Fungal Diversity
Marasmius sensu stricto in Peninsular Malaysia
Tan, Y-S.1, Desjardin, D.E.2*, Perry, B.A.2, Vikineswary, S.1 and Noorlidah, A.1
1
Institute of Biological Sciences, Faculty of Science, University of Malaya, 50603 Kuala Lumpur, Malaysia.
Department of Biology, San Francisco State University, 1600 Holloway Avenue, San Francisco, CA 94132, USA.
2
Tan, Y.-S., Desjardin, D.E., Perry, B.A., Vikineswary, S. and Noorlidah, A. (2009). Marasmius sensu stricto in
Peninsular Malaysia. Fungal Diversity 37: 9-100.
A total of 43 species of Marasmius sensu stricto belonging to six sections (Globulares, Hygrometrici,
Leveilleani, Marasmius, Neosessiles and Sicci) are reported from Peninsular Malaysia.
Macro- and
micromorphological characters in combination with ITS sequence data were used to circumscribe species. Of the 43
species reported herein, 9 represent new species, viz., M. angustilamellatus, M. diminutivus, M. distantifolius, M. iras,
M. kanchingensis, M. kuthubutheenii, M. musicolor, M. ochropoides, and M. olivascens. An additional 19 species are
new records for Peninsular Malaysia. Phylogenetic analyses of 36 Malaysian Marasmius species were conducted based
on sequence data from nrDNA ITS-1–5.8S–ITS-2 gene regions. The ITS data support the circumscription of species
based on morphological characters and are useful in aiding species delimitations. The ITS data do not, however,
support the current sectional delimitation of Marasmius wherein most sections are not monophyletic. Comprehensive
descriptions, illustrations and photographs of each species, a key to aid in diagnosis, comparisons with allied species,
and a preliminary phylogenetic construct are provided.
Key words: agarics, Agaricales, mating studies, phylogeny, taxonomy
Article Information
Received 31 Dec 2008
Accepted 2 February 2009
Published online 1 August 2009
*Corresponding author: Dennis E. Desjardin; email: ded@sfsu.edu
Introduction
Malaysia is one of the mega-diverse
countries for most groups of organisms
including fungi; however, the diversity of
macrofungi and in particular Marasmius
species in Malaysia is poorly known. The
genus Marasmius (Basidiomycota, Agaricales,
Marasmiaceae) was first established in 1835 by
Elias Magnus Fries. Species in the genus are
decomposer fungi that colonize leaf litter,
twigs, trapped debris, lianas and dead wood
(Hedger et al., 1993; Singer, 1986). More than
1700 epithets have been published to date in
Marasmius although it is suspected that these
represent only about 500-600 species
worldwide (Desjardin unpubl.) The diversity of
Marasmius is strongly correlated with the
diversity of plants in the habitats where they
occur, with a higher diversity in tropical areas
and fewer species in temperate areas (Lodge et
al., 1995; Desjardin unpubl.). Because
Marasmius species are saprotrophic they serve
many important ecological roles in litter
decomposition, nutrient cycling, soil genesis
and as a food source for myriad molluscs,
arthropods and small vertebrates. Favourable
temperature and humidity conditions in Asian
tropical forests support a wide diversity of
Marasmius species. However, the unpredictable seasonality of fruiting and the shortlived and often solitary fruiting bodies are
major problems encountered when documenting their diversity in tropical forests.
E.J.H. Corner was one of the early
researchers who studied macrofungi from the
Malesian region. He published a preliminary
account of Marasmius sensu lato in which he
reported 121 species of which 103 represented
new species (Corner, 1996). Corner had
accepted a much broader generic concept of
Marasmius than most contemporary workers,
and he included taxa that are now accepted in
the
genera
Gymnopus,
Hemimycena,
Marasmiellus, Moniliophthora, Rhodocollybia,
Setulipes and Tetrapyrgos. Although most of
9
the species described as Marasmius by Corner
(1996) either do not belong to that genus or are
superfluous epithets, Corner's study provided
essential preliminary data and quality specimens (at E) of Marasmius sensu lato from the
Malesian area. Since Corner, there has been no
monographic study on the genus Marasmius in
Malaysia.
Singer (1958a,b) was one of the pioneers
in developing the generic and infrageneric
circumscriptions of Marasmius and allied
genera. In subsequent years (Singer, 1976,
1986) he refined his taxonomic construct and
restricted Marasmius to include only species
with smooth, hyaline, inamyloid, acyanophilic
basidiospores in combination with macro- and
micromorphological characters distinguished
mainly by the absence of features: viz., an
absence of long dextrinoid hairs, an absence of
strongly gelatinized tissues, and by a pileipellis
that lacks a trichodermial palisade of smooth
and elongated cells. Singer (1976) recognized
11 sections within Marasmius and this
classification system has been followed (with a
few exceptions) by contemporary researchers
(Gilliam, 1976; Antonín and Noordeloos, 1993;
Desjardin et al., 2000). In this study,
Marasmius was restricted to a monophyletic
lineage containing only sections Marasmius,
Sicci, Hygrometrici, Globulares, Neosessiles,
and Leveilleani based on Wilson and Desjardin
(2005). Marasmius is the sister taxon of
Crinipellis and Chaetocalathus species
(Moncalvo et al., 2002; Wilson and Desjardin,
2005). A re-evaluation of the diversity of
Marasmius sensu stricto, based on extensive
new field studies in numerous primary forests
in Peninsular Malaysia, on the examination of
type specimens and additional exsiccati (dried
specimen), and on molecular sequences data
was the focus of this research.
by distilled water, 3% potassium hydroxide
(KOH) or Melzer’s reagent. The terms used to
describe lamellae spacing refer to the number
of lamellae that reach from the stipe to the
pileus margin and do not include the lamellulae,
whose spacing is indicated by the number of
series present. Spore statistics include: xm, the
arithmetic mean of the spore length by spore
width (± standard deviation) for n spores
measured in a single specimen; xmr, the range
of spore means, and xmm, the mean of spore
means (± SD) when more than one specimen is
available; Q, the quotient of spore length by
spore width in any one spore, indicated as a
range of variation in n spores measured; Qm,
the mean of Q-values in a single specimen; Qmr,
the range of Qm values and Qmm, the mean of
Qm values where more than one specimen is
available; n, the number of spores measured
per specimen; s, the number of specimens
involved. Dried collections were accessioned
into the Mycology Herbarium (KLU-M) at
University of Malaya, Kuala Lumpur and into
the Harry D. Thiers Herbarium (SFSU) at San
Francisco State University.
Phylogenetic study
Materials and methods
Taxon Sampling
Molecular sequence data were generated
for 40 species of Marasmius. Of these, 36
represent Malaysian species and three are
North American in distribution. A sequence of
Marasmius rotula (DQ182506), type species of
Marasmius, was obtained from GenBank and
included in the analyses. Three species of
Crinipellis were chosen as outgroup taxa for
rooting purposes, based on previous results
(Moncalvo et al, 2002; Wilson and Desjardin,
2005) that indicated a sister-relationship
between Crinipellis and Marasmius. All
specimens used in this study are listed in Table
2.
Morphological descriptions
Samples of the genus Marasmius were
collected in various forests in Peninsular
Malaysia as itemized in Table 1 and Fig. 1.
Color terms and notations in parentheses are
those of Kornerup and Wanscher (1978). All
measurements and colors reported for
microscopic features were observed from dried
material rehydrated in 100% ethanol followed
Molecular techniques
The entire region of the internal
transcribed spacers (ITS-1 and ITS-2),
including the 5.8S gene, was analyzed in this
study. Genomic DNA was extracted from dried
herbarium material using an E.Z.N.A. forensic
DNA kit (Omega Bio-tek Inc., Doraville,
Georgia), and amplified using primers ITS1-F
and ITS4-B (Gardes and Bruns, 1993) or ITS4
10
Fungal Diversity
Table 1. Collecting sites sampled for this study.
No.
1.
Area
Ulu Gombak forest area, Selangor.
2.
University of Malaya’s Field Study Centre, Ulu Gombak, Selangor.
3.
Sungai Chongkak Forest Reserve, Ulu Langat, Selangor.
4.
Mount Nuang Forest Reserve, Ulu Langat, Selangor
5.
Kanching Forest Reserve, Rawang, Selangor.
6.
MNS Heritage Trail, MNS Kuala Lumpur.
7.
Fraser’s Hill, Pahang.
8.
Cameron Highlands, Pahang.
9.
Langkawi Island, Kedah.
10.
Pasoh Forest Reserve, Negeri Sembilan.
11.
Ulu Bendul Forest Reserve, Negeri Sembilan
12.
Endau-Rompin National Park, Johore
(White et al., 1990). The amplification reaction
mixture typically contained the following: 2.5
µl of dNTP, 1.25 µl of MgCl2, 1.25 µl of each
primer, 1.9 µl of BSA, 0.1 µl of taq polymerase,
2.5-5 buffer, and 2.5 µl of DNA template,
eluted to a final volume of 25 µl. The
amplification protocol consisted of 2 min at 95
ºC, followed by 35 cycles of 30 sec at 94 ºC, 30
sec at 46 ºC to 54 ºC, and 1 min at 72 ºC,
followed by a final extension of 5 min at 72 ºC.
Sequencing reactions utilized the same primers
as PCR, and were purified using an isopropanol
precipitation. 125 µl of 65% isopropanol was
added to the sequencing reations, vortexed and
centrifuged at 15000 rpm for 35 min. The
supernatant was removed, 125 µl of 75 %
isopropanol was added, vortexed and
centrifuged at 15000 rpm for additional 10 min.
Again, the supernatant was removed and the
pellet was air-dried for 30 min. Clean
sequencing reactions were suspended in 15 µl
Hi-Di formamide (Applied Biosystems), and
visualized on an ABI 3100 Genetic Analyzer
(Applied Biosystems, Foter City, CA).
Sequence fragments were edited and
assembled with Sequencher v.3.1.1 (Gene
Codes Corp., Ann Arbor, Michigan). ITS
sequence data were initially aligned with
Location
N 03˚19’,
E 101˚45’
N 03˚19’,
E 101˚45’
N 03˚12’,
E 101˚ 50’
N 03˚12’,
E 101˚52’
N 03˚20’,
E 101˚35’
N 03° 09’,
E101˚41’
N 03˚42’,
E 101˚44’
N 04˚27’,
E 101˚22’
N 06˚25’,
E 99˚45’
N 02°59’,
E 102°19'
N 2° 42’,
E 101° 54’
N 02˚25’,
E 103˚5’
Type of Forest
Tropical lowland rainforest
Tropical lowland rainforest
Tropical lowland rainforest
Highland forest
Tropical lowland rainforest
Tropical lowland rainforest
Highland forest
Highland forest
Tropical lowland rainforest
Tropical lowland forest
Tropical lowland rainforest
Virgin lowland rainforest
Clustal X v.1.81 (Thompson et al., 1997) using
the default settings, followed by manual
alignment with MacClade v.4.03 (Maddison
and Maddison, 2001). Edited sequences have
been deposited in GenBank (Table 2), and the
aligned dataset used for all analyses is
available on TreeBase (S2230; http://www.tree
base.org).
Phylogenetic Analyses
Parsimony analyses were performed in
PAUP* v.4.0b (Swofford, 2002) utilizing
heuristic search methods, with 100 random
stepwise sequence addition replicates, tree
bisection-reconnection (TBR), collapse of zero
length branches, and maxtrees set to 10,000.
All characters were equally weighted and
unordered, and gaps were treated as missing
data. Parsimony bootstrap (PB) analysis was
performed using 1000 bootstrap replicates,
each with a limit of 1,000,000 rearrangements,
closest taxon addition, and TBR branch
swapping.
The appropriate model of sequence
evolution for the ITS data was estimated using
the Akaike Information Criterion (AIC) as
implemented in Modeltest version 3.7 (Posada
and Crandall, 1998) and MrModeltest
11
Fig.1. Collecting sites for the genus Marasmius in Peninsular Malaysia. A: Langkawi Island, Kedah; B: Cameron
Highlands, Pahang; C: Fraser’s Hill, Pahang; D: Kanching Forest Reserve, Selayang, Selangor; E: Ulu Gombak Forest
Reserve area, Selangor; F: MNS Heritage Trail, Kuala Lumpur; G: Sungai Chongkak Forest Reserve, Ulu Langat,
Selangor; H: Mount Nuang Forest Reserve, Ulu Langat, Selangor; I: Ulu Bendul Forest Reserve, Negeri Sembilan; J:
Pasoh Forest Reserve, Negeri Sembilan; K: Endau-Rompin National Park, Johore.
(Nylander, 2004). Maximum likelihood (ML)
and searches employed the TrN+I+G model of
sequence evolution. ML topology searches
were performed in PAUP* v.4.0b, and
followed an iterative approach. First, a
neighbor-joining tree was constructed from
which the ML model parameters were
estimated. Next, the estimated parameter
values were fixed and a ML search was
performed. The resulting tree(s) from this
search were then used to re-estimate the
parameter values, fixed, and another likelihood
search was conducted. This re-estimation and
search step was repeated two additional times,
for a total of three ML searches, to assure that
all three searches had converged on equally
likely topologies. ML bootstrap analyses were
performed in GARLI v.0.951 (Zwickl, 2006)
and consisted of 300 replicates under a
GTR+I+G model of sequence evolution, using
12
the default run termination settings.
Bayesian analyses were performed in
MrBayes v.3.1.2 (Huelsenbeck and Ronquist
2001, Ronquist and Huelsenbeck 2003) under a
GTR+I+G model of sequence evolution.
Analyses consisted of two parallel searches,
run for 6 million generations each and initiated
with random starting trees. The default settings
in MrBayes were used for the incremental
heating scheme, and to set unconstrained
branch lengths [unconstrained: exponential
(10.0)] and uninformative topology priors.
Chains were sampled every 500 generations for
a total 12,001 trees from each run, sampled
from the posterior distribution. Trees sampled
prior to the chains reaching a split deviation
frequency of 0.03 were discarded as the “burnin”, while the remaining trees were used to
calculate the posterior possibilities (PP) of the
clades.
Fungal Diversity
Table 2. Species, collection information and GenBank accession numbers of specimens used in the phylogenetic analyses.
Sample
Crinipellis brunneipurpurea Corner
Crinipellis dipterocarpi Singer
Crinipellis malesiana nom. prov.
Marasmius abundans Corner var. abundans
Marasmius abundans Corner var. abundans
Marasmius abundans Corner var. abundans
Marasmius abundans var. aurantiacus Corner
Marasmius acerosus Y.S. Tan & Desjardin
Marasmius acerosus Y.S. Tan & Desjardin
Marasmius aciebrunneus Corner
Marasmius adhaesus Corner
Marasmius adhaesus Corner
Marasmius angustilamellatus Y.S. Tan & Desjardin
Marasmius angustilamellatus Y.S. Tan & Desjardin
Marasmius angustilamellatus Y.S. Tan & Desjardin
Marasmius aratus Massee nom. illegit.
Marasmius araucariae var. siccipes Desjardin, Retn. & E. Horak
Marasmius araucariae var. siccipes Desjardin, Retn. & E. Horak
Marasmius bambusiniformis Singer
Marasmius berambutanus Desjardin, Retn. & E. Horak
Marasmius berambutanus Desjardin, Retn. & E. Horak
Marasmius berambutanus Desjardin, Retn. & E. Horak
Marasmius brevicollus Corner
Marasmius aff. camerunensis Antonín & Mossebo
Marasmius cohaerens var. lachnophyllus (Berk.) Gilliam
Marasmius crinis-equi F. Muell. ex Kalchbr.
Marasmius crinis-equi F. Muell. ex Kalchbr.
Marasmius crinis-equi F. Muell. ex Kalchbr.
Marasmius crinis-equi F. Muell. ex Kalchbr.
