rather inconspicuous. Hyphae of pileus trama and hymenophore
varying from slightly and incompletely pseudoamyloid to distinctly and strongly pseudo-
amyloid, all hyaline and with clamp connections. Cortical layer: epicutis of pileus con-
sisting of subglobose, pedicellate or clavate-vesiculosecells which are hymeniformly
arrangedand either completely smooth and entire, or in some sections frequently nodulose
or hook-bearingor 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 branchesrotting on the ground in subtropical,
tropical-montane,tropical and subtropical-montaneforests, gregariousto 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).
Singer (1965) pl 12, fig 64.
ILLUSTRATION.
231d.Marasmiuscohortalisvaralachuanus(Murrill)Singer,comb nov
MiarasmiusalachuanusMurrill.Lloydia5: 140. 1942.
TYPE. Murrill,from Florida,U.S.A.
Pileus "CreamBuff" to "CartridgeBuff" (Ridgway),hygrophanous,becomingrather
uniformlymilk-white dried or faded, transparentlyreticulatewhen moist, scrobiculate-
rugosewhen dried, besides coarsely sulcate over about three quartersof the radiusat least
when freshand faded and over one side of the pileus, more rarelyslightly sulcate to sub-
smooth on the margin,glabrous,not viscid, convex, soon umbilicateor with depressed
center, eventually often applanateor irregular,8-33 mm broad. Lamellaewhite to buff-
ish white but sometimeseventually cream-grayish,intervenose,even anastomosingin some
old specimens,horizontal,variablein number,varyingfrom subclose to subdistant(distant
to very distant if only the through-lamellaeare taken into consideration),1-4 mm broad,
not ventricose,adnexed, often adnexed to a widened apex of the stipe and the eventually
either appearingdecurrentor separatingfrom the apex but not truly collariate. Stipe
white becomingtawny, russetand Marsbrown (in this order)when matureor when old,
remainingpallid at apex, smooth but sometimescanaliculate,macroscopicallyglabrous,
+ subpruinateundera lens, hollow, equal or taperingdownwards,30-48 X 1.5-4 mm;
basalmycelium abundant,sometimesextensive and membranous,white or cinnamon-pallid.
Context very thin, white, unchanging,with an odor of crabswhen quite fresh (of "anise"
accordingto Murrill);taste mild.
Spores (4.5-)6-8(-8.5) X (2.3-(3-4(-4.5) u, fusoid or ovoid-oblongto almost ellipsoid,
in profile mostly somewhatapplanateon inner side but generallywithout a distinct sup-
rahilardepression,thin-walled,smooth, hyaline, inamyloid. Hymenium:basidiarelatively
long, 4-spored;cystidia few like the cheilocystidiaand near the edges or none whatever:
cheilocystidianot very numerousin most collections and not makingthe edge of the
lamellaeheteromorphous,24-29 X 5.5-10.8 u, versiform,often clavate to vesiculose,hy-
aline. Hyphae of the regularhymenophoraltramahyaline, interwoven,thin- to slightly
thick-walled,not gelatinized,with clamp connections, stronglypseudoamyloid;in pileus
tramasimilar;in stipe tramaparallel. Corticallayer: epicutis of the pileus hymeniform,
consistingof erect smooth cells, 13-25 X 7.5-20 u, mostly vesiculoseto piriform,obtuse
or a few with appendage,without setulae, hyaline or subhyalinein KOH.
On leaf mold in humid forest, low, mesophyticand high hammocktype but also
in tropicalhammock, on dicotyledonous leaves, often of Quercusbut also other trees,
in the Gulf region includingall of Florida,Louisiana,Puerto Rico, fruitingfrom June
until October.
282 Flora Neotropica
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.