Marasmius crinis-equi F. Muell. ex Kalchbr.
Marasmius crinis-equi F. Muell. ex Kalchbr.
Marasmius crinis-equi F. Muell. ex Kalchbr.
Marasmius diminutivus Y.S. Tan, Desjardin & Vikineswary
Marasmius distantifolius Y.S. Tan & Desjardin
Marasmius florideus Berk. &Broome
Specimen ID
JFK 84
TYS 111
TYS 346
TYS 425
TYS 460
TYS 513
TYS 515
TYS 458-Holotype
TYS 427
TYS 457
TYS 464
TYS 467
TYS 437
TYS 524-Holotype
TYS 528
KUM 152
TYS 529
MY 38
TYS 511
TYS 337
TYS 398
TYS 503
TYS 517
KUM 60134
DED 4071
TYS 338
TYS 341
TYS 366
TYS 367
TYS 412
TYS 447
TYS 466
TYS 514-Holotype
TYS 478-Holotype
TYS 480
Location
Indonesia
Malaysia-Pahang
Malaysia-Kedah
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Johore
Malaysia-Johore
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Johore
Malaysia-Kedah
Malaysia-Selangor
Malaysia-Johore
Malaysia-Johore
Malaysia-Kedah
USA-Tennessee
Malaysia-Kedah
Malaysia-Kedah
Malaysia-Negeri Sembilan
Malaysia-Negeri Sembilan
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Johore
Malaysia-Selangor
Malaysia-Selangor
Herbarium
SFSU
SFSU
SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
Genbank
FJ167646
FJ431208
FJ167628
FJ431212
FJ431210
FJ431209
FJ431211
FJ431213
FJ431214
FJ431215
FJ431217
FJ431216
FJ431218
FJ431219
FJ431220
FJ431221
FJ431222
FJ431223
FJ431224
FJ431225
FJ431227
FJ431226
FJ431228
FJ431229
FJ431230
FJ431233
FJ431232
FJ167669
FJ431234
FJ431236
FJ431235
FJ431231
FJ431238
FJ431239
FJ431240
13
Table 2 (Continued). Species, collection information and GenBank accession numbers of specimens used in the phylogenetic analyses.
Sample
Marasmius gracilichorda Corner
Marasmius gracilichorda Corner
Marasmius gracilichorda Corner
Marasmius gracilichorda Corner
Marasmius guyanensis Mont.
Marasmius guyanensis Mont.
Marasmius guyanensis Mont.
Marasmius haematocephalus (Mont.) Fr.
Marasmius iras Y.S. Tan & Desjardin
Marasmius kanchingensis Y.S. Tan & Desjardin
Marasmius kuthubutheenii Y.S. Tan, Desjardin, Vikineswary & Noorlidah
Marasmius kuthubutheenii Y.S. Tan, Desjardin, Vikineswary & Noorlidah
Marasmius leucorotalis Singer
Marasmius aff. leoninus Berk.
Marasmius leveilleanus (Berk.) Pat.
Marasmius leveilleanus (Berk.) Pat.
Marasmius luteomarginatus Desjardin, Retn. & E. Horak
Marasmius micraster Petch
Marasmius micraster Petch
Marasmius micraster Petch
Marasmius micraster Petch
Marasmius musicolor Y.S. Tan & Desjardin
Marasmius ochropoides Y.S. Tan & Desjardin
Marasmius ochropoides Y.S. Tan & Desjardin
Marasmius olivascens Y.S. Tan & Desjardin
Marasmius olivascens Y.S. Tan & Desjardin
Marasmius oreades (Bolton) Fr.
Marasmius oreades (Bolton) Fr.
Marasmius purpureostriatus Hongo
Marasmius rotula (Scop.) Fr.
Marasmius ruforotula Singer
Marasmius ruforotula Singer
Marasmius ruforotula Singer
Marasmius selangorensis Y.S. Tan & Desjardin
Marasmius selangorensis Y.S. Tan & Desjardin
Marasmius selangorensis Y.S. Tan & Desjardin
14
Specimen ID
TYS 396
TYS 411
TYS 442
TYS 472
TYS 028
TYS 329
TYS 395
TYS 523
TYS 483-Holotype
TYS 415-Holotype
TYS 348
TYS 364-Holotype
TYS 489
TYS 423
KUM 60142
KUM 116a
TYS 477
TYS 229
TYS 237
TYS 391
DED 7647
TYS 417-Holotype
TYS 384-Holotype
TYS 416
TYS 424
TYS 426-Holotype
PBM 2701
HDT 51606
TYS 328
PBM 2563
TYS 369
TYS 438
TYS 509
TYS 435
TYS 453-Holotype
TYS 454
Location
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Kedah
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Negeri Sembilan
Malaysia-Johore
Malaysia-Selangor
Malaysia-Kedah
Malaysia-Kedah
Malaysia-Selangor
Malaysia-Pahang
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
USA-Massachusetts
USA-Santa Barbara
Malaysia-Kuala Lumpur
New Hampshire
Malaysia-Negeri Sembilan
Malaysia-Selangor
Malaysia-Johore
Malaysia-Selangor
Malaysia-Selangor
Malaysia-Selangor
Herbarium
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
Genbank
FJ431242
FJ431244
FJ431241
FJ431243
FJ431245
FJ431247
FJ431246
FJ431248
FJ431249
FJ431250
FJ431252
FJ431251
FJ431253
FJ431254
FJ431256
FJ431255
FJ431257
FJ431259
FJ431260
FJ431258
FJ431261
FJ431262
FJ431263
FJ431264
FJ431266
FJ431265
DQ490641
FJ431267
FJ431268
DQ182506
FJ431269
FJ431271
FJ431270
FJ431274
FJ431273
FJ431275
Fungal Diversity
Table 2 (Continued). Species, collection information and GenBank accession numbers of specimens used in the phylogenetic analyses.
Sample
Marasmius siccus (Schwein.) Fr.
Marasmius tenuissimus (Jungh.) Singer
Marasmius tenuissimus (Jungh.) Singer
Marasmius tenuissimus (Jungh.) Singer
Marasmius tubulatus Petch
Marasmius tubulatus Petch
Marasmius tubulatus Petch
Specimen ID
DED 5255
TYS 345
TYS 468
DED 7659
TYS 490
TYS 502
TYS 518
Location
USA-Tennessee
Malaysia-Kedah
Malaysia-Selangor
Malaysia-Pahang
Malaysia-Johore
Malaysia-Johore
Malaysia-Johore
Herbarium
SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
KLU, SFSU
Genbank
FJ431272
FJ431277
FJ431276
FJ431278
FJ431281
FJ431280
FJ431279
15
Results and Discussion
Seventy-seven new ITS rDNA sequences
were generated as part of this study. Sequence
data were not obtained for five additional
Malaysian Marasmius species (M. nummularioides, M. persicinus, M. purpureobrunneolus, M. purpureisetosus and M.
somalomoensis) due to the presence of multiple
ITS amplification products. Sequence length
ranged from 695 bp in M. crinis-equi to 963 bp
in M. micraster. Several regions of the
sequence data, totaling 117 nucleotides, were
ambiguous in their alignment and excluded
from all analyses. Sequence ends were trimmed
to account for missing data in several taxa,
creating a final alignment of 551 characters for
77 ingroup and three outgroup taxa. Of the 551
characters comprising the dataset, 153 were
parsimony informative.
Parsimony search found 29822 equally
parsimonious trees of 680 steps long (CI=
0.4088, RI= 0.7561). The maximum likelihood
search converged on a stable average
likelihood of –lnL = 6033.7657 (Fig. 2). The
topology of the parsimony and ML trees were
similar, varying only in the arrangement of a
few species within terminal clades. Bayesian
analyses reached a split deviation frequency <
0.03 after six million generations, and the first
5812 trees were excluded as the burn-in.
All Marasmius species included in these
analyses belong to a monophyletic lineage
supported by 96% ML bootstrap (BS) and 1.0
posterior probability (PP) values. The nodes
representing potential infrageneric groups are
not well supported. Species currently
recognized as belonging to the sections Sicci
and Globulares fall into a clade with limited
support (< 50% BS and 0.79 PP); however,
neither section is monophyletic. Sect. Marasmius, representing species with a collarium, is
not monophyletic. Sect. Marasmius subsect.
Sicciformes (species with a collarium and
Siccus-type broom cells) is not monophyletic,
with members clustering into four separate
clades with statistical support for each clade
ranging from < 50% BS and 0.85 PP to 91%
BS and 1.0 PP. Sect. Marasmius subsect.
Marasmius (species with a collarium and
Rotalis-type broom cells) is monophyletic for
16
the four included species with 96% BS and 1.0
PP support. Only a single member of sect.
Hygrometrici was included in the analyses and
it is sister of two members of sect. Marasmius
subsect. Sicciformes but without statistical
support. The single member of sect. Neosessiles is sister of two members of Marasmius
subsect. Sicciformes but without statistical
support, and the single member of sect.
Leveilleani is sister of these clades but again
without statistical support.
In general, sequences representing
individuals belonging to the same morphospecies form well-resolved clades with high
statistical support. For example, three specimens of the new species M. angustilamellatus
that were collected in geographically distant
sites form a clade with 100% BS and 1.0 PP
support and show little genetic diversity. Only
M. crinis-equi broke this rule with a single
collection (TYS 412) resolving as sister of a
clade containing M. guyanensis plus the
remaining M. crinis-equi specimens but with <
50% BS and 0.85 PP support. Five additional
specimens of M. crinis-equi form a clade with
90% BS and 1.0 PP support. In a few cases
genetic diversity within species is quite high.
For example, the two specimens of the new
species M. olivascens that form a clade
supported by 83% BS and 1.0 PP values were
collected on the same day at the same site but
TYS 426 is on a long branch relative to TYS
424.
The genus Marasmius was restricted
according to Wilson & Desjardin (2005) to
include only members belonging to the sections
Marasmius, Hygrometrici, Leveilleani, Scotophysini, Sicci, Neosessiles and Globulares as
defined by Singer (1976, 1986). Members of
all of these infrageneric groups, except
Scotophysini, occur in Malaysia and are
represented in the molecular phylogenetic
analyses (Fig. 2). Representatives of the other
sections traditionally recognized within
Marasmius by Singer (1976, 1986) and others
were not included in this study; viz.,
Marasmius sect. Androsacei as the genus
Setulipes (Antonín and Noordeloos, 1993); sect.
Fusicystides as a synonym of Setulipes
(Desjardin, unpubl.); sect. Alliacei as the genus
Mycetinis (Wilson and Desjardin, 2005); and
Fungal Diversity
Fig. 2. Maximum likelihood tree using a GTR+I+G model of evolution. Numbers above branches at the left are
bootstrap percentage and the right are posterior probability values (Bayesian MCMC analysis).
17
sect. Epiphylli as a distinct genus allied with
Gloiocephala in the Physalacriaceae (Desjardin,
unpubl.). As a consequence of our delimitation
of Marasmius sensu stricto and a sampling
design influenced by the nLSU DNA dataset of
Wilson and Desjardin (2005), this study based
on ITS sequences also resolved a monophyletic
Marasmius with high statistical support.
Species concepts based on shared
morphological features were well supported by
the ITS data. In nearly all cases, specimens
assigned to a morphospecies clustered in wellresolved clades supported by high BS and PP
values. The same cannot be said for the ability
of ITS data to resolve infrageneric groups. Of
the six sections of Marasmius recognized in
this study, none are monophyletic. It should be
noted, however, that sect. Leveilleani is
monotypic (M. leveilleanus is the only species)
and we included only one representative each
of sects. Hygrometrici and Neosessiles so
monophylly of these groups could not be tested.
Nonetheless, the diverse sects. Marasmius,
Sicci and Globulares are not monophyletic in
these analyses. The addition of sequences from
other genes (nLSU, mtSSU, rpb1, rpb2) will be
necessary to clearly delimit infrageneric groups
within Marasmius. Until those data become
available, the infrageneric delimitations as
circumscribed by Singer (1976, 1986) and
Desjardin et al. (2005) will be recognized.
All species of Marasmius with Siccustype broom cells or Globulares-type pileipellis
cells in combination with a non-insititious stipe
and the absence of a collarium (viz., sects.
Sicci and Globulares) formed a monophyletic
group with limited statistical support (Fig. 2).
The type species of sect. Sicci, M. siccus
(Schwein.) Fr., is included in the analyses. The
type species of sect. Globulares, M. wynneae
Berk. & Broome, is not included although a
common European and North American
representative, M. oreades (Bolton: Fr.) Fr., is
included. The data suggest that the presence of
Siccus-type broom cells versus Globularestype cells forming the pileipellis tissue, which
is the defining character distinguishing sect.
Sicci from sect. Globulares, is not phylogenetically significant. This observation is not
surprising in that a number of species
traditionally accepted in sect. Sicci have both
Siccus-type broom cells and Globulares-type
18
cells forming the pileipelles. Desjardin and
Horak (1997) recognized this when they
described M. nexus from material collected in
Papua New Guinea and noted that the species
was morphologically intermediate between
these two sections (and hence their use of the
epithet nexus). The data support recognition of
a single infrageneric group to accommodate
species currently accepted in sects. Sicci and
Globulares.
Section Sicci was subdivided into two
subsections by Singer (1976) based on whether
the pileus tramal tissues are inamyloid (subsect.
Inaequales) or dextrinoid (subsect. Siccini).
Subsect. Inaequales is monotypic, and the
single species, M. inaequalis Berk. & M.A.
Curtis, is known only from the New World
(Caribbean and South America) and is not
included in these analyses. Subsect. Siccini is
further subdivided into series based on the
presence or absence of pleurocystidia, setae
and cylindrical (non-setoid) caulocystidia.
Currently recognized series include: ser.
Spinulosi, with setae on the pileus, lamellae
and/or stipe (type – M. cohaerens (Pers.: Fr)
Cooke & Quél.); ser. Atrorubentes, with
cylindrical, non-setoid cystidia on stipe (type –
M. atrorubens (Berk.) Berk.); ser. Leonini,
without pleurocystidia (type – M. leoninus
Berk.); and ser. Haematocephali, with
pleurocystidia (type – M. haematocephalus
(Mont.) Fr.). The ITS data indicate that ser.
Atrorubentes and Leonini are not monophyletic.
Representatives of ser. Atrorubentes, viz., M.
ochropoides, M. musicolor and M. iras, do not
form a single clade, although the latter two
species are monophyletic with 91% BS and 1.0
PP values. Hence, the presence of cylindrical,
non-setoid caulocystidia plus absence of
Siccus-type caulocystidia are not phylogenetically informative features. The monophylly of
ser. Spinulosi, represented in Malaysia by only
one species, M. nummularioides, could not be
tested. We were unable to obtain quality
sequences of M. nummularioides because of
multiple different ITS copies in the specimens
studied; therefore, no data on the phylogenetic
placement of this species is available. The type
species of ser. Spinulosi, M. cohaerens, was
sister of all other members of sect. Sicci with
66% BS and 0.98 PP values. The three
members of ser. Haematocephali included in
Fungal Diversity
the analyses, viz., M. siccus, M. distantifolius
and M. haematocephalus (type species of the
series), did cluster in one clade but without
statistical support. It is possible that the
presence of pleurocystidia is a phylogenetically
informative feature but more data are needed
before this supposition is supported. Representatives of ser. Leonini are scattered through
the sects. Sicci + Globulares clade, suggesting
that the absence of pleurocystidia alone is not
phylogenetically informative.
All species of Marasmius that produce a
collarium to which the lamellae adhere (viz.,
sect. Marasmius) do not form a monophyletic
clade in the ITS analyses (Fig. 2). Historically,
sect. Marasmius has been circumscribed to
include species with an insititious, wiry stipe
and lamellae attached to a collarium. It is
subdivided into subsect. Marasmius that
includes species with Rotalis-type broom cells,
and subsect. Sicciformes that includes species
with Siccus-type broom cells. In the ITS
dataset, only subsect. Marasmius, here represented by the type species of Marasmius (M.
rotula (Scop.: Fr.) Fr.) and three additional
Malaysian species, is monophyletic with strong
support (96% BS, 1.0 PP). In comparison,
members of subsect. Sicciformes form four
distinct clades that do not share a common
ancestor. Interestingly, sect. Hygrometrici that
shares the features of an insititious, wiry stipe
and Rotalis-type broom cells with sect.
Marasmius subsect. Marasmius is not sister of
the latter group but instead is sister of two
species in subsect. Sicciformes that form
Siccus-type broom cells. This was a surprising
result, although it should be noted that there is
no statistical support for this topology and the
nodes delimiting the various clades containing
members of subsect. Sicciformes and sects.
Hygrometrici, Neosessiles and Leveilleani
collapse in a consensus tree.
Although there is no statistical support
for definitive conclusions regarding infrageneric circumscriptions and relationships, it is
interesting that M. tenuissimus, the only
representative of sect. Neosessiles (a group
defined by the absence of a stipe or the
presence of a rudimentary, lateral stipe in
combination with Siccus-type broom cells) is
sister of two members of subsect. Sicciformes
(species with a collarium) and not sister of or
within sect. Sicci (species lacking a collarium).
Based on its overall basidiome size and lack of
a collarium on those basidiomes that form a
rudimenary stipe it was expected to have a
close relationship with sect. Sicci. The
morphology of the Siccus-type broom cells
developed by M. tenuissimus (with short,
knobby or forked setulae) are similar to those
formed by M. ruforotula and M. curreyi, and
distinct from those formed by most members of
sect. Sicci (with long, acutely conical setulae),
thereby supporting the putative relationship
indicated in Fig. 2. Marasmius leveilleanus, the
only member of sect. Leveilleani, also shares
this distinctive form of Siccus-type broom cells
in combination with a wiry, insititious stipe, so
it is not surprising that it resolves sister of the
clade containing M. tenuissimus, M. ruforotula
and M. curreyi (but again with no statistical
support). Whether any of these suggested but
unsupported relationships are resolved in
phylogenetic analyses based on multiple loci
datasets remains to be seen.
Basidiospore size, another useful taxonomic feature, is apparently not phylogenetically
significant beyond informing of closely related
species. For an example in sect. Sicci, M.
aratus and M. aciebrunneus, species with
basidiospore mean length between 25-27 µm
form a clade supported by 94% BS and 1.0 PP
values, but they are not sister of M. adhaesus, a
species with basidiospore mean length of 26
µm. Rather M. adhaesus is sister of M.
bambusiniformis (with 71% BS and 0.98 PP
support), a species with basidiospore mean
length of 17 µm.
In summary, the ITS data support the
recognition of morphospecies but do not
support the current infrageneric delimitation of
Marasmius although they do help focus the
direction of further research
Synopsis of Marasmius sensu stricto of
Peninsular Malaysia
Sect. Marasmius, subsect. Marasmius
1. Marasmius tubulatus Petch
2. Marasmius somalomoensis Antonín
3. Marasmius
diminutivus
Y.S.
Desjardin & Vikineswary, sp. nov.
4. Marasmius leucorotalis Singer
Sect. Marasmius, subsect. Sicciformes
Tan,
19
5.
6.
Marasmius purpureisetosus Corner
Marasmius berambutanus Desjardin, Retn.
& E. Horak
7. Marasmius brevicollus Corner
8. Marasmius purpureobrunneolus Henn.
9. Marasmius guyanensis Mont.
10. Marasmius crinis-equi F. Muell. ex
Kalchbr.
11. Marasmius kuthubutheenii Y.S. Tan,
Desjardin, Vikineswary & Noorlidah, sp.
nov.
12. Marasmius gracilichorda Corner
13. Marasmius ruforotula Singer
Sect. Globulares
14. Marasmius aff. camerunensis Antonín &
Mossebo
15. Marasmius purpureostriatus Hongo
Sect. Neosessiles
16. Marasmius tenuissimus (Jungh.) Singer
Sect. Hygrometrici
17. Marasmius micraster Petch
Sect. Leveilleani
18. Marasmius leveilleanus (Berk.) Pat.
Sect. Sicci, subsect. Siccini, ser. Spinulosi
19. Marasmius nummularioides Desjardin &
Y.S. Tan
Sect.Sicci, subsect. Siccini, ser. Atrorubentes
20. Marasmius
musicolor
Y.S.Tan
&
Desjardin, sp. nov.
21. Marasmius ochropoides Y.S. Tan &
Desjardin, sp. nov.
22. Marasmius iras Y.S. Tan & Desjardin, sp.
nov.
Sect. Sicci, subsect. Siccini, ser. Leonini
23. Marasmius aff. leoninus Berk.
24. Marasmius araucariae var. siccipes
Desjardin, Retn. & E. Horak
25. Marasmius luteomarginatus Desjardin,
Retn. & E. Horak
26. Marasmius jasminodorus Wannathes,
Desjardin & Lumyong
27. Marasmius angustilamellatus Y.S. Tan &
Desjardin, sp. nov.
28. Marasmius olivascens Y.S. Tan &
Desjardin, sp. nov.
29. Marasmius florideus Berk. & Broome
30. Marasmius acerosus Y.S. Tan & Desjardin
31. Marasmius bambusiniformis Singer
32. Marasmius
abundans
Corner
var.
abundans
33. Marasmius abundans var. aurantiacus
Corner
20
34. Marasmius abundans var. campanulatus
Corner
35. Marasmius persicinus Desjardin, Retn. &
E. Horak
36. Marasmius corneri Wannathes, Desjardin
& Lumyong
37. Marasmius selangorensis Y.S. Tan &
Desjardin
38. Marasmius kanchingensis Y.S. Tan &
Desjardin, sp. nov.
39. Marasmius aratus Massee
40. Marasmius aciebrunneus Corner
41. Marasmius adhaesus Corner
Sect. Sicci, subsect. Siccini, ser. Haematocephali
42. Marasmius distantifolius Y.S. Tan &
Desjardin, sp. nov.
43. Marasmius haematocephalus (Mont.) Fr.
Artificial Key to Marasmius of Peninsular
Malaysia
1.
Collarium present, stipe insititious..........................
................................................ (sect. Marasmius) 2
1.* Collarium absent, stipe insititious or non-insititious
............................................................................ 14
2.
Pileipellis composed of Rotalis-type broom cells
........................................... (subsect. Marasmius) 3
2.* Pileipellis composed of Siccus-type broom cells
........................................... (subsect Sicciformes) 6
3.
Pileus grayish orange, yellowish brown or light
brown, lacking a dark papilla ................................ 4
3.* Pileus beige, pale yellow, or white, with dark
papilla.................................................................... 5
4.
Pileus 4-12 mm diam; lamellae distant (10-13);
basidiospore mean 9.3 × 4 µm ....... 1. M. tubulatus
4.* Pileus 2-6 mm diam; lamellae subdistant (11-15);
basidiospore mean 8.8 × 4.3 µm .............................
..............................................2. M. somalomoensis
5.
Pileus < 1mm diam, beige to pale yellow; lamellae
distant (8-10) .............................. 3. M. diminutivus
5.* Pileus 1.5-7 mm diam, white; lamellae subdistant
(10-14)........................................ 4. M. leucorotalis
6.
(from 2) Pileus hispidulous from presence of
pileosetae or very long setulae .............................. 7
6.* Pileus glabrous to minutely pruinose or velutinous;
pileosetae and long setulae absent......................... 8
Pileosetae 16-200+ µm long; pileipellis broom cells
and cheilocystidia with 2-4 apical setulae; tipe
rarely arising from rhizomorphs..............................
.............................................5. M. purpureisetosus
7.* Pileosetae 26-60 µm long; pileipellis broom cells
7.
Fungal Diversity
and cheilocystidia with numerous (2-20) pical
setulae; stipe mostly arising directly from
rhizomorphs............................ 6. M. berambutanus
Basidiospores mean length ≥ 14.7 µm, length range
12-20 µm ............................................................... 9
8.* Basidiospores mean length ≤ 14.2 µm, length range
8-14 (-16) µm ...................................................... 10
8.
9.
Lamellae distant (9-11); pileus with dark papilla;
basidiospore mean 18.8 × 3.7 µm ..........................
......................................................7. M. brevicollus
9.* Lamellae subdistant (10-14); pileus lacking a dark
papilla; basidiospore mean 14.7 × 4.1 µm...............
......................................8. M. purpureobrunneolus
10. Pileus orange, yellowish orange or pale yellow
overall.......................................... 9. M. guyanensis
10.* Pileus brownish orange, grayish orange, light
brown, reddish brown or brown .......................... 11
11. Stipe arising directly from copious black
rhizomorphs................................10. M. crinis-equi
11.* Stipe not arising or rarely arising directly from
rhizomorphs......................................................... 12
12. Pileus 0.5-3 mm diam; lamellae remote to distant
(6-9); stipe 2-4 mm long; basidiomes on bamboo
leaves.................................. 11. M. kuthubutheenii
12.*Pileus 2-8 mm diam; lamellae distant (6-13); stipe 523 mm long; basidiomes on dicotyledonous and
monocotyledonous leaves, including bamboo..... 13
13. Pileipellis with many thick-walled, darkly
pigmented broom cells that have lumpy or warted
setulae 5-12 mm long 12. ............ M. gracilichorda
13.* Pileipellis with thinner-walled broom cells that
have smooth setulae 2-5 µm long............................
.....................................................13. M. ruforotula
14. (from 1) Pileipellis composed of a hymeniform
layer of clavate to pyriform, non- setulose cells
............................................... (sect. Globulares) 15
14.* Pileipellis composed of Rotalis-type or Siccus-type
broom cells .......................................................... 16
15. Pileus grayish brown, brownish orange or grayish
orange; basidiospore mean length ca 28 µm ...........
........................................ 14. M. aff. camerunensis
15.* Pileus purplish gray to grayish violet with
yellowish white radial stripes; basidiospore mean
length ca 25 µm................ 15. M. purpureostriatus
16. Stipe none or rudimentary, lateral to strongly
eccentric, less than 2 mm long ...............................
.................... (sect. Neosessiles) 16. M. tenuissimus
16.* Stipe central, well-developed. ............................. 17
17. Pileipellis composed of Rotalis-type broom cells ...
.................... (sect. Hygrometrici) 17. M. micraster
17.* Pileipellis composed of Siccus-type broom cells 18
18. Lamellae free, well separated from stipe apex; stipe
insititious; coarse black rhizomorphs abundant .....
....................(sect. Leveilleani) 18. M. leveilleanus
18.* Lamellae attached, typically adnexed to adnate;
stipe non-insititious; coarse black rhizomorphs
absent ..............................................(sect. Sicci) 19
19. Caulosetae present; stipitipellis with a combination
of Siccus-type broom cells, setae-like cells with
setulae up to 60 µm long and setae; (ser. Spinulosi)
..........................................19. M. nummularioides
19.* Caulosetae
absent;
stipitipellis
lacking
caulocystidia or with cylindrical non- setulose
caulocystidia or Siccus-type broom cells ............ 20
20. Stipe pruinose overall, prunoisity formed from
irregularly cylindrical, obtuse to subacute, nonsetulose cells, Siccus-type cells absent on stipe ......
............................................. (ser. Atrorubentes) 21
20.* Stipe glabrous or pruinose, if pruinose, then
pruinosity formed from Siccus-type broom cells 23
21. Pileus light yellow (banana colored); basidiospore
mean length 16.1 µm................... 20. M. musicolor
21.* Pileus brownish orange, light brown, reddish
browm or brown; basidiospore mean length 10.614.1 µm ............................................................... 22
22. Stipe 11-21 mm long; cheilocystidia with only 2-4
setulae up to 20 µm long; 10.6 × 3.7 µm ................
.................................................21. M. ochropoides
22.* Stipe 21-45 mm long; cheilocystidia with numerous,
crowded setulae 4-6 µm long; basidiospore mean
14.1 × 3.7 µm ........................................ 22. M. iras
23. Pleurocystidia absent.....................(ser. Leonini) 24
23.* Pleurocystidia present, often refractive and
conspicuous.....................(ser. Haematocephali) 42
24. Stipe with caulocystidia (Siccus-type broom cells
sometimes in combination with small cylindrical to
fusoid, non-setulose cells)................................... 25
24.* Stipe lacking caulocystidia.................................. 28
25. Basidiospores < 12 µm long, mean length range
9.2-11.4 µm......................................................... 26
25.* Basidiospores > 12 µm long, mean length range
12.7-17.9 µm....................................................... 27
26. Pileus pale yellow to pale orange with darker disc;
lamellae with non-pigmented edges........................
................................................ 23. M. aff. leoninus
26.* Pileus brownish orange to light brown with darker
disc; lamellae with brownish orange edges.............
..............................24. M. araucariae var. siccipes
27. Pileus orange to pale orange; lamellae with yellow
edges; stipe < 3.5 mm long, eccentric; odor none;
cheilocystidia irregularly cylindrical, non-setulose.
..........................................25. M. luteomarginatus
27.* Pileus brownish orange to orangish red with darker
disc; lamellae with orangish red edges; stipe > 16
21
mm long, central; odor like jasmine tea;
cheilocystidia of Siccus-type broom cells ...............
............................................... 26. M. jasminodorus
28. Basidiospore length in the range 8-14 µm with
mean length 9.9-12.7 µm .................................... 29
28.* Basidiospore length in the range 12-30 µm with
mean length 14.4-27 µm ..................................... 32
29. Lamellae crowded (24-40), very narrow; pileus 1850 mm diam, yellowish brown to grayish orange or
light brown with a black disc ..................................
........................................ 27. M. angustilamellatus
29.* Lamellae subdistant to distant (10-20); pileus 3-18
mm diam, olive, reddish brown to brown or
cinnamon-ochraceous to pinkish fulvous ............ 30
30. Pileus olive to olive-brown; basidiospore length in
the range 11-14 µm with mean 12.7 µm .................
.....................................................28. M. olivascens
30.* Pileus reddish brown to brown or cinnamonochraceous to pinkish fulvous; basidiospore length
in the range 9.5-12 µm with mean 10.4-10.7 µm 31
31. Pileus reddish brown to brown, 3-6 mm diam,
striate in age; lamellae close to subdistant (10-12).
........................................................ 29 M. florideus
31.* Pileus cinnamon-ochraceous to pinkish fulvous,
typically >10 mm diam, smooth in age; lamellae
close to crowded (17-20).........................................
......................34. M. abundans var. campanulatus
32. Basidiospore length in the range 12-18 µm with
mean length 14.4-16.9 µm .................................. 33
32.* Basidiospore length in the range 16-30 µm with
mean length 17.7-27 µm ..................................... 36
33. Lamellae crowded (16-25 with 3-4 series of
lamellulae); basidiospore mean length 14.4 µm;
setulae on cheilocystidia and pileipellis broom cells
up to 13 µm long ........................... 30. M. acerosus
33.* Lamellae close to subdistant (12-19); basidiospores
mean length 15.4-16.9 µm; setulae on
cheilocystidia and pileipellis broom cells up to 7
µm long ............................................................... 34
34. Lamellae subdistant (12-13); pileus 1.5-5 mm diam,
striate to sulcate in age .....31. M. bambusiniformis
34.* Lamellae close (14-19); pileus 8-30 mm diam,
smooth to striatulate in age.................................. 35
35. Pileus grayish orange to golden yellow, brownish
yellow or ferruginous ..............................................
.............................32. M. abundans var. abundans
35.* Pileus orange to deep orange...................................