233. Marasmiussilvicola 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.," "amberbr.," "Martin-
ique" in the center, otherwise 9 F 6 to "walnut taffy," hygrophanous,fading to avellan-
eous, dried colored like M. pampicola or somewhat paler, glabrous,smooth or shallowly
to coarsely sulcate on margin,especially when dry, transparentlystriate over 1/7 to 4/5
of radiuswhen wet, convex, eventually applanate or with slightly depressedcenter, often
umbonate, 17-95 mm broad. Lamellaepale and sordid grayishcinnamon ("almond") but
in some collections more yellowish ("golden corn" to "saratoga")or more gray but in
many cases on sides and lamellae-groundin the part near the marginof the pileus concol-
orous with the latter, regularlyintermixed with lamellulae,the lamellulaeoften crisp
or venose, ventricose or subventricose,ratherbroad to broad (2-4 mm, mostly 4 mm
broad), with entire edge, rounded-adnateto rounded subfree or narrowlyadnexed,
rarely broadly adnexed to adnate, not intervenose,subclose to distant. Stipe pallid to
cream-grayish(concolorous with the edges of the lamellae), fresh subglabrousto 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, other-
wise smooth, tubular-hollowat least in upper portion, eventually always entirely hollow,
equal or towards the base slightly to distinctly subbulbousor attenuate, 20-118 X 1.5-7
Marasmius 283
many with swollen base (7-18 wide) and slightly narrowedto long-effilate apex, the
latter entirely or in the lower portion thick-walled(wall 0.5-2 p1thick), hyaline, some-
times melleous-hyaline,some with resinous or mucilaginous-granular, amorphousincrus-
tations, inamyloid, in some hairs with an inner wall-layervery slightly pseudoamyloid or
an outer layer amyloid; vestiment of the stipe consisting of similarhairs, some of these
as long as about 1 mm, the lower portion often pale melleous, the apex often long-eff-
ilate and acute, 11-70(-300) X (at base) 3-4 p, wall at base inamyloid or slightly amy-
loid, wall 0.3-0.7 , thick; near apex hairs with thinner wall and swollen base. 4-25 X
2.5-6 p, cylindric, obtuse-conicalor pulvinate, wall hyaline, 0.2-0.41thick.
On leaves and petioles as well as herbaceousstems of all kinds of hosts (Pteridophyta
to Myrtaceae)in tropical-montaneor fog forest. subtropical-montane,and rain forest
of the plains, also in inundated forest on earth-coveredfoliage, solitary or gregarious.
MATERIALSTUDIED. BRAZIL. Serrados Orgoos.1500 in alt. 25 Oct 1961, SingerB
4033 (F). ECUADOR. Napo, Lago Agrio, 8 May 1973, SingerB 7274 (F). BOLIVIA. Pando:Man-
uripi, Santa Rosa, 22 Mar 1956, SingerB 2146 (LIL): La Paz: Nor-Yungas.CarmenPainpa,200 m
alt, 26 Feb 1956, SingerB 1535 (LIL). ARGENTINA. Tucuman,between CiudadUniversitaria
and EstanciaSan 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 distributionas well as the size of the metuloid hairs of the
pileus and the stipe is apparentlysomewhat variable,and it is possible that some vari-
eties may be distinguished. The amyloid reaction of these bodies was observed in the
collection B 4033; the hairs were much more evenly distributedup to the apex of the
stipe in coll~ction B 1535 (on Pteridophyta). The size and distributionof the cystidia
also appearsto be variable.
but very scattered in the hymenium, 31-5-44 X 6.8-7 ,, ventricose with capitate apex,
constriction underneaththe capitulum about 2 A,across, capitulumabout 4.8-5 , diameter,
wall thin, hyaline, sometimes found only rarelyand on or near edge of lamellae. Hyphae:
subhymeniumsubcellular,consisting of very irregular,small elements, hyaline, not gela-
tinized. Hymenophoraltrama regular,consisting of hyaline to stramineous,thin, fila-
mentous, interwoven hyphae which are not or scarcely pseudoamyloid. Hyphae of the
stipe parallel,the thin-walledones mostly inamyloid or scarcely pseudoamyloid,the
thicker-walledones in superpositionweakly pseudoamyloid,with clamp connections, but
many clampless secondary septa present. Corticallayers: epicutis of pileus consisting of
a hymeniform layer of elements 16-25 X 7.5-9 A,,these fusiform to ventricose or vesicu-
lose-subclavate,smooth, non-incrusted,thin-walled;in distances between 25 and 40 p,
there are dermatocystidiainterspersed,these 24-29 X 10-14 1, varyingfrom ventricose-
subampullaceousand short-pedicellateto vesiculose-subclavate,thick-walled,cinnamon.
stramineousto melleous; furthermore,there are also long-setose hairs, remindingone of
those of Crinipellis,these (37-)130-255 X 8-16 u, not very numerousbut very conspicu-
ous, broadestjust above the hyaline to melleous base, equal above and there hyaline,
more rarely slightly taperingupwards,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 pseudoamyloidbut
often slightly amyloid; covering of the stipe consisting of the same hairs but these rather
scattered, between them surface layer of hyphae not differentiatedfrom the other cor-
tex-hyphae of the stipe.