......................... 33. M. abundans var. aurantiacus
36. Basidiospore length in the range 16-24 µm with
mean length 17.7-21.8 µm .................................. 37
36.* Basidiospore length in the range 20-30 µm with
mean length 24.8-27 µm ..................................... 40
22
37. Pileus light orange to light pinkish tan or peachcolored................................................................. 38
37.* Pileus more deeply pigmented, brownish yellow,
brownish orange, brown or dark brown .............. 39
38. Pileus 3-10 mm diam; lamellae 8-16; basidiospore
mean length 17.7-18.3 µm ..........35. M. persicinus
38.* Pileus 10-43 mm diam; lamellae 12-18;
basidiospore mean length 19-21.8 µm ....................
.........................................................36. M. corneri
39. Pileus 10-23 mm diam, brown to dark brown with
pinkish tones, gradually becoming yellowish brown
to pinkish brown or golden brown on the margin
with a dark brown disc; basidiospore mean length
20.9 µm ................................. 37. M. selangorensis
39.* Pileus 2-12 mm diam, brown to brownish orange
with pale grayish orange stripes (lacking pinkish
tones); basidiospore mean length 19.5 µm..............
............................................. 38. M. kanchingensis
40. Pileus 4-13 mm diam, blood red to deep brownish
purple; basidiospore mean length 27 µm ................
.......................................................... 39. M. aratus
40.* Pileus often up to 20 mm or more in diameter,
brown, brownish olive or ferruginous; basidiospore
mean length 24.8-25.9 µm .................................. 41
41. Pileus 7-20 mm diam, brown to ferruginous with a
darker disc; lamellae 11-14; basidiospore length
range 20-27 µm with mean length 24.8 µm ............
.............................................. 40. M. aciebrunneus
41.* Pileus 6-40 mm diam, brown to brownish olive
with a darker disc; lamellae 10-20; basidiospore
length range 25-30 with mean length 27.9 µm........
..................................................... 41. M. adhaesus
42. (from 23) Lamellae distant (7-8); basidiospore
mean length 21.4 µm.............. 42. M. distantifolius
42.* Lamellae subdistant (12-13); basidiospore mean
length 18.2 µm ................ 43. M. haematocephalus
Taxonomy
Marasmius Fr. sect. Marasmius, subsect.
Marasmius
= sect. Pararotulae Singer, Sydowia 18: 339.
1965. [Type species: Marasmius pararotula Singer]
= subsect. Pararotulae (Singer) Singer, Fl.
Neotrop. Monogr. 17: 92. 1976.
1. Marasmius tubulatus Petch, Trans. Brit.
Mycol. Soc. 31: 42. 1947. (Fig. 3, Plate 1A)
Type: Sri Lanka, Central Prov., Kandy
District, Peradeniya, 1 Nov. 1914, Petch 4243
(K).
Pileus 4-12 mm diam, broadly conical
and truncate to broadly convex or convex with
a shallowly depressed disc, striate to sulcate;
Fungal Diversity
surface dry, dull, minutely pruinose; grayish
yellow (4B4-5) to grayish orange (5B3-5),
yellowish brown (5D5), light brown (6D5-6) or
brown (6E6-5) with a central black dot.
Lamellae adnate to a collarium, distant (10-13),
yellowish white, non-marginate or more
commonly with light brown marginate edges.
Stipe 10-32 × 0.5 mm, central, terete, equal,
cylindrical, dry, dull, glabrous, insititious,
yellowish white to light brown at apex, dark
brown to dark brown at base.
Basidiospores (7-) 8-10.5 × 3-4.5 (-5) μm,
[xmr = 8.7-10.3 × 3.9-4.1 μm, xmm = 9.3 ± 0.6 ×
4 ± 0.2 μm, Q = 1.7-3.3, Qmr = 2.2-2.7, Qmm =
2.4 ± 0.1, n = 23 spores per 9 collections],
ellipsoid, smooth, hyaline, inamyloid. Basidia
21-27 × 6-7 μm, 2-spored and 4-spored.
Basidioles 20-28 × 5-8 μm, fusoid to clavate.
Cheilocystidia abundant, composed of Rotalistype broom cells; main body 12-27 × 7-18 (-22)
μm, subclavate to clavate or subglobose,
hyaline, inamyloid, thin-walled or apically
thick-walled; divergent setulae 1-3 × 0.5-1 μm,
abundant over upper half of cell, cylindrical to
conical, subacute to obtuse, light yellowish
brown to light brown, weakly dextrinoid.
Pleurocystidia absent. Pileipellis mottled,
composed of a hymeniform layer of Rotalistype broom cells; main body 10-35 (-41) ×
(7-)10-26 μm, cylindrical to clavate, subclavate,
turbinate or subglobose, rarely lobed, hyaline
to light yellow, inamyloid, thin-walled or
apically thick-walled; divergent setulae 1-4 ×
0.5-1 μm, abundant over upper half of cell,
cylindrical to conical, subacute to obtuse,
brownish yellow to brown, inamyloid, thinwalled. Pileus trama interwoven; hyphae 3-6
μm diam, hyaline, inamyloid to weakly dextrinoid, thin-walled, non-gelatinous. Lamellar
trama regular to subparallel; hyphae 4-8 μm
diam, cylindrical, hyaline, dextrinoid, thinwalled, non-gelatinous. Stipe tissue monomitic;
cortical hyphae 5-10 μm diam, parallel,
cylindrical, smooth, yellowish brown to brown,
weakly dextrinoid, thick-walled (up to 1 μm),
non-gelatinous; medullary hyphae 4-8 μm diam,
parallel, cylindrical, smooth, hyaline, inamyloid, thin-walled, non-gelatinous. Caulocystidia
absent. Clamp connections present.
Habit, habitat and known distribution:
gregarious on leaves of undetermined
dicotyledonous trees. Malaysia (Johore), Sri
Lanka.
Material examined: Malaysia, Johore, Endau
Rompin National Park, way to Dato’s Ghani Trail +
Kuala Jasin, 13 Jul. 2005, Yee-Shin Tan, TYS 492
(KLU-M#109, SFSU); same location, Yee-Shin Tan,
TYS 490 (KLU-M#105, SFSU); same location, along
the way to NERC, Peta Village and Temiang Trail, 14
Jul. 2005, Yee-Shin Tan, TYS 502 (KLU-M #17,
SFSU); same location, 14 Jul. 2005, Yee-Shin Tan,
TYS504 (KLU-M #18, SFSU); same location, 14 Jul.
2005, Yee-Shin Tan, TYS 505 (KLU-M #21, SFSU);
same location, 14 Jul. 2005, Yee-Shin Tan, TYS 510
(KLU-M #19, SFSU); same location, 14 Jul. 2005, YeeShin Tan, TYS 512 (KLU-M #20, SFSU); same location,
14 Jul. 2005, Yee-Shin Tan, TYS 518 (KLU-M #22,
SFSU).
Notes: Marasmius tubulatus is characterized by a light brown or pale yellowish
brown pileus with a dark central spot, distant
lamellae (10-13) that are non-marginate or
have light brown edges, relatively small
basidiospores in the range 8-10 × 3-4.5 μm
with mean of 9.3 × 4 μm, broad, light brown
Rotalis-type broom cells on the pileipellis and
lamellar edge, and by growth on dicotyledonous leaves. The material from Malaysia
differs only subtly from the protologue (Petch,
1947) and redescription (Pegler, 1986) of Sri
Lanka specimens, in forming slightly larger
pilei (up to 12 mm diam) that are more evenly
pigmented and lack a distinct white margin. A
tetrapolar (bifactorial) mating system was
reported for M. tubulatus by Tan et al. (2007)
based on Malaysian material.
2. Marasmius somalomoensis Antonín,
Mycotaxon 88: 66. 2003.
(Fig. 4)
Type:
Cameroon,
Pud
Province,
Somalomo, Dja Biospher Reserve, 8 Apr. 2001.
(BRNM 666108)
Pileus 2-6 mm diam, convex, truncately
convex with a shallowly depressed disc, no
papilla, striate to sulcate; surface dry, dull,
minutely pruinose; yellowish white to light
brownish white, golden brown (5E7), grayish
orange (6B5), or light brown (6D6), with a
central dark brown or black dot. Lamellae
adnate to a collarium, subdistant (11-15),
yellowish white with light brown to brown
marginate edges. Stipe 10-15 × 0.5 mm, central,
terete, equal, cylindrical, dry, dull, glabrous,
insititious, yellowish white at apex, dark brown
at base.
23
Plate 1. Marasmius s.s. in section Marasmius, subsection Marasmius and subsection Sicciformes. A: Marasmius
tubulatus (x 1); B: Marasmius leucorotalis (x 1); C: Marasmius purpureisetosus (x 1); D: Marasmius
purpureobrunneolus (x 2); E: Marasmius guyanensis (x 1); F: Marasmius kuthubutheenii (x 3); G: Marasmius
gracilichorda (x 2); H: Marasmius ruforotula (x 2).
24
Fungal Diversity
Fig. 3. Marasmius tubulatus (TYS 502 = KLU-M#17). a. Basidiomes. b. Basidiospores. c. Basidia and basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a= 10 mm, b-e=10 µm.
Fig. 4. Marasmius somalomoensis (TYS 330 = KLU-M#107). a. Basidiomes. b. Basidiospores. c. Basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e =10 µm.
25
Basidiospores (7-) 8-10 × 3-4 (-5) μm,
[xmr = 8.1-9.2 × 3.6-4.8 μm, xmm = 8.8 ± 0.5 ×
4.3 ± 0.5 μm, Q = 1.7-3.3, Qmr = 1.9-2.3, Qmm =
2.1 ± 0.2, n = 20 spores per 4 specimens],
ellipsoid, smooth, hyaline, inamyloid. Basidia
18-23 × 4-7 μm, 4-spored. Basidioles 22-25 ×
5-7 μm, fusoid to clavate. Cheilocystidia
abundant, composed of Rotalis-type broom
cells; main body (12-) 13-20 (-22) × (8-) 10-14
(-15) μm, subclavate to clavate or subglobose,
rarely lobed, hyaline, inamyloid, thin-walled or
apically thick-walled; divergent setulae 2-3 ×
0.5-1 μm, abundant over upper half of cell,
cylindrical to conical, subacute to obtuse, light
yellow to light brown, inamyloid. Pleurocystidia absent. Pileipellis mottled, composed
of a hymeniform layer of Rotalis-type broom
cells; main body (15-) 17-28 (-34) × (10-) 1220 (-22) μm, cylindrical to clavate, subclavate,
turbinate or subglobose, sometimes lobed, light
yellow to light brown, inamyloid, thin-walled
or apically thick-walled; divergent setulae 2-3
× 0.5-1 μm, abundant over upper half of cell,
cylindrical to conical, subacute to obtuse,
yellowish brown to brown, inamyloid, thinwalled. Pileus trama interwoven; hyphae 3.5-5
μm diam, hyaline, inamyloid, thick-walled,
non-gelatinous. Lamellar trama regular to
subparallel; hyphae 3-5 μm diam, cylindrical,
hyaline, inamyloid, thin-walled, non-gelatinous.
Stipe tissue monomitic; cortical hyphae 4-6 μm
diam, parallel, cylindrical, smooth, brown,
weakly dextrinoid, thick-walled (up to 1 μm),
non-gelatinous; medullary hyphae 4-7 μm diam,
parallel, cylindrical, smooth, light yellow,
inamyloid, thin-walled, non-gelatinous. Caulocystidia absent. Clamp connections present.
Habit, habitat and known distribution:
gregarious on leaves of undetermined dicotyledonous trees. Africa (Cameroon, DR Congo),
Malaysia (Kedah), Thailand.
Material examined: Malaysia, Kedah, Langkawi
Island, Mt. Raya, N 6°25’ 52.8, E 99° 49’ 26.5, elev. 652
m, 6 Apr. 2004, Yee-Shin Tan, KUM 60023 (KLUM#106, SFSU); same location, Matchinchang Forest
Reserve, 30 Aug. 2004, Yee-Shin Tan, TYS 330 (KLUM#107, SFSU); same location, 1 Sept. 2004, Yee-Shin
Tan, TYS 335 (KLU-M#108, SFSU); same location, 1
Sept. 2004, Yee-Shin Tan, TYS 336 (KLU-M#130,
SFSU).
Notes: Marasmius somalomoensis was
described recently from material collected in
Africa. It is distinguished by light brown to
26
brownish orange, sulcate-depressed pilei 1-5
mm diam, 11-15 collariate lamellae with pale
brown edges, basidiospores in the range 7-10 x
3.7-5.5 µm, Rotalis-type cheilocystidia (14-30
× 8-18 µm) and pileipellis broom cells (18-33
× 10-19 µm), and growth on dicotelydonous
leaves (Antonín, 2007). The Malaysian
specimens differ from the protologue of the
African material only in having weakly
marginate lamellar edges. The species is
similar to M. apatelius Singer, but this has
basidiospores with mean 7.2 ×3.9 µm and nonmarginate lamellae. It is also similar to M.
tubulatus Petch, but the latter forms larger pilei
(4-12 mm diam), fewer (10-13) and broader
lamellae and slightly narrower basidiospores
(mean width 4.0 µm).
3. Marasmius diminutivus Y.S. Tan,
Desjardin & Vikineswary, sp. nov.
(Fig. 5)
MycoBank: MB512624
Etymology: deminutio (Latin) = to make small,
diminutive; referring to the tiny basidiomes.
Pileus 0.5-1 mm latus, conico-umbilicatus, papilla
atrobrunnea conica instructus, striatus vel sulcatus,
minutae pruinosus, stramineus vel aurantio-griseus.
Lamellae adnato-collariatae, distantes (8-10), stramineae,
haud marginatae. Stipes 6-17 × 0.1 mm, filiformis,
glabrus, insititius, apicaliter stramineus, basim rubrobrunneus. Basidiosporae 7-9 × 3-4.5 μm, ellipsoideae,
leves, hyalinae, inamyloideae. Basidiola 18-20 × 4.5-5
μm, fusiodeae vel clavatae. Cheilocystidia rara, typi
Rotalis, 12-18 ×11-14 μm, subclavata vel clavata,
hyalina; setulae divergentes 1.5-2 × 1 μm, cylindricae
vel conicae, obtusae, stramineae. Pleurocystidia nulla.
Pileipellis hymeniformis, typi Rotalis, cellulae 19-25 ×
12-19 μm, subclavatae vel turbinatae, hyalinae,
inamyloideae; setulae divergentes 2 × 0,5-1 μm,
cylindricae
vel
conicae,
obtusae,
stramineae.
Caulocystidia nulla. Fibulae presentes. Gregarius, ad
folia putrida plantarum dicotyledonearum. Holotypus:
Malaysia, Johore, Endau Rompin National Park, 14 Jul.
2005, Yee-Shin Tan, TYS 514 (Holotypus: KLU-M#66).
Pileus 0.5-1 mm diam, obtusely conical
with narrowly depressed to umbilicate disc,
with a tiny, acute papilla in the umbilicus;
margin striate to sulcate; surface dry, dull,
minutely pruinose; beige to pale yellow or
grayish orange (5B3-4) with dark brown
papilla. Lamellae adnate to a collarium, distant
to subdistant (8-10), yellowish white, nonmarginate. Stipe 6-17 × 0.1 mm, central, terete,
equal, filiform, dry, dull, glabrous, insititious,
sometimes with a few scattered nodes, orangish
white at apex, reddish brown to brown at base.
Fungal Diversity
Fig. 5. Marasmius diminutivus (Holotype: TYS 514 = KLU-M#66). a. Basidiomes. b. Basidiospores. c. Basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a = 3 mm, b-e =10 µm.