On dead fallen dicotyledonous leaves in tropical-montaneforest, Yungas.
MATERIALSTUDIED. BOLIVIA. La Paz, Nor-Yungas,CarmenPampa,200 m alt, 26 Feb
1956, SingerB 15285 (LIL), type.
ILLUSTRATION.Singer(1960), pl. 38, fig. 4.
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-
fidateportion about 5-6 wide, apex 1.2-2.5(-3) wide with rounded tip, inamyloid.
Hyphae with clamp connections, inamyloid, parallelin the stipe, thin-filamentousand
not gelatinized in the pileus, hyaline or subhyaline. Cortical layers: epicutis of the
pileus hymeniformly arrangedconsisting of thin- to moderately thick-walledvesiculose
elements which may also be ampullaceousor ventricose, 18-27 X 10-13.5 u, hyaline to
brownish;among these short thick-walledelements, 12-19 X 6.5-10 ,, distributedin
almost irregularpattern among the basic elements, broadly clavate, wall increasingly
thicker towards the apex and reachinga thickness of 2 p, also increasinglymore pig-
mented 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. Hairsconspicuous, as describedabove.
On dead leaves of dicotyledonous trees in tropical forest. Solitary. Mexico.
MATERIALSTUDIED. MEXICO.Chiapas,between Finca Sospiro,and El Pozo 4 Aug
1969, SingerM18971. (F), type.
Gloiocephalasect Gloiocephala
Extremely long hairs absent; pileus not sessile; spores largerthan 6 u.
TYPE SPECIES. Gloiocephala epiphylla Massee
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.
(MarasmiuscaucasicusSinger 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 measurementsof the stipe and less grayish
colors on the marginalsurface of the pileus. An illustrationof the Caucasianspecies
given (Fig 104) for comparison.
with septa at intervalsof 6-70 p, not forming moniliform rows, thin-walled,with oily
contents, not agglutinatedat the surface;cystals 12 ,wide, tetrahedral,abundantin the
flesh, particularlyin the pseudostipe;hyphal tips 1.5-2.5 P wide. Corticallayer: pseudo-
stipe surface with dermatocystidiato 20 j long, 2.5-6 p broad, subclavateor subventricose,
with thin to slightly thickened wall, divergent,loose, smooth.
According to Corner(1953), the habit and generalstructureof the carpophoreis
the same as in Hormomitariasulphurea,from which it differs in white fruiting bodies,
smallerspores, narrowerhyphae, not forming moniliform rows of cells on becoming
secondarily septate, shorter caulocystidia, and absence of oleocystidia.
This latter characterkeeps me from transferringthis species to Physalacriaalthough
Corner(1950, 1970) recognizesa species in Physalacriawhich is likewise devoid of oleo-
cystidia, P. stilboides. I know neither this species nor Hormomitariaalbidula. The des-
cription of the latter has been adapted from Corner(1953).
While I have no doubt but that Hormomitariaas originallydescribedis congeneric
with Phvsalacria,I am not certain about the species described above, nor am I any more
convinced than Cornerwith regardto the position within the genusPhysalacriaof Phy-
salacriastilboides.
Nevertheless,for completeness sake, Hormomitariaalbidula is treated here. It was
12Basionym. Hormomitaria sulphurea Corner, Monograph of Clavaria and Allied Genera p 649,
1950.
306 Flora Neotropica
Physalacriasect Physalacria
Characters:Those of the genus, but with neither elongated and solid nor with hat-
or shield-likepileus. See fig. 118.
TYPESPECIES.Physalacriainflata (Schweinitz) Peck.
5. Physalacriaconcinna 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-likeprocesses up to 8 p long, irregularlyarrangedat 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-resinousmass, 10-17 , wide, thin-walled
or with firm walls, usually a long base, projectingup to 30 p; transitionsto cystidioles
few. Hyphae of the head 2-4.5 uwide, thin-walled,(?) subgelatinous,not inflated,
branchedat a wide angle, hyphae of the stipe composed of compact longitudinalhy-
phae 2-7 p wide, clamped. Cortical layers: base of the head apparentlynot different-
iated as a sterile zone though some of the cystidia of this part ratherlargerand slight-
ly thick-walled;superficialhyphae 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;dermatocystidiaas 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).