Basidiospores (6-) 7-10 × 3-4.5 μm, [xmr
= 8-9.3 × 3.5-3.9 μm, xmm = 8.9 ± 0.8 × 3.7 ±
0.2 μm, Q = 1.8-2.7, Qmr = 2.3-2.5, Qmm = 2.4
± 0.1, n = 18 spores per 3 specimens], ellipsoid,
smooth, hyaline, inamyloid. Basidia not
observed. Basidioles 18-20 × 4.5-5 μm, fusoid
to clavate. Cheilocystidia rare, composed of
Rotalis-type broom cells; main body (10-) 1218 (-25) × (7-) 11-14 μm, subclavate to clavate
or subglobose, lobed, hyaline, inamyloid, thinwalled or apically thick-walled; divergent
setulae 1.5-2 × 1 μm, abundant over upper half
of cell, cylindrical to conical, obtuse, light
yellow, inamyloid, thin-walled. Pleurocystidia
absent. Pileipellis weakly mottled, composed
of a hymeniform layer of Rotalis-type broom
cells; main body (15-) 19-25 × (9-) 12-19 μm,
subclavate to turbinate or subglobose, lobed,
hyaline, inamyloid, thin-walled or apically
thick-walled; divergent setulae 2 × 0.5-1 μm,
abundant over upper half of cell, cylindrical to
conical, obtuse, light yellow, inamyloid, thinwalled. Pileus trama interwoven; hyphae 3-4
μm diam, hyaline to light yellow, dextrinoid,
thin-walled, non-gelatinous. Lamellar trama
subparallel; hyphae 3.5-4 μm diam, cylindrical,
hyaline, inamyloid, thin-walled, non-gelatinous.
Stipe tissue monomitic; cortical hyphae 3.5-5
μm diam, parallel, cylindrical, smooth,
yellowish brown, weakly dextrinoid, thickwalled (1 μm), non-gelatinous; medullary
hyphae 2.5-4.5 μm diam, parallel, cylindrical,
smooth, hyaline, inamyloid to weakly
dextrinoid, thin-walled, non-gelatinous. Caulocystidia absent. Clamp connections present.
Habit, habitat and known distribution:
gregarious on leaves of undetermined
dicotyledonous trees. Malaysia (Johore).
27
Material examined: Malaysia, Johore, Endau
Rompin National Park, 14 Jul. 2005, Yee-Shin Tan, TYS
514 (Holotype: KLU-M#66; Isotype: SFSU); Selangor,
Sungai Chongkak Forest Reserve, N 03° 12.705’, E 101°
50.472’, elev. 188m, 7 Jan. 2005, Yee-Shin Tan, TYS
403 (KLU-M#67, SFSU); same location, 16 May 2005,
Yee-Shin Tan, TYS 473 (KLU-M#68, SFSU).
Notes: Marasmius diminutivus is
characterized by the following features: a
pileus less than 1 mm diam that is beige to pale
yellow with a tiny dark brown papilla; 8-10
collariate lamellae; a wiry stipe 6-17 × 0.1 mm
with a few nodes; basidiospores 7-10 × 3-4.5
µm with mean 8.9 × 3.7 µm; Rotalis-type
cheilocystidia and pileipellis broom cells; and
growth on dicotyledonous leaves. It is the
smallest Marasmius species known from
Malaysia and unlike any species described
from Malesia. Marasmius capillaris Morgan,
from eastern North America, differs in forming
much larger pilei (2-7 mm diam.), more
numerous lamellae (13-18), and a much longer
stipe (15-45 mm) (Holotype: ISC!; Desjardin,
1989).
4. Marasmius leucorotalis Singer, Sydowia
18: 337. 1965.
(Fig. 6, Plate 1 B)
Type: Bolivia, La Paz; Nor-Yungas, Rio
Yariza, 23 Feb 1956, Singer B 1414 (LIL).
Pileus 1.5-7 mm diam, convex to broadly
convex with an umbilicate disc and tiny papilla;
margin sulcate; surface dry, dull, minutely
pruinose; white with dark brown papilla.
Lamellae adnate to a collarium, subdistant (1014), white, non-marginate. Stipe 8-35 × 0.5 mm,
central, terete, equal, cylindrical, dry, dull,
glabrous, insititious, beige at apex, dark brown
to black at base.
Basidiospores (5-) 6-7 (-7.5) × (2-) 2.5-3
(3.5) μm, [x = 6.6 ± 0.7 × 2.8 ± 0.5 μm, Q = 23.5, Qm = 2.4 ± 0.4, n = 25 spores per 2
specimens], ellipsoid, smooth, hyaline,
inamyloid. Basidia not observed. Basidioles
18-25 × 5-8 μm, fusoid to clavate.
Cheilocystidia abundant, composed of Rotalistype broom cells; main body 14-20 (-26) × (9-)
10-16 (-22) μm, subclavate to clavate or
subglobose, rarely lobed, hyaline, inamyloid,
thin-walled or apically thick-walled; divergent
setulae 1-2 × 0.5-1 μm, abundant over upper
28
half of cell, cylindrical to conical, obtuse, light
yellow, inamyloid, thin-walled. Pleurocystidia
absent. Pileipellis weakly mottled, composed
of a hymeniform layer of Rotalis-type broom
cells; main body (14-) 16-29 (-37) × 10-17 (-21)
μm, subclavate to turbinate or subglobose,
rarely lobed, hyaline, inamyloid, thin-walled or
apically thick-walled; divergent setulae 1.5-2 (2.5) × 0.5-1 μm, abundant over upper half of
cell, cylindrical to conical, subacute to obtuse,
light yellow, inamyloid, thin-walled. Pileus
trama interwoven; hyphae 3-4 μm diam,
hyaline, inamyloid, thin-walled, non-gelatinous.
Lamellar trama regular; hyphae 4-5 μm diam,
cylindrical, hyaline, inamyloid, thin-walled,
non-gelatinous. Stipe tissue monomitic; cortical
hyphae 4-6 μm diam, parallel, cylindrical,
smooth, yellowish brown to brown, weakly
dextrinoid, thick-walled (1-2 μm), nongelatinous; medullary hyphae 4-8 μm diam,
parallel, cylindrical, smooth, hyaline, inamyloid, thin-walled, non-gelatinous. Caulocystidia
absent. Clamp connections present.
Habit, habitat and known distribution:
gregarious on leaves of undetermined dicotyledonous trees. Indonesia (Java), Malaysia
(Johore), South America (Bolivia).
Material examined: Malaysia, Johore, Endau
Rompin National Park, Dato’s Ghani Trail and trail to
Upeh Geling, 12 Jul. 2005, Yee-Shin Tan, TYS 486
(SFSU). Malaysia, Johore, Endau Rompin National Park,
Kuala Jasin, 13 Jul. 2005, Yee-Shin Tan, TYS 489
(KLU-M#126, SFSU).
Notes: The Malaysian material differs
slightly from the Indonesian populations in
forming shorter and narrower basidiospores
with mean 6.6 × 2.8 µm versus 8.4 × 3.7 µm
(Desjardin et al. 2000).
Marasmius sect. Marasmius, subsect.
Sicciformes Antonín, Acta Moraviae, Sci. Nat.
76: 145. 1991.
5. Marasmius purpureisetosus Corner, Beih.
Nova Hedwigia 111: 90. 1996.
(Fig. 7, Plate 1C)
Type: Malaysia, Johore, Malayan
Peninsula, Corner s.n., 1 Sept. 1940
(E#206731!).
Fungal Diversity
Fig. 6. Marasmius leucorotalis (TYS 486). a. Basidiomes. b. Basidiospores. c. Basidioles. d. Cheilocystidia. e.
Pileipellis. Bars: a = 5 mm, b-e =10 µm.
Pileus 0.5-2.5 mm diameter, obtusely
conical or conical to convex, with central
umbilicus and a small, acute papilla; margin
smooth to striate; surface dry, dull, hispid
overall with long and brown setae, background
colour yellowish brown to grayish orange
(5B4), with a pale yellow or light yellow
umbilicus. Lamellae adnate to a collarium,
subdistant to close (7-18), yellowish white,
non-marginate. Stipe 3.5-10 × 0.1 mm,
cylindrical, central, terete, wiry, dry, dull,
glabrous, insititious, yellowish brown to dark
brown, black rhizomorphs present, some stipes
arising from rhizomorphs.
Basidiospores (7-) 8-14 × 3-5 µm [xmr =
8.6-12 × 3.5-4.8 μm, xmm = 9.8 ± 1.4 × 4.1 ±
0.5 μm, Q = 1.9-3.3, Qmr = 2.2-2.6, Qmm = 2.4
± 0.2, n = 22-35 spores per 7 specimens],
ellipsoid, hyaline, smooth, inamyloid. Basidia
20-28 × 4-6 µm, clavate, 4-spored. Basidioles
21-25 × 5-7 µm, clavate to fusoid.
Pleurocystidia absent. Cheilocystidia numerous,
composed of 2 types of cells: a) Siccus-type
broom cells; main body 8-22 × 6-15 µm,
clavate to broadly clavate, cylindrical or
irregular in outline, rarely lobed, hyaline,
inamyloid, thick-walled; apical setulae 3-12 ×
2-4 µm, only 2-4 per cell, cylindrical to conical,
subacute to obtuse, light yellow, inamyloid,
thick-walled; b) non-setulose cells, 23-46 × 810 µm, cylindrical to clavate or subclavate,
hyaline, inamyloid, thick-walled (-1 µm).
Pileipellis mottled, composed of a hymeniform
layer of 2 types of cells and pileosetae: a)
29
Fig. 7. Marasmius purpureisetosus (KUM 60018 = KLU-M#114). a. Basidiomes. b. Basidiospores. c. Basidia and
basidioles. d. Cheilocystidia. e. Pileipellis. f. Pileosetae. Bars: a = 10 mm, b-e =10 µm.
Siccus-type broom cells; main body 7-28 × 613 µm, turbinate to clavate, broadly clavate or
irregular in outline, hyaline, inamyloid, thickwalled; apical setulae 6-20 × 1-4 µm, only 2-4
per cell, cylindrical to conical, rarely forked,
light yellow, dextrinoid, thick-walled, most
cells have long setae; b) transitional elements,
15-55 × 7-12 µm, cylindrical to clavate, light
yellow, dextrinoid, thick-walled with 2-3 apical
setulae 8-41 × 4-8 µm, cylindrical to conical,
subacute, light brown, dextrinoid, thick-walled;
c) pileosetae 16-200+ × 3-13 µm, brown,
dextrinoid, thick-walled (1-3 µm broad). Pileus
trama interwoven; hyphae 7-9 µm diam,
hyaline, inamyloid, non-gelatinous. Lamellar
trama regular; hyphae 2-6 µm diam, cylindrical,
30
hyaline, dextrinoid, thin-walled. Stipe tissue
monomitic; cortical hyphae 3.5-6 µm diam,
parallel, cylindrical, smooth, brownish yellow,
weakly dextrinoid, thick-walled (0.5-1 µm),
non-gelatinous; medullary hyphae 3-5 µm diam,
parallel, cylindrical, smooth, hyaline, inamyloid, thin-walled, gelatinous. Caulocystidia
absent. Clamp connections present.
Habit, habitat and known distribution:
gregarious on leaves. Indonesia (Java),
Malaysia (Kedah, Negeri Sembilan, Pahang,
Selangor), Singapore, Thailand.
Material examined: Malaysia, Kedah, Langkawi
Island, Gunung Raya, N 06°22’42.3, E 99° 49’21.5, 640
m elev., 6 Apr. 2004, Yee-Shin Tan, KUM 60018 (KLUM#114, SFSU); same location, 6 Apr. 2004, Yee-Shin
Tan, KUM 60020 (KLU-M#115, SFSU); same location,
Fungal Diversity
6 Apr. 2004, Yee-Shin Tan, KUM 60022 (KLU-M#116,
SFSU); Langkawi Island, Lubuk Semilang Forest
Reserve, N 06° 21’58.4, E 99° 47’29.6, 59m elev., 8 Apr.
2004, Yee-Shin Tan, KUM 60143 (KLU-M#117, SFSU);
Negeri Sembilan, Ulu Bendul Forest Reserve, 23 Oct.
2004, Yee-Shin Tan, TYS 371 (KLU-M#119, SFSU);
Pahang, Cameron Highlands, Tanah Rata, Jungle walk
no 9 & 9A, 9 Jan. 2004, Yee-Shin Tan, TYS 224 (KLUM#118, SFSU); Selangor, Kanching Forest Reserve, N
03° 17.958’, E 101° 31.15’, 110 m elev., 9 Jan. 2005,
Yee-Shin Tan, TYS 432 (KLU-M#120, SFSU).
Singapore, Johore, 1 Sept. 1940, Corner (Holotype
E#206731).
Notes: Basidiomes of M. purpureisetosus
have the following features: a small, hispid,
yellowish brown to grayish orange or pale
yellow pileus covered with dark brown to
purplish brown pileosetae, subdistant to close
(7-18) lamellae attached to a collarium, and
glabrous stipes attached to leafy substrate or
directly to rhizomorphs. Our Malaysian
material matches quite closely with Corner’s
protologue and the holotype specimen except
for the absence of purple tones on the pilei. In
addition, the Malaysian specimen differs from
Indonesian material in having narrower
lamellae and slightly larger basidiospore mean
(9.8 × 4.1 μm versus 7.9 × 3.7 µm in Java
material; Desjardin et al., 2000).
6. Marasmius berambutanus Desjardin,
Retn. & E. Horak, Sydowia 52: 116. 2000.
(Fig. 8)
Type: Indonesia, Java, West Java, Mt.
Halimun National Park, loop trail from
Cikaniki, 6 Jan. 1999, A. Retnowati 081 (BO!).
Pileus 1-6 mm diam, paraboloid to
obtusely conical, convex or broadly convex,
umbilicate, with an acute central papilla,
sulcate to plicate to the umbilicus; surface dry,
dull, minutely velutinous; grayish yellow (4B5)
to grayish orange (5B4-5), pale orange or
brownish orange (5C5). Lamellae adnate to a
collarium, distant (7-12) with no lamellulae,
broad, pale yellowish white (< 4A2) with
grayish orange marginate edges. Stipe 3-24 ×
0.1 mm, central, terete, equal, filiform, dry,
dull, glabrous, insititious, yellowish white at
apex, dark brown at base, arising directly from
black rhizomorphs; rhizomorphs often with
scattered nodes.
Basidiospores (7-) 8-11 (-12) × 3-5 (-5.5)
μm [xmr = 8.2-9.5 × 3.3-4.5 μm, xmm = 9.1 ± 0.8
× 4.0 ± 0.5 μm, Q = 1.5-3.3, Qmr = 2.0-2.5,
Qmm = 2.3 ± 0.3, n= 30 spores per 12
collections], ellipsoid, smooth, hyaline,
inamyloid. Basidia 22-30 × 5-7 μm, clavate, 4spored. Basidioles 18-26 × 5-7 μm, fusoid to
clavate. Cheilocystidia abundant, composed of
Siccus-type broom cells; main body 10-20 (-28)
× 5-10 (-13) μm, subclavate to broadly clavate
or subcylindrical, seldom lobed, hyaline,
inamyloid, thin- to thick-walled; apical setulae
4-8 (-12) × 1-2.5 μm, cylindrical to conical,
subacute to obtuse, hyaline to light yellow or
sometimes pale yellowish brown, inamyloid,
thick-walled. Pleurocystidia absent. Pileipellis
weakly mottled, composed of a hymeniform
layer of Siccus-type broom cells and pileosetae:
1) Siccus-type cells with main body 10-24 × 613 μm, subclavate to clavate or irregular in
outline, often lobed, hyaline to brownish
yellow, inamyloid, thick-walled; apical setulae
5-16 (-26) × 1-2 μm, cylindrical to conical or
irregular, subacute to obtuse, light yellow to
yellowish brown, inamyloid, thick-walled; 2)
pileosetae 26-60 × 4-7 μm, filiform to
lanceolate or irregular in outline, sometimes
branched with apical setulae up to 100 µm long,
base often swollen up to 10 µm diam,
inamyloid, thick-walled; numerous cells
transitional between typical broom cells and
setae, with 4-6 simple or forked setulae up to
40 µm long. Pileus trama subparallel to
interwoven; hyphae 2-6 μm diam, cylindrical,
hyaline, inamyloid, thin-walled, non-gelatinous.
Lamellar trama regular; hyphae 3-7 μm diam,
cylindrical, hyaline, inamyloid to weakly
dextrinoid, thin-walled, non-gelatinous. Stipe
tissue monomitic; cortical hyphae 4-6 μm diam,
parallel, cylindrical, smooth, dark brown,
inamyloid, thick-walled (0.5-1 μm); medullary
hyphae 2-5 μm diam, parallel, cylindrical,
smooth, hyaline, inamyloid, thin- to thickwalled. Caulocystidia absent. Clamp connections present.
Habit, habitat and known distribution:
gregarious on leaves of undetermined
dicotyledonous plants. Indonesia (Java),
Malaysia (Johore, Kedah, Kuala Lumpur),
Thailand.
Material examined: Malaysia, Johore, Endau
Rompin National Park, trail to Upeh Geling and trail to
Dato’s Ghani, 12 Jul. 2005, Yee-Shin Tan, TYS 485
(KLU-M #9, SFSU); same location, 12 Jul. 2005, YeeShin Tan, TYS 487 (KLU-M #10, SFSU); Johore, Endau
31
Fig. 8. Marasmius berambutanus (TYS 485 = KLU-M#9). a. Basidiomes. b. Basidiospores. c. Basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e =10 µm.
Rompin National Park, Kuala Jasin, 13 Jul. 2005, YeeShin Tan, TYS 493 (KLU-M #12, SFSU); same location,
13 Jul. 2005, Yee-Shin Tan, TYS 496 (KLU-M #13,
SFSU); same location; 13 Jul. 2005, Yee-Shin Tan, TYS
499 (KLU-M#133, SFSU); same location, Peta village,
14 Jul. 2004, Yee-Shin Tan, TYS 503 (KLU-M#45,
SFSU); Kedah, Langkawi Island, Matchinchang Forest
Reserve, 1 Sept. 2004, Yee-Shin Tan, TYS 337 (KLU-M
#11, SFSU); Kuala Lumpur, MNS heritage trail, 6 May
2004, Yee-Shin Tan, KUM 60168 (KLU-M #14, SFSU);
Selangor, Hulu Langat, Sungai Chongkak Forest Reserve,
7 Jan. 2005, Yee-Shin Tan, TYS 398 (KLU-M#44,
SFSU).
Notes: Marasmius berambutanus was
described from material collected from a single
site on Mt. Halimun, Java (Desjardin et al.,
32
2000). The tiny, collariate basidiomes with
pilosetae and Siccus-type pileipellis broom
cells with elongated setulae are diagnostic for
the species. This is the first report of the
species from Malaysia. The Malaysian
specimens differ subtly from the Java
specimens in forming pilei coloured grayish
orange when young and often become darker in
age (i.e., more deeply pigmented), and they
have up to 12 lamellae per basidiome (versus
7-10 in Java material).
A tetrapolar (bifactorial) mating system
was reported for M. berambutanus by Tan et al.
(2007) based on Malaysian material.
Fungal Diversity
7. Marasmius brevicollus Corner, Beih.
Nova Hedwigia 111: 37. 1996. (Fig. 9 & 10)
Type: Singapore, Corner s.n., 2 Apr.
1941 (E #205870!).
Pileus 2-7 mm diam, obtusely conical
when young, becoming broadly and truncately
conical with an umbilicate disc containing a
small fuscous papilla, sulcate to umbilicus;
surface dry, dull, minutely velutinous; dark
brown (7F8) to reddish brown (8F7-8), paler
surrounding the papilla. Lamellae adnate to a
collarium, distant (9-11), yellowish white with
dark brown marginate edges. Stipe 15-23 × 0.20.5 mm, central, cylindrical, terete, dry, dull,
glabrous, insititious, yellowish white at apex,
golden yellow (5B7-8) to reddish brown at the
base, arising directly from the substrate or
rarely from brownish orange rhizomorphs.
Basidiospores (16-) 17-20 (-21) × 3-4 (4.5) μm [x = 18.8 ± 1.4 × 3.7 ± 0.4 μm, Q = 46.6, Qm = 5.1 ± 0.7, n = 30 spores], elongatelacrimoid to fusoid or clavate, smooth, hyaline,
inamyloid. Basidia 20-30 × 7-10 μm, clavate,
4-spored. Basidioles 21-25 × 6-9 μm, fusoid to
clavate. Cheilocystidia numerous, composed of
Siccus-type broom cells; main body 13-20 (-26)
× 6-10 (-17) μm, cylindrical to clavate or
broadly clavate, hyaline, inamyloid, thin- to
thick-walled; apical setulae 2-5 × 0.5-1 μm,
crowded, cylindrical to conical, subacute to
obtuse, pale yellowish brown, inamyloid, thinto thick-walled. Pleurocystidia absent.
Pileipellis mottled, composed of a hymeniform
layer Siccus-type broom cells; main body 1220 × 7-16 μm, cylindrical to clavate, broadly
clavate or more commonly irregular in outline,
often lobed, hyaline to light yellow, weakly
dextrinoid, thin- to thick-walled; apical setulae
2-5 × 0.5-1 μm, cylindrical to conical, pale
brown to tawny, weakly dextrinoid, thickwalled. Pileus trama subparallel; hyphae 3-6
μm diam, cylindrical, hyaline, weakly dextrinoid, thin-walled, non-gelatinous. Lamellar
trama regular; hyphae 5-9 μm diam, cylindrical,
hyaline, weakly dextrinoid, thin-walled, nongelatinous. Stipe tissue monomitic; cortical
hyphae 5-7 μm diam, parallel, cylindrical,
smooth, yellow (apex) to brownish yellow
(base), dextrinoid, thick-walled (-2 μm);
medullary hyphae 3.5-6 μm diam, parallel,
cylindrical, smooth, hyaline, weakly dextrinoid,
thin-walled. Caulocystidia absent. Clamp
connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves. Malaysia (Johore), Singapore, Thailand.
Material examined: Malaysia, Johore, Endau
Rompin National Park, NERC, Peta Village, on the way
to jetty, 14 Jul. 2005, Yee-Shin Tan, TYS 517 (KLU-M
#15, SFSU). Singapore, 2 Apr. 1941, Corner s.n. (E
#205870).
Notes: Diagnostic features of this species
include: a small sulcate-umbilicate pileus
coloured dark brown to reddish brown or
fuscous bay with a paler zone in the umbilicus
surrounding a small fuscous papilla; collariate
lamellae with dark brown edges; a reddish
brown stipe; long and narrow, clavate
basidiospores in the range 17-20 × 3-4 μm
(mean 18.8 × 3.7 μm); and Siccus-type
cheilocystidia and pileipellis broom cells.
Marasmius brevicollus differs from M.
purpureobrunneolus Henn. [= M. acierufus
Corner (E!)] primarily in forming longer
basidiospores and a pileus without purple tones
(Desjardin et al., 2000).
A tetrapolar (bifactorial) mating system
was reported for M. brevicollus by Tan et al.
(2007) based on Malaysian material.
Type study of Marasmius brevicollus:
The holotype specimen (Corner s.n., 2
Apr. 1941, E #205870) consists of approx. 12
basidiomes, whole or fragmented, in fair
condition, attached to dicot leaves. As dried:
Pileus 1.5-3 mm diam, convex, shallowly
umbilicate with a small dark papilla, sulcate,
glabrous, deep reddish brown. Lamellae
narrowly adnate to a narrow collarium,
subdistant to distant, broad, cream with cream
or slightly darker edges. Stipe 15-30 × 0.1 mm,
central, filiform, wiry, insititious, glabrous,
brown; with numerous hair-like, wiry, dark
brown rhizomorphs; basidiomes not arising
from rhizomorphs.
Basidiospores 17-19.2 × 4.0-4.8 µm,
clavate to subfusoid, sometimes slightly curved,
smooth, hyaline, inamyloid, thin-walled.
Hymenial cells reviving poorly: no basidia,
basidioles
or
pleurocystidia
observed.
Cheilocystidia abundant, edge sterile, of
Siccus-type broom cells; main body 9.5-16 × 58 µm, clavate to irregular in outline, hyaline,
thin-walled or apically thick-walled; apical
33
Fig. 9. Marasmius brevicollus (TYS 517 = KLU–M#15). a. Basidiomes. b. Basidiospores. c. Basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
Fig. 10. Marasmius brevicollus (Holotype: E#205870). a. Basidiospores. b. Cheilocystidia. c. Pileipellis. Bar = 10 µm.
setulae 1.5-5 (-6.5) × 0.5-1.2 µm, cylindrical to
conical, pale tawny, weakly dextrinoid, thin- to
thick-walled. Pileipellis weakly mottled, a
34
hymeniform layer of Siccus-type broom cells;
main body 7-18 × 6-10 µm, cylindrical to
clavate or irregular in outline, often lobed,
Fungal Diversity
basally thin-walled to thick-walled, apically
thick-walled, hyaline to pale yellowish brown;
some cells thicker walled and with fewer more
knob-like setulae; apical setulae 1.2-5 × 0.5-2
µm, irregularly cylindrical to knob-like, obtuse,
yellowish brown in KOH, weakly dextrinoid,
thick-walled. Stipe cortical hyphae cylindrical,
parallel, smooth, brown, strongly dextrinoid,
thick-walled; medullary hyphae dextrinoid.
Caulocystidia absent. Clamp connections
present.
8. Marasmius purpureobrunneolus Henn.,
Monsunia 1: 151. 1900
(Fig. 11 & 12, Plate 1D)
= Marasmius acierufus Corner, Beih. Nova
Hedwigia 111: 25. 1996 (Type: E #205854!)
Type: Java, Bogor Botanical Garden, 12
Feb. 1898, E. Nyman. Not extant.
Neotype: Java, Bogor Botanical Garden,
11 Mar. 1977, E. Horak 77-179 (SFSU!).
Pileus 1.5-6 mm diam, convex with
distinctly depressed to umbilicate disc, papilla
absent; surface dry, dull, pruinose; margin
sulcate to plicate; dark brown (8F8), reddish
brown (9E7-8) to violet brown (10F8) overall.
Lamellae adnate to a collarium, subdistant (1014) with no lamellulae, yellowish white with
reddish brown to dark brown edges. Stipe 1238 × 0.5 mm, central, terete, equal, wiry, tough,
solid, dry, dull, glabrous, insititious, pale
yellow (3A3) at apex, gradually grading to
orange to brownish orange (6B5-6) at the base.
Basidiospores 12-17 (-19) × 3.5-5 μm [x
= 14.7 ± 2.3 × 4.1 ± 0.7 μm, Q = 3-4.3, Qm =
3.6 ± 0.4, n = 34 spores], elongate-ellipsoid to
fusoid, smooth, hyaline, inamyloid, thin-walled.
Basidia 20 × 5 μm, 4-spored. Basidioles 25 × 6
μm, fusoid. Lamellar edge sterile, with a broad
band of cheilocystidia; cheilocystidia of
Siccus-type broom cells; main body 10-23 (-25)
× 7-12 (-15) μm, cylindrical to clavate or
vesiculose, rarely lobed, hyaline to yellowish
brown, inamyloid to weakly dextrinoid, thinwalled; apical setulae (2-) 3-5 × 0.5-1 μm,
crowded, bluntly cylindrical to broadly conical,
sometimes forked, light yellow, inamyloid to
dextrinoid, thin- to thick-walled. Pleurocystidia
absent. Pileipellis mottled, composed of a
hymeniform layer of Siccus-type broom cells;
main body (10-) 12-20 (-24) × 7-12 (-14) μm,
clavate to broadly clavate or irregular in outline,
rarely lobed, hyaline, inamyloid to weakly
dextrinoid, thick-walled; apical setulae (2-) 3-4
× 1 μm, broadly acute, yellow to yellowish
brown, inamyloid, thick-walled. Pileus trama
interwoven; hyphae 3.5-5 μm diam, cylindrical,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Lamellar trama regular; hyphae 3-5 μm diam,
cylindrical, hyaline, dextrinoid, thin-walled,
non-gelatinous. Stipe tissue monomitic; cortical
hyphae 3-5 μm diam, parallel, cylindrical,
smooth, yellow, dextrinoid, thick-walled, nongelatinous; medullary hyphae 5-7 μm, parallel,
cylindrical, smooth, hyaline, inamyloid, thinwalled, non-gelatinous. Caulocystidia absent.
Clamp connections present.
Habit, habitat and known distribution:
solitary and gregarious on undetermined
dicotyledonous leaves and bamboo leaves.
Indonesia (Java), Malaysia (Negeri Sembilan),
Singapore.
Material examined: Malaysia, Johore, EndauRompin National Park, Dato’s Ghani Trail, 12 Jul. 2005,
Yee-Shin Tan, TYS 484 (KLU-M#113); Negeri
Sembilan, Ulu Bendul Forest Reserve, 23 Oct. 2004,
Yee-Shin Tan, TYS 374 (KLU-M#112, SFSU). Type of
M. acierufus: Singapore, Botanic Garden, 19 March
1943, Corner s.n. (E #205854).
Notes: Marasmius purpureobrunneolus is
characterized by a small, convex, dark brown
to reddish brown or violet brown pileus that
lacks a papilla, subdistant (10-14), collariate
lamellae with marginated edges, a long, wiry
glabrous stipe, relatively short basidiospores
(12-17 × 3.5-5 μm with mean 14.7 × 4.1 μm),
pigmented broom cells and growth on
dicotyledonous leaves. Marasmius acierufus,
described recently by Corner (1996) from
material collected in Singapore, is accepted
here as a colour form of M. purpureobrunneolus. The protologue of the former reports
the pileus as purple rufous to purple madder
with the margin becoming cinnamon rufous in
age, whereas in M. purpureobrunneolus the
pileus is more dark purplish brown overall.
Type study of Marasmius purpureobrunneolus:
For a study of the neotype refer to
Desjardin et al. (2000).
35
Fig. 11. Marasmius purpureobrunneolus (TYS 374 = KLU-M#112). a. Basidiomes. b. Basidiospores. c. Basidia and
basidiole. d. Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
Type study of Marasmius acierufus:
The holotype specimen (Singapore,
Botanic Garden, Corner s.n., 19 March 1943, E
#205854) consists of approx. 16 basidiomes in
fair condition, most attached to dicot leaves,
rarely to twigs. As dried: Pileus 3-5 mm diam,
convex, umbilicate, with or without a small
papilla, sulcate-plicate, glabrous, dark reddish
brown. Lamellae adnate to a collarium, distant
with no lamellulae, broad, cream, edges
concolorous. Stipe 15-22 × 0.2 mm, central,
terete, filiform, insititious, reddish brown to
dark brown; some leaves with wiry, hair-like,
glabrous rhizomorphs.
36
Basidiospores 14-17.5 × 3.5-4.5 µm,
subfusoid to elongate-clavate, sometimes
curved, smooth, hyaline, inamyloid, thinwalled. Basidia 4-spored, clavate. Basidioles
clavate to subfusoid. Cheilocystidia abundant,
edge sterile, of Siccus-type broom cells; main
body 11-16 × 6-10 µm, clavate to pyriform,
sometimes lobed, hyaline, thin-walled; apical
setulae 1.5-3 (-4) × 0.8-1.8 µm, broadly conical,
obtuse to subacute, thin-walled to firm-walled
(< 0.5 µm), hyaline to pale yellow in KOH.