MATERIALSTUDIED. none.
Physalacria 309
I have not seen the type of this species but suspect strongly that Sydow's orig-
inal collection does not refer here but to P. tenera. In this case, the Florida species
describedby Cornerfrom whom I have adapted my descriptionabove would have to
be renamed. This problem can be solved by studies of the type locality of P. concinna.
ILLUSTRATION. Corner (1953) fig 4.
from Miami, Fla. (F 992 and F 1259, both determinedas P. langloisii(F) on coconut
shell) belong here.
A collection by Rick (MO 140328) and one by Furtado both from southern
Brazilare mentioned by Corner(1970) underP. aggregata,but the carpophoresare
somewhat larger(head 2-3 mm wide). I have not seen this materialwhich has the
spores of P. sanctaemartaeand the size almost of P. brasiliensis.
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 uncertainunless, as
we have done in the second and third edition of "Agaricalesin moderntaxonomy"
Singer(1962, 1974), it is derived from the charactersof the second species described
below. If so, it would turn out to be so close to Physalacriasection Pileolina that it
cannot be classified in any tricholomataceoustribe except Marasmieae.
An effort to collect topotypical materialin Ecuadorwas unfortunatelyunsuccess-
ful.
ILLUSTRATION. Patouillard(1900), fig 65, but see fig 118.
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Cap epic utiand -el
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s
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i r the
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t Fh rv e e oa, p. f p o"%
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smot Ls t idi um; broomcell;
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Marasmius 321
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Carpophore,upper right profile of pileus X 11/2,spore, and broom cell. 50. Marasmius
niebuilanimy.
hlajleicola.Carpophores,spores and broom celL 51. Marasmzius varderubnicans.Carpo-
variabdliceps
phore, spore, two broom cells. 52. Marasmius baeocephalius(B) and Ml.peckii (P). Cheilocystidium
(ch), broom cell of epicutis, spores, carpophore.53. Mlarasmiusnmagnisetulosus. Carpophoreand
elements of epicutis, type 1 and type 2. 54. Mfarasmiuspolycladus. Young carpophoreattached to
black rhizomorph(slightlyenlarged),maturepileus,two sporesand two broom cells.
322 Flora Neotropica
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324 Flora Neotropica
7,
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Marasmius 325
,.., . . ,. > a !
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den-natocystidia (of the pileus, dce, of the stipe, dcs), carpophore. 93. Marasniius latitiscitlospermux~~~~
haro h fes(i');dnaoysii'fth
the of'?,' ies(:p::hiocsii ch;salr lmnso
co~~~~~~~~~~~~~~~~~~~~~~~~~vering
thestipe.(blw.'.-:i''.C.
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).
326 Flora Neotropica
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LiteratureCited 329
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330 FloraNeotropica
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9
_____ . 1919. Bijdrage tot de kennis der Krullotenziekte. Bull. Dep. Landb. Suriname
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32(369).
List of Taxa 331
NUMERICALLISTOF TAXA
1. Crinipellis 3. Lachnella
1-1. C. macrosphaerigera 3-1. L. cecropiae
1-2. C. carecomoeis 3-2. L. subfalcispora
1-3. C. gracilis 3-3. L. caracassana
1-4. C. urbica 3-4. L. villosa
1-5. C. subtomentosa 4. Amyloflagellula