Pleurocystidia absent. Pileipellis not mottled to
weakly mottled, a hymeniform layer of Siccustype broom cells; main body 8-16 × 4.8-12 µm,
irregularly clavate to pyriform, sometimes
Fungal Diversity
Fig. 12. Marasmius acierufus (Holotype: E#205854). a. Basidiospores. b. Cheilocystidia. c. Pileipellis. Bar = 10 µm.
lobed, thin-walled below, apically thick-walled,
hyaline to pale tawny; apical setulae 1.5-3 (-4)
× 0.8-2 µm (rather short and broad), broadly
conical, sometimes forked, obtuse to subacute,
pale tawny, inamyloid, thick-walled. Pileus
trama and lamellar trama regular; hyphae
inamyloid. Stipe cortical hyphae 3-5 µm diam,
parallel, cylindrical, brown, strongly dextrinoid,
thick-walled; medullary hyphae similar but
hyaline, dextrinoid, thin-walled. Caulocystidia
absent. Clamp connections present
9. Marasmius guyanensis Mont., Ann. Sci.
Nat. Bot. sér. 4, 1: 114. 1854.
(Fig. 13, Plate 1E)
Type:
French
Guyana,
Leprieur.
Authentic specimen, Herb. Berk. (K).
Pileus 1-5 mm diam, convex to
campanulate, umbilicate, with or without a
papilla; margin sulcate to plicate; surface dry,
dull, glabrous to minutely pruinose; colour
ranging from orange (5A6) to yellowish orange
(4A6-7) or pale yellow (3A3) overall with the
umbilicus black or brownish orange. Lamellae
adnate to a collarium, distant to subdistant (812) with no lamellulae, not intervenose, broad,
yellowish white. Stipe 12-23 × 0.1-0.2 mm,
central, terete, wiry, dry, dull, glabrous,
insititious, dark brown to black overall; black
rhizomorphs present.
Basidiospores 9-16 × (2.2-) 3-5 μm [xmr =
10.1-14.2 × 3.3-4.7 μm, xmm = 12.5 ± 1.5 × 3.9
± 0.4 μm, Q = 2.5-5.7, Qmr = 2.4-4.2, Qmm =
3.1 ± 0.8, n = 24 spores per 9 specimens]
ellipsoid, smooth, hyaline, inamyloid. Basidia
not observed. Basidioles 22-24 × 5-6.4 μm,
clavate to fusoid. Pleurocystidia absent.
Cheilocystidia numerous, composed of Siccustype broom cells; main body 6-13 (-16) × 3.8-8
μm, clavate to subclavate, cylindrical or
irregular in outline, rarely lobed, hyaline,
inamyloid, thick-walled; apical setulae 1.6-3.2
× 0.6-1.3 μm, crowded, cylindrical to conical,
obtuse, light yellow, inamyloid to dextrinoid,
thick-walled. Pileipellis weakly mottled, a
hymeniform layer of Siccus-type broom cells;
main body 6.4-14.4 × 4.8-9.6 μm, cylindrical
to clavate, broadly clavate or irregular in
outline, hyaline, inamyloid, thick-walled;
apical setulae 1-3.2 × 0.5-1 μm, conical, obtuse
to subacute, rarely forked, hyaline to light
yellow, inamyloid, thin- to thick-walled. Pileus
trama interwoven; hyphae 2.6-3.2 μm diam,
light yellow, inamyloid, thin-walled, nongelatinous. Lamellar trama regular; hyphae
37
Fig. 13. Marasmius guyanensis (TYS 329 = KLU-M#135). a. Basidiomes. b. Basidiospores. c. Basidia and basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
3.2-4.2 μm diam, cylindrical, hyaline,
inamyloid, thin-walled, non-gelatinous. Stipe
tissue monomitic; cortical hyphae 2-3 μm diam,
parallel, cylindrical, smooth, brownish yellow
to brown, inamyloid to weakly dextrinoid,
thick-walled, non-gelatinous; medullary hyphae 3-5 μm diam, parallel, cylindrical, smooth,
hyaline, inamyloid to weakly dextrinoid, thinwalled, gelatinous. Caulocystidia absent.
Clamp connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves and twigs. Africa (Benin, Cameroon,
DR Congo, Nigeria, Sierra Leone), Indonesia
(Java), Malaysia (Selangor), South America
(Brazil, Bolivia, French Guyana, Trinidad,
Venezuela), Thailand.
Material examined: Malaysia, Johore, EndauRompin National Park, Trail to Upeh Guling, 12 Jul.
2005, Yee-Shin Tan, TYS 488 (KLU-M#158); same
location, way to Dato’s Ghani Trail and Kuala Jasin, 13
Jul. 2005, Yee-Shin Tan, TYS 494 (KLU-M#159); same
location, way to NERC, Peta Village and Temiang Trail,
13 Jul. 2005, Yee-Shin Tan, TYS 520 (KLU-M#160);
Kedah, Langkawi Island, Matchinchang Forest Reserve,
30 Aug. 2004, Yee-Shin Tan, TYS 329 (KLU-M#135,
SFSU); same location, 30 Aug. 2004, Yee-Shin Tan,
38
TYS 332 (KLU-M#139); same location, 1 Sept. 2004,
Yee-Shin Tan, TYS 342 (KLU-M#140); Mount Raya, N
06° 25’ 52.8, E 99° 49’ 26.5, 652 m elev., 6 Apr. 2004,
Yee-Shin Tan, KUM 60017 (KLU-M#143); Lubuk
Semilang Forest Reserve, N 06° 21’ 55.2, E 99° 47’ 26.5,
59 m elev., 8 Apr. 2004, Yee-Shin Tan, KUM 60144
(KLU-M#141); same location, 8 Apr. 2004, Yee-Shin
Tan, KUM 60146 (KLU-M#142); same location, 8 Apr.
2004, Yee-Shin Tan, KUM 60147 (KLU-M#144); Kuala
Lumpur, MNS heritage trail, 6 May 2004, Yee-Shin Tan,
KUM 60167 (KLU-M#150); Selangor, Ulu Gombak,
University of Malaya’s Field Study Center, 28 Jul. 2003,
Yee-Shin Tan, TYS 028 (KLU-M#137, SFSU); same
location, 28 Jul. 2003, Yee-Shin Tan, TYS 030 (KLUM#151, SFSU); same location, 28 Jul. 2003, Yee-Shin
Tan, TYS 031 (KLU-M#152); same location, 6 Sept.
2003, Yee-Shin Tan, TYS 080 (KLU-M#153, SFSU) ;
same location, 8 Nov. 2003, Yee-Shin Tan, TYS 215
(KLU-M#154, SFSU); same location, 16 Oct. 2004,
Yee-Shin Tan, TYS 349 (KLU-M#155); same location,
16 Oct. 2004, Yee-Shin Tan, TYS 354 (KLU-M#156);
same location, 16 Oct. 2004, Yee-Shin Tan, TYS 355
(KLU-M#157); Negeri Sembilan, Ulu Bendul Forest
Reserve, 23 Oct. 2004, Yee-Shin Tan, TYS 373 (KLUM#162, SFSU); same location, 23 Oct. 2004, Yee-Shin
Tan, TYS 372 (KLU-M#148); same location, 23 Oct.
2004, Yee-Shin Tan, TYS 375 (KLU-M#149); Selangor,
Hulu Langat Sungai Chongkak Forest Reserve, N
03°12.705’, E101° 50.472’, 188 m elev., 7 Jan. 2005,
Yee-Shin Tan, TYS 395 (KLU-M#136, SFSU); same
Fungal Diversity
location, 7 Jan. 2005, Yee-Shin Tan, TYS 401 (KLUM#145); same location, 7 Jan. 2005, Yee-Shin Tan, TYS
404 (KLU-M#146); Selangor, Selayang, Kanching
Forest Reserve, N 03° 17.958;, E 101° 37.151’, 110 m
elev., 8 Jan. 2005, Amy Honan, TYS 407 (KLU-M#161);
same location, 9 Jan. 2005, Yee-Shin Tan, TYS 433
(KLU-M#147).
Notes: Marasmius guyanensis is a
widespread, apparently pantropical species
distinguished by a small, orange, yellowish
orange or pale yellow plicate pileus with or
without a central black papilla, collariate
lamellae, and an insititious stipe that does not
arise directly from rhizomorphs. The
Malaysian material differs subtly from that
reported in Indonesia is having up to 12
lamellae per basidiome (versus 7-10 lamellae
in Java material; Desjardin et al., 2000).
A tetrapolar (bifactorial) mating system
was reported for M. guyanensis by Wannathes
et al. (2007) based on Thai material.
10. Marasmius crinis-equi F. Muell. ex
Kalchbr., Grevillea 8: 153. 1880.
(Fig. 14)
= Marasmius equicrinis F. Muell. ex Berk., J.
Linn. Soc., Bot. 18: 383. 1881.
= Androsaceus crinis-equi (F. Muell. ex Kalchbr.)
Overeem, Hoofd van Het Mus. Econ. Bot. Buitenzorg 1:
69. 1927.
= Marasmius graminum Berk. & Broome var.
equicrinis (F. Muell. ex Berk.) Dennis, Trans. Brit.
Mycol. Soc. 34: 416. 1951.
= Marasmius repens Henn. Engl. Bot. Jahrb. 23:
548. 1897
= Marasmius ramentaceus (Pat.) Sacc. &
Traverso, Syllog. Fung. 20: 21. 1911.
= Androsaceus ramentaceus Pat., Ann. Jard. Bot.
Buitenzorg 1: 107. 1897.
Type: Australia, North Queensland,
Rockingham Bay, F. von Mueller (K).
Pileus 1-4 mm diam, convex to broadly
convex, umbilicate, without a papilla; margin
sulcate; surface dry, dull, minutely pruinose;
brownish orange (6C8) to grayish orange (5B3).
Lamellae adnate to a collarium, distant (6-8)
with no lamellulae, yellowish white, nonmarginate. Stipe 4-5 × 0.5 mm, cylindrical,
central, terete, wiry, dry, dull, glabrous,
insititious, yellowish white at the apex, black at
the base, arising directly from wiry, black
rhizomorphs.
Basidiospores (7-) 9-13.5 × 3-4 μm [xmr =
8.9-13 × 3.4-4.5 μm, xmm = 10.7 ± 1.3 × 3.9 ±
0.4 μm, Q = 1.8-3.3, Qmr = 2.4-3.2, Qmm = 2.8
± 0.3, n = 20 spores per 13 specimens],
ellipsoid, hyaline, smooth, inamyloid, thinwalled. Basidia 28-30 × 5-6 μm, fusoid, 4spored. Basidioles 16-25 × 6-7 μm, clavate to
fusoid. Cheilocystidia numerous, composed of
Siccus-type broom cells; main body (5-) 9-16 (28) × 5-10 μm, clavate to broadly clavate or
cylindrical, rarely lobed, hyaline, inamyloid to
weakly dextrinoid, thick-walled; apical setulae
(2-) 3-6 × 0.5-1 μm, crowded, rarely forked,
cylindrical to conical, obtuse, light yellow,
inamyloid to weakly dextrinoid, thin- to thickwalled. Pleurocystidia absent. Pileipellis
mottled, a hymeniform layer of Siccus-type
broom cells; main body 9-20 × 5-14 μm,
clavate to subclavate or irregular in outline,
rarely lobed, hyaline to brownish yellow,
inamyloid, thick-walled; apical setulae 2-6 ×
0.5-1.5 μm, cylindrical to conical, subacute,
rarely forked, light yellow to yellow or brown,
inamyloid, thin- to thick-walled. Pileus trama
interwoven; hyphae 3-5μm diam, hyaline,
inamyloid, thin-walled, non-gelatinous. Lamellar trama regular; hyphae 5-8 μm diam,
cylindrical, hyaline to light yellow, weakly
dextrinoid to dextrinoid, thin-walled, nongelatinous. Stipe tissue monomitic; cortical
hyphae 3-5 μm diam, parallel, cylindrical,
smooth, yellowish brown at apex, brown at
base, weakly dextrinoid, thick-walled (1 μm);
medullary hyphae 3-4 μm diam, parallel,
cylindrical, smooth, hyaline, inamyloid, thinwalled, non-gelatinous. Caulocystidia absent.
Clamp connections present.
Habit, habitat and known distribution:
solitary on rhizomorph or sometimes on leaves
or twigs. Putatively pantropical; Africa, South
America, Indonesia (Java), Malaysia (Kedah,
Negeri Sembilan, Pahang, Selangor), Sri Lanka,
Thailand.
Material examined: Malaysia, Kedah, Langkawi
Island, Matchinchang Forest Reserve, 1 Sept. 2004, YeeShin Tan, TYS 338 (KLU-M#61, SFSU); same location,
1 Sept. 2004, Yee-Shin Tan, TYS 339 (KLU-M#62);
same location, 1 Sept. 2004, Yee-Shin Tan, TYS 341
(KLU-M#63, SFSU); Negeri Sembilan, Ulu Bendul
Forest Reserve, 23 Oct. 2004, Yee-Shin Tan, TYS 367
(KLU-M#55, SFSU); same location, 23 Oct. 2004, YeeShin Tan, TYS 365 (KLU-M#56); same location, 23 Oct.
2004, Yee-Shin Tan, TYS 366 (KLU-M#57, SFSU);
same location, 23 Oct. 2004, Yee-Shin Tan, TYS 381a
(KLU-M#58); same location, 23 Oct. 2004, Yee-Shin
Tan, TYS 382 (KLU-M#59); same location, 23 Oct.
2004, Yee-Shin Tan, TYS 383 (KLU-M#60); Pahang,
Fraser’s Hill, Kindersley’s Trail, 16 Jan. 2004, Yee-Shin
39
Fig. 14. Marasmius crinis-equi (TYS 466 = KLU-M#48). a. Basidiomes. b. Basidiospores. c. Basidia and basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
Tan, TYS 251 (KLU-M#131, SFSU); Selangor, Hulu
Langat, Sungai Chongkak Forest Reserve, N 03° 12.705’,
E101° 50.472’, 188 m elev., 16 May 2005, Yee-Shin Tan,
TYS 466 (KLU-M#48, SFSU); same location, 7 Jan.
2005, Yee-Shin Tan, TYS 397 (KLU-M#49); same
location, 6 Mar. 2005, Yee-Shin Tan, TYS 443 (KLUM#50, SFSU); same location, 6 Mar. 2005; Yee-Shin
Tan, TYS 444 (KLU-M#51); same location,16 May
2005, Yee-Shin Tan, TYS 459 (KLU=M#52); Selangor,
Ulu Gombak, University of Malaya’s Field Study Center,
6 Sept. 2003, Yee-Shin Tan, TYS 081 (KLU-M#53);
same location, 16 Oct. 2004, Yee-Shin Tan, TYS 361
(KLU-M#54); Selangor, Rawang, Kanching Forest
Reserve, N 03°17.958’, E 101° 37.151’, 110 m elev., 8
Jan. 2005, Yee-Shin Tan, TYS 412 (KLU-M#127,
SFSU); same location, 8 Jan. 2005, Yee-Shin Tan, TYS
422 (KLU-M#64); same location, 9 Mar. 2005, Yee-Shin
Tan, TYS 447 (KLU-M#65, SFSU).
40
Notes: Marasmius crinis-equi is the most
common member of the subsect. with brownish
orange to deep orange, sulcate pilei and
basidiomes that arise directly from copious
rhizomorphs. The Malaysian populations differ
slightly from the Indonesian and Sri Lankan
material in typically forming evenly pigmented
pilei that lack a papilla and dark central spot.
In all other features they are indistinguishable.
11. Marasmius kuthubutheenii Y.S. Tan,
Desjardin, Vikineswary & Noorlidah sp. nov.
(Fig. 15, Plate 1F)
MycoBank: MB 512635
who
Etymology: Named in honor of Dr. Kuthubutheen
contributed to mycology including applied
Fungal Diversity
mycology research in Malaysia.