1-6. C. megalospora 4-1. A. pseudoarachnoidea
1-7. C. perpusilla 5. Flagelloscypha
1-8. C. bisulcata 5-1. F. polylepidis
1-9. C. pseudostipitaria 6. Marasmius
a. var pseudostipitaria 6-1. M. yalae
b. var mesites 6-2. M. polylepidis
1-10. C. catamarcensis 6-3. M. cyrillidis
1-11. C. schini 6-4. M. bactrosporus
1-12. C. atrobrunnea 6-5. M. leguminosarum
1-13. C. missionensis 6-6. M. earlei
1-14. C. herrerae 6-7. M. liquidambari
1-15. C. septotricha 6-8. M. splachnoides
1-16. C. podocarpi 6-9. M. hakgalensis
1-17. C. stupparia 6-10. M. cryptotrichus
1-18. C. coroicae 6-11. M. chiapasensis
1-19. C. tenuipilosa 6-12. M. radicellicola
1-20. C. mexicana 6-13. M. dysodes
1-21. C. foliicola 6-14. M. atroincrustatus
1-22. C. phyllophila a. var atroincrustatus
1-23. C. dicotyledonum b. var inodorosus
1-24. C. commixta 6-15. M. atlanticus
a. var commixta 6-16. M. eorotula
b. var junia 6-17. M. defibulatus
1-25. C. excentrica 6-18. M. tomentellus
1-26. C. myrti 6-19. M. llicis
1-27. C. albipes 6-20. M. crescentiae
1-28. C. alcalivirens 6-21. M. sphaerodermus
1-29. C. trichialis 6-22. M. magnoliae
1-30. C. sapindacearum 6-23. M. echinosphaerus
1-31. C. tucumanensis 6-2 3A. M. kroumirensis
1-32. C. dusenii 6-24. M. leveilleanus
1-33. C. rubida 6-25. M. scotophysinus
1-34. C. austrorubida 6-26. M. euosmus
1-35. C. purpurea 6-27. M. martini
1-36. C. insignis 6-28. M. carpenterianus
1-37. C. eggersii 6-29. M. munyozii
a. var eggersii 6-30. M. sanctixaverii
b. var flavipes 6-31. M. caliensis
c. var epiphyllus 6-32. M. multiceps
d. var lilaceipes 6-33. M. cupressiformis
1-38. C. sublivida 6-34. M. rotuloides
1-39. C. siparunae 6-35. M. panamensis
1-40. C. trinitatis 6-36. M. pararotula
1-41. C. perniciosa 6-37. M. vergeliensis
2. Chaetocalathus 6-38. M. apatelius
2-1. C. columellifer 6-39. M. rotalis
2-2. C. liliputianus 6-40. M. castellanoi
2-3. C. niduliformis 6-41. M.pandoanus
2-4. C. asperifolius 6-42. M. oaxacanus
2-5. C. carnelioruber 6-43. M. manuripiensis
2-6. C. aurantiacus 6-44. M. psychotriophilfis
332Flora Neotropica
INDEX OF E XSICCATAE
Baker, R. Dodge, B. G. J.
1488 (6-160). (no number) (1-15)
Balansa, B. Donk, M. A.
3379 (6-83); 3380 (6-231b); 4284 11554 (2-1)
(6-181). Dumont, K. P. et al.
Bartholomew, E. VE-234 (2-2); '.'6-llOa); 725
1935; 1735 (1-5) (6-207); 1919
Batista. Chaves 3859 (2-2); 3S -2); 3938 (6-189b);
2224 (6-141) 4194 (6-148); (6-206b); 4300
Bertero, C. G. (6-117); 4676 5a); 4877 (6-189c);
(M. berteroi) 5124 (6-19); 6-160); 5145
Bon, H. F. (6-38); 5173 ( ); 5980 (6-139);
4362 (1-12); 4397 (6-76) 5867 (6-188); (6-78); 6555
Boutan (6-31); 6580 ( 6707 (6-31);
341b (1-12) "
7287 (7-16); 6-5); 7375 (6-170)
Britton, E. G. Dusen, P.
611 (6-187); 612 (6-160) 320 (7-13)
Britton, N. L. & Seaver, F. J. Duss, P.
1352; 1540 (1-9b) 35 (2-4); 67 C 93 (2-2); 445 (2-2);
Brown, A., Britton, N. L. & Seaver, F. J. 484 (1-15); 48o ti-i5) 492 (1-9);
1350 (2-3) 513 (1-15); 539 (6-215); 594 (6-186);
Bruch, C. 1765 (1-9); (6-189b); 1769 (6-221);
67 (6-167). 1877 (1-9b); 1888 (1-9)
Buza, J. J. & Dennis, R. W. G. Earle, F. S.
173 (6-2) 36 (6-231d); 85 (6-154); 184 (6-20);
Cardenas, M. 269(6-6); (6-6); 501 (6-114)
243 (6-226) Eggers, H.
Carpenter, J. B. s n (1-371)
136 (6-28) Eiten, G. & L. T. & Felipe, G. M.
Cook, 0. F. 5254 (6-127)
1538 (6-160) Gaillard, A.
Corner, E. J. H. 21 (6-122); 52 (6-160b); 116 (6-145a);
769 (Hormomitaria albidula) 128 (1-25); 136 (1-8)
Curtis, M. A. (see Wright) Ghesquihre, J.
Demange 1355 (6-33)
67 (6-76) Glaziou, A. F. M.
Dennis, R. W. G. 9171 (6-231a)
42 A (6-173); J40 (6-3); 56 (6-161a); Goossens-Fontana, M.
101 A (6-106a); 101 B (6-106a); 137 109 (6-75); 605 (1-32)
(4-1); 158 (6-135); 164 (6-162); 231 Goretta, Pedro
(6-603); 232 (6-34); 246 A (6-140); 92 (6-145a)
255 (6-lOa); 265 B (6-227); 274 (6-211); Grisol
339 (6-86); 376 (1-40); 391 (6-47); 397 31 (1-33)
(6-188); 409 (6-76); 411 (6-106a); 414 Guerrero, R. T.
1011 (6-159); 1020 (6-112); 1022 50 (6-148b); 61 (6-168); 68 (6-150a);
(6-184); 1023 (6-84b); 1052 (6-196); 71 (6-231b); 73 (6-125); 74 (6-150a);
1968 (6-57); 1071 (1-29); 1071 A (1-21); 91 (6-181); 92 (6-149); 102 (6-148d);
1103 (6-151b); 1107 (6-175); 114 (6-148d); 118 (6-83); 132 (6-223);
1157 (6-112); 1165 (2-2); 1172 133 (6-126b)
(6-158); 1427 (6-201); 1472 (3-2); 1775 Guzmain, G.
(6-55); 1787 (11-1); 1800 (1-17); 1809 948 (6-228); 1582 (6-216); 1869
(5-1); 1876 (3-3); (6-180); 2601 (6-164); 2786 (6-228);
de Witte 2800 B (6-143b); 3666 (6-7); 4524
9432 (6-132)
(2-2); 4580 (6-52); 4630 (6-8); 6148
Digilio, P. L. & Singer, R. (6-182)
T 488 (6-138)
336 Flora Neotropica
Holden, G. C. Ocampo
35 (1-32) 516 (6-181)
Hostmann Occhioni, P.
297 (6-189b) 11 (RB 5.2.04) (6-227); 15 (RB 5.2.17)
JeEek (6-1 50a)
4 (1-4) Petch, T.
Johnston, J. R. 3920 (16-9); 4171 (6-198); 4238 (6-76)
556 (6-158); 579 (6-158); 580 (6-169b) Piedras, M.
Kermes 108 (6-182)
732 (6-101) Prance, G. T., Pefia, B.S. & Ramos, J. F.
Krieger, K. W. Fungi Saxonici 3461 (6-160)
1457 (3-4) Puiggari, J. I.
Langlois, A. B. 1644 (1-7)
350 (8-9) Puttemans, A.
Lazo, W. 860 (6-94a)
PU-124 (1-24) Ravenel, H. W.
Lebrun, J. 816 (6-15a)
3009 (6-102) Rick, J.
Leprieur, F. R. 213 (1-17); 422 (6-150a); 477 (1-39);
989 (1-4); 990 (6-165); 1018 (6-202); 576 (6-148d); 642 (6-83)
1023 (6-165); 1038 (6-227) SaInchez
Libert, M. A. 132 (6-180)
119 (6-106a) Sandwith, N. Y.
Lopez Gonzalez, A. 1580 (6-190)
119 (6-182) Schmitz-Levecq
Lorentz, P. G. 169 (6-23)
3035 (6-142) Schultes, R. E. & Cabrera, I.
Louis, J. 12869 (6-80)
11010 (6-24); 14862 (1-32); 14934 Sehnem. A.
(6-39); 14935 (6-75); 15216 (6-188) 1917 (1-1)
Lowy, B. Singer, R.