Pileus 1-3 mm latus, convexus vel late convexus,
papillatus ad centrum, striatus, glaber, brunneoaurantiacus. Lamellae adnato-collariatae, distantes (6-9),
albidae vel pallide stramineae, subbrunneo-marginatae.
Stipes 2-4 × 0.2 mm, cylindricus, glabrus, insititius,
brunneus, rhizomorpha glaber sed haud exoriens.
Basidiosporae 7.5-9 × 3-4 μm, ellipsoideae, leves,
hyalinae, inamyloideae. Basidia 20-25 × 6 μm, clavatae,
4-spora. Cheilocystidia typi Sicci, abundantia, 8-15 × 614 μm, subcylindracea vel clavata, hyalina; setulae ad
apicem 3-5 × 0.5-1 μm, cylindricae vel conicae,
subacutae vel obtusae, luteae, inamyloideae, crassetunicatae. Pleurocystidia nulla. Pileipellis hymeniformis,
typi Sicci, cellulae 10-16 × 7-12 μm, clavatae,
subclavatae vel irregularia, hyalinae, inamyloideae,
crasse-tunicatae; setulae ad apicem 4-8 × 0.5-1 μm,
cylindricae vel conicae, subacutae vel obtusae, luteobrunneae, inamyloideae, crasse-tunicatae. Caulocysitidia
nulla. Fibulae presentes. Gregarius ad folia putrida
bambusarum. Holotypus: Malaysia, Negeri Sembilan,
Ulu Bendul Forest Reserve, 23 Oct. 2004, Yee-Shin Tan,
TYS 364 (Holotypus: KLU-M#76).
Pileus 1-3 mm diam, convex to broadly
convex, disc depressed with an acute papilla;
margin striate; surface dry, dull, glabrous to
minutely pruinose; brownish orange (6C4-5) to
light brown (7D5) overall with a brown to
black papilla, drying tan to pale brown.
Lamellae adnate to a collarium, distant (6-9),
yellowish white (2A2) with light brown edges.
Stipe 2-4 × 0.2 mm, central, cylindrical,
filiform, dry, dull, glabrous, insititious,
yellowish white at apex, brown at base;
associated with brown, wiry, glabrous
rhizomorphs but not arising from them.
Basidiospores 7.5-9 (-10) × 3-4 (-4.5) μm
[xmr = 7.9-8.7 × 3.7-3.9 μm, xmm = 8.3 ± 0.6 ×
3.8 ± 0.1 μm, Q = 1.8-3, Qmr = 1.6-3, Qmm =
2.2 ± 0.3, n = 26 spores per 2 specimens],
ellipsoid, smooth, hyaline, inamyloid. Basidia
20-25 × 6 μm, clavate, 4-spored (few observed).
Basidioles 19-25 × 5-7 μm, fusoid to clavate.
Cheilocystidia numerous, composed of Siccustype broom cells; main body 8-15 (-17) × (5-)
6-14 (-17) μm, clavate to broadly clavate or
subcylindrical, rarely lobed, hyaline, inamyloid,
thick-walled; apical setulae (2-) 3-5 × 0.5-1 μm,
forked, cylindrical to conical, subacute to
obtuse, seldom forked, yellow, inamyloid, thinto thick-walled. Pleurocystidia absent.
Pileipellis mottled, composed of a hymeniform
layer of Siccus-type broom cells; main body (7-)
10-16 × (5-) 7-12 μm, clavate to subclavate or
sometimes irregular in outline, rarely lobed,
hyaline, inamyloid, thick-walled; apical setulae
(3-) 4-8 × 0.5-1 μm, cylindrical to conical,
subacute to obtuse, seldom forked, brownish
yellow to dull yellow, inamyloid, thick-walled.
Pileus trama interwoven; hyphae 3-4 μm diam,
cylindrical, hyaline, inamyloid, thin-walled,
non-gelatinous. Lamellar trama regular;
hyphae 5-7 μm diam, cylindrical, hyaline,
inamyloid to weakly dextrinoid, thin-walled,
non-gelatinous. Stipe tissue monomitic; cortical
hyphae 2-5 μm diam, parallel, cylindrical,
smooth, yellow at apex, dull yellow at base,
dextrinoid to strongly dextrinoid, thick-walled;
medullary hyphae 3-4 μm diam, parallel,
cylindrical, smooth, hyaline, inamyloid, thickwalled. Caulocystidia absent. Clamp connections present.
Habit, habitat and known distribution:
gregarious on bamboo leaves. Malaysia
(Negeri Sembilan, Selangor).
Material examined: Malaysia, Negeri Sembilan,
Ulu Bendul Forest Reserve, 23 Oct. 2004, Yee-Shin Tan,
TYS 364 (Holotype: KLU-M#76; Isotype: SFSU);
Selangor, Ulu Gombak, University of Malaya’s Field
Study Centre, 16 Oct. 2004, Yee-Shin Tan, TYS 348
(KLU-M#77, SFSU).
Notes: Marasmius kuthubutheenii is
characterized by the following features: a
convex-depressed pileus 1-3 mm diam
coloured light brown to brownish orange (dries
tan to pale brown) with a black papilla; 6-9
collariate lamellae with light brown edges; a
short wiry stipe (2-4 × 0.1 mm) that is
associated with wiry, glabrous rhizomorphs but
does not arise directly from them;
basidiospores 7.5-10 × 3-4.5 µm (xmm = 8.3 ×
3.8 µm); Siccus-type cheilocystidia with
narrowly conical, acute setulae 3-5 × 0.5-1 µm;
Siccus-type pileipellis broom cells with
narrowly conical, acute setulae 4-8 × 0.5-1 µm;
and growth on bamboo leaves. The new species
is phenetically similar to M. subruforotula
Singer from Africa, M. aripoensis (Dennis)
Singer from Trinidad, and M. foliicola Singer
from Argentina. Marasmius subruforotula
differs in forming larger pilei (3-10 mm diam),
more lamellae (up to 12), a longer stipe (7-25
mm), and grows on dicotyledonous leaves.
Marasmius aripoensis differs in forming a
brightly pigmented pileus (deep orangish
ferruginous to mahogany red), has more
41
Fig. 15. Marasmius kuthubutheenii (Holotype: TYS 364 = KLU-M#76). a. Basidiomes. b. Basidiospores. c. Basidia and
basidioles. d. Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
lamellae (7-13), a longer stipe (up to 30 mm),
and grows on dicotyledonous leaves.
Marasmius foliicola differs in forming a larger
pileus (4-7 mm diam) coloured more intensely
orange, has more lamellae (10-13), and a much
longer stipe (15-75 mm); it does, however,
grow on bamboo leaves.
12. Marasmius gracilichorda Corner, Beih.
Nova Hedwigia 111: 57. 1996.
(Fig. 16 & 17, Plate 1G)
Type: Singapore, Gardens Jungle, Corner
s.n., 15 Sept. 1940 (E #206712!).
Pileus 2-6.5 mm diam, convex,
umbilicate, sulcate; surface dry, dull, minutely
velutinous; brownish orange (6C7-8, 7C8) to
light brown (5D6-7,7D8), reddish brown (8D8,
8E7) or dark brown (6E8, 8F7-8, 9F8) overall
with a tiny black dot in the umbilicus.
Lamellae adnate to a collarium, distant (6-12),
yellowish white (2A2) with brown marginate
edges. Stipe 8-23 × 0.2 mm, central, cylindrical,
filiform, dry, dull, glabrous, insititious,
yellowish white at apex, dark brown to black at
42
base, often associated with fine, dark
rhizomorphs.
Basidiospores 8-11 × (3-) 3.5-5 μm [xmr =
9.2-10 × 4-4.6 μm, xmm = 9.6 ± 0.3 × 4.3 ± 0.3
μm, Q = 1.8-3.3, Qmr = 2.1-2.4, Qmm = 2.3 ±
0.2, n = 30 spores per 5 specimens], ellipsoid,
smooth, hyaline, inamyloid. Basidia 26-32 × 68 μm, clavate, 4-spored. Basidioles 23-28 × 4-7
μm, fusoid to clavate. Cheilocystidia numerous,
composed of Siccus-type broom cells; main
body 10-25 (-30) × 4-12 (-15) μm,
subcylindrical to clavate or broadly clavate,
rarely lobed, light yellow to light brown,
inamyloid, thin-walled; apical setulae 5-8 (-10)
× 0.5-2 (-2.5) μm, crowded, rarely forked,
cylindrical to conical, subacute to obtuse,
brownish yellow to light brown, inamyloid,
thick-walled. Pleurocystidia absent. Pileipellis
strongly mottled, composed of a hymeniform
layer of Siccus-type broom cells; main body
10-23 (-30) × 6-12 (-15) μm, turbinate to
clavate, broadly clavate or irregular in outline,
often lobed, light brown to reddish brown,
Fungal Diversity
Fig. 16. Marasmius gracilichorda (TYS 442 = KLU-M#16). a. Basidiomes (x2). b. Basidiospores. c. Basidia and
basidioles. d. Cheilocystidia. e.Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
inamyloid, some thin-walled and with shorter
setulae and others thick-walled with longer and
more coarsely warted setulae; apical setulae 512 (-13) × 0.5-1.5 (-3) μm, cylindrical to
conical, many warted or lumpy, often forked,
brownish yellow to brown, reddish brown or
dark brown, inamyloid, thick-walled. Pileus
trama interwoven; hyphae 3-5 μm diam,
cylindrical, hyaline to light yellow, inamyloid,
thin-walled, non-gelatinous. Lamellar trama
regular; hyphae 3-5 μm diam, cylindrical,
hyaline, inamyloid to weakly dextrinoid, thinwalled, non-gelatinous. Stipe tissue monomitic;
cortical hyphae 4-6 μm diam, parallel,
cylindrical, smooth, brown, inamyloid, thickwalled (-0.5 μm); medullary hyphae 2-5 μm
diam, parallel, cylindrical, smooth, hyaline,
inamyloid, thin-walled. Caulocystidia absent.
Clamp connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves.
Malaysia
(Pahang,
Selangor),
Singapore.
Material examined: Malaysia, Pahang, Fraser's
Hill, Kindersley's Trail, 23 September 2003, Yee-Shin
Tan, TYS 190 (KLU-M#132, SFSU); Selangor, Hulu
Langat, Sungai Chongkak Forest Reserve, N 03°12.705’,
E 101°50.472’, elev. 122 m, 7 Jan 2005, Yee-Shin Tan,
TYS 396 (KLU-M#70, SFSU); same location, 6 Mar.
2005, Yee-Shin Tan, TYS 442 (KLU-M #16, SFSU);
same location, 16 May 2005, Yee-Shin Tan, TYS 472
(KLU-M#71, SFSU); Selangor, Selayang, Kanching
Forest Reserve, N 03°17.958’, E 101°37.151’, elev. 110
m, 8 Jan 2005, Yee-Shin Tan, TYS 411 (KLU-M#69,
SFSU); Singapore, Gardens Jungle, 15 Sept. 1940,
Corner s.n. (Holotype E #206712).
43
Fig. 17. Marasmius gracilichorda (Holotype: E#206712). a. Basidiospores. b. Cheilocystidia. c. Pileipellis. Bar = 10
µm.
Notes: The most distinctive feature of M.
gracilichorda is the pileipellis anatomy. In this
species, the pileipellis is mottled, composed of
a combination of relatively thin-walled Siccustype broom cells with conical setulae 5-7 μm
long, and many thicker-walled and more
deeply pigmented broom cells with setulae 512 μm long that are forked or lobed and often
lumpy or warted. Corner (1996) described the
latter as “antler-like lobulate”. Macroscopically,
M. gracilichorda looks like M. ruforotula
Singer, but the characteristic pileipellis of the
former clearly distinguishes it.
A tetrapolar (bifactorial) mating system
was reported for M. gracilichorda by Tan et al.
(2007) based on Malaysian material.
Type study of Marasmius gracilichorda:
The holotype specimen (Singapore,
Gardens Jungle, Corner s.n., 15 Sept. 1940, E
#206712) consists of approx. 20 basidiomes in
good condition, pressed flat and attached to
several dicot leaves. As dried: Pileus 1.5-3 mm
diam, when young with a tall, obtusely conical
papilla surrounded by a short skirt, in age the
"skirt" expands to become the pileus that is
broadly convex, striate to sulcate, with a
depression surrounding the central papilla, dull,
minutely granulose; papilla brown to dark
44
brown, elsewhere brownish orange. Lamellae
adnate to a collarium, subdistant (10-13), no
lamellulae, broad, apparently non-marginate.
Stipe 10-18 × 0.1 mm, filiform, wiry, insititious,
glabrous, dark brown to black; with or without
fine, wiry, dark brown rhizomorphs; basidiomes not arising from rhizomorphs.
Basidiospores 8.6-11.2 × 4-5 µm,
ellipsoid to subfusoid or subclavate, smooth,
hyaline, inamyloid, thin-walled. Basidia 4spored. Basidioles subfusoid. Cheilocystidia
abundant, of Siccus-type broom cells; main
body 10-16 × 4.5-7 (-9.5) µm, clavate, seldom
lobed, hyaline, thin-walled or apically thickwalled; apical setulae 2.2-6 × 0.6-1.5 µm,
irregularly conical, smooth or with scattered
warts, thin- to thick-walled, hyaline to pale
yellow. Pleurocystidia absent. Pileipellis not
mottled to mottled, a hymeniform layer of
Siccus-type broom cells; main body 8-18 x 512 µm, irregularly cylindrical to broadly
clavate or irregular in outline, often lobed,
hyaline at base, yellowish brown at apex,
apically thick-walled or entirely thick-walled (1.5 µm), with scattered very thick-walled cells
interspersed; apical setulae 2-10 × 1-1.5(-2.5)
µm, irregularly conical, smooth to wavy in
outline and often with warts or small branches,
yellowish brown, inamyloid, thick-walled.
Tramal hyphae inamyloid. Stipe cortical
Fungal Diversity
hyphae cylindrical, smooth, brown, strongly
dextrinoid, thick-walled; medullary hyphae
dextrinoid. Caulocystidia absent. Clamp
connections present.
13. Marasmius ruforotula Singer, Sydowia 2:
34. 1948.
(Fig. 18, Plate 1H)
Type: United States, Florida, Dade Co.,
Matheson Hammock, 3 Nov. 1942, Singer F
1456 (FH!).
Pileus 2-8 mm diam, convex to broadly
convex or sometimes plano-convex, disc
depressed with an acute papilla; margin striate
to sulcate; surface dry, dull, glabrous to
minutely velutinous; brownish orange (6C4-5)
to reddish brown (7C8) or dark brown (7F8)
when young, becoming grayish orange (6B5)
to brownish orange (6C6-7) to light brown
(7D5) overall with a black papilla at maturity.
Lamellae adnate to a collarium, distant (8-13),
yellowish white (2A2), non-marginate. Stipe 515 × 0.2 mm, central, cylindrical, terete, equal,
dry, dull, glabrous, insititious, yellowish white
at apex, brown on the base, associated with
black, wiry, tough, black rhizomorphs and
rarely arising from them.
Basidiospores 9-11 (-12) × 4-5 (-6) μm
[xmr = 8.9-10.5 × 4.2-5 μm, xmm = 9.6 ± 0.6 ×
4.6 ± 0.3 μm, Q = 1.8-2.7, Qmr = 1.5-3.3, Qmm
= 2.1 ± 0.1, n = 29 spores per 5 specimens],
ellipsoid, smooth, hyaline, inamyloid. Basidia
20-28 × 6-10 μm, clavate, 4-spored. Basidioles
28-30 × 5-8 μm, fusoid to clavate. Cheilocystidia numerous, composed of Siccus-type
broom cells; main body 10-18 (-20) × 7-13 (-15)
μm