365 (6-223); 367 a(6-175); 423 B 53 (6-233); 54 (6-198); 88 (6-19):
(6-180); 447 (6-181) 88a (6-19); 403 (6-213); 426 (7-14);
Mc Farlin 436 (6-150a): 437 (6-169b);
56 (6-3) 531 (6-195): 532 (6-195); 535
Maguire, B. & Wurdack, J. (6-11Ob); 549 (6-198): 558 t1-18):
34796 (6-118) 590 (6-161a); 492 (6-160g); 656
Mains, E. B. (6-55); 668 (6-138); 669 (6-141):
3628 (6-150a); 3660 (6-126b) 670 (6-160i); 676 (6-148b); 686
Martin, G. W. 693 (6-126b); 714 (6-231b); 720
2260 ; 3469 (6-27); 3485 (8-8); (1-8): 722 (6-151c); 725 (6-126a):
8360 (6-150a) 734 (6-160d); 735 (6-170); 736
Martin, G.W. & Bouche, A. M. (6-113); 737 (6-160e): 738 (6-160i);
7549 (6-160c) (6-165); 739 (6-160i); 740 (6-160);
Martin, G. W. & Welden, A. 743 (6-160e); 747 (1-37c) 755 (6-
7279 (2-1); 7553 (6-152); 7606 114); 758 (6-19); 776 (6-156); 779
(6-204); 7673 (6-150a); 8406 (6-204); (1-37a); 782 (6-231b); 800 (6-64);
8562 (2-1); 8580 (2-1); 8682 (1-2); B 800a (6-69a); 860 (1-35); 871 (6-64);
8702 (6-26); 8733 (6-35) 873 (6-64); 935 (1-35);
Martin, G. W. et al. 944 (6-170); 945 (6-160e); 958
7552 (6-150a) (1-35); 980 (6-207); 1043
Martinez, M. (6-126b); 1045 (6-160); 1048
89 (6-145a) (6-2) (6-3); 1067 (6-160); 1068
Mexia, Ynes (6-175); 1079 (6-195); 1098
4429 (6-160) (6-82); 1106 (6-170); 1121
Mille, L. (6-188); 1126 (6-231c); 1134
14 (2-2) (6-214); 1168 (6-145a); 1180
Murrill, F. (6-165a); 1181 (1-37a); 1193
19885 (6-231d) (6-77); 1194 (6-176); 1206
Murrill, W. A. & Murrill, E. (6-192); 1210 (6-69b); 1219
84 (6-196); 142 (1-20); 163 (6-218); (11-1); 1245 (6-64); 1270
632 (6-217); 1047 (1-38) (6-169a); 1318 (1-35); 1323
Noack, F. (1-20); 1369 (6-176); 1378
34 (1-7) (2-2); 1394 (6-126b); 1395
Exsiccatae 337
HOSTINDEX
Caprifoliaceae 41 Magnolia 80
Carapa guianensis 147 Magnolia grandiflora 83
Carpinus 277 Malvaceae 233
Carya 156 Melanostomataceae 45, 213
Cecropia 58, 205, 212, 216, 243, 256, 299 Metahybe 37, 39
Cecropia pamata 233 Miconia 299
Celtis 310 Morinda citrifolia 56
Chrysophyllum 240 Musaceae (Musa, banana) 141, 159, 206, 224
Chusquea 20, 29, 55, 129, 224 Myrtaceae 68, 142, 288
Clibadium 300, 304 Myrtus 36, 253
Clibadium erosum 55
Clusia 307 Nectandra 241
Clusia rosea 307 Nectandra coriacea 158
Clusiaceae 142 Nothomyrcia 114
Coccolobis 156
Cocos (coconut) 311 Oleaceae 41
Coffea (arabica) 25, 31, 67, 253, 270, 277
Compositae 150, 300 Palmae 9, 198, 247, 293, 302, 310, 315
Crescentia 82 Panicum 224
Croton 299 Panicum maximum 243
Cynodon 20, 152 Panicum mexicanum 19
Cyphellopsis subglobispora 59 Paritium 233
Cyrilla racemosifolia 67 Paspalum 14
Persea 81
Dryopteris 213 Phoebe porphyria 103, 104, 119, 173, 253,
Dunalia breviflora 33 309
Durantia serratifolia 112, 25 3 Pinus (pine) 236
Piperaceae 30
Enterobolium 283 Pithecollobium scalare 148
Ericaceae 116 Platanus occidentalis 238
Eryngium 81 Podocarpus 11, 173, 250, 294
Eugenia 137 Podocarpus coriaceus 307
Eugenia pungens 25 3 Podocarpus parlatorei 26, 295
Host Index341
Agaricus setulosa268
atrorubens 241 strictipes276
bambusinus 224 stupparia26
calopus 221 urbica 15
carecomoeis 13 Collybiopsis62
ceratopus 247 CrinipeUis1, 3, 9
chortophilus 17 albipes 35, 36
cohaerens 247 alcalivirens37
columellifer 53 asperifolius56
ferrugineus 222 atrobrunnea11, 21
fulvus 172 austrorubida37, 43
griseoroseus 256 bambusae17
haematocephalus 210 bisulcata11, 18
hymeniicephalus 278 calosporus54
liliputianus 54 carecomoeis11, 13, 14, 15
oreades 273 catamarcensis10, 20
spinuliferus 247 commixta 12, 28, 32, 33, 34, 39, 41
splachnoides 70 coroicae 12, 18
splitgerberi 216 corticalis41
tenuissimus 258 dicotyledonum 13, 32
trichialis 38 dipterocarpi31
trichophorus 39 dusenii 37, 41
urbicus 15 eggersii6, 43, 45, 49, 51
Amyloflagellula 9, 60 epiphyllus(v.) 47, 48, 49
pseudoarachnoides 60 excentrica6, 34, 35
Androsaceus 62 flavipes(v.) 47, 48
corrugatus 191 foliicola 12, 30, 31, 32
epodius var. microsprus 220 ghanaensis23
glaucopus 240 gracilis 11, 14, 18
haematocephalus 210 herrerae12, 23, 28
leoninus 186 hirticeps29
longisporus 169 insignis45
nigrobrunneus 128 iopus 43, 44
rhodocephalus 210 junia (v.) 32, 33, 34
splachnoides 70 lilaceiceps(v.) 47, 49
litseae (v.) 14
Baumanniella 304 macrosphaerigera 10
brasiliensis 307 megalospora11, 17
mesites (v.) 12, 19
Chaetocalathus 1, 3, 9, 53 mexicana 12, 29, 32, 39
africanus 53 minutula 12, 34
aurantiacus 53, 57 mirabilis37
carnelioruber 53, 57 missionensis12, 22
columellifer 5, 53, 56 molfmoana 13, 25
craterellus 53 myrti 35, 36
liliputianus 53, 54, 56 nidulus54
niduliformis53, 56 occidentalis(ssp) 19
Chamaeceras 62 orientalis(ssp) 20
Collybia 272 patouillardii28, 30, 41
albogrisea 274 pemiciosa 2, 45, 51, 52
bisulcata 18 perpusilla11, 17, 18, 39
carecomoeis 13 phyllophila31
excentrica 35 podocarpi11, 25
fimicola 274, 275 pseudoarachnoides60
hymeniicephala 278 pseudostipitaria12, 19
nivea 276 purpurea44
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
ROLF SINGER
Rolf Singer was born in Schliersee, Bavaria, Germany, June 23, 1906, is a
naturalized American citizen, received his Ph.D. at the University of Vienna under
Richard Wettstein, a Dr. biol. sc. at the Academy of Sciences USSR, is Professor
honoris causa at the Universidade Federal de Pernambuco and honorary Profesor
Emerito of the University of Buenos Aires, has worked as a researcher and teacher
in Vienna, Barcelona, Paris, Leningrad, Cambridge, Massachusetts (Harvard
University), Tucuman and Buenos Aires, Argentina, Lincoln, Nebraska, Chicago,
Illinios (Field Museum of Natural History), Santiago, Chile and Mexico. Since 1968
he is Visiting Professor at the University of Illinois at Chicago Circle and is now also
Research Associate at the Field Museum after retiring as Curator in 1973. His travels
in the Neotropics include extensive and often repeated trips to South Florida,
Mexico, Colombia, the Bolivian, Ecuadorian and Brazilian Amazonia, the entire
Andine chain (Colombia, Ecuador, Peru, Bolivia, Argentina and Chile), and the
lowland regions of the tropics and subtropics of the Atlantic Coast in Brazil,
Paraguay, Uruguay and Argentina. Many of these trips were made during his ex-
tended residence in Tucuman, Argentina. His life-long interest is in the taxonomy of
the Higher Basidiomycetes, and the largest number of his nearly 300 publications
deal with this subject. He has also worked on mycorrhiza, particularly ecology of
the ectotroph, and on hallucinogenic fungi. He is corresponding member of the
Sociedad Botanica do Brasil, honorary member of the Sociedad Mexicana de
Micologia, life member of the Societe Mycologique de France, and a member of the
Mycological Society of America. He is a member of the Organization for Flora
Neotropica and served as Scientific Director 1966-1968.
Flora Neotropica is designed to present in monographic form taxonomic
accounts of all plants growing spontaneously within the WesternHemisphere
tropics. Geographic, ecologic, cytologic, anatomic, morphologic, chemical and
economic data will provide complementary information for each contribution.
Bibliography, citation of specimens and indexes are intended to facilitate
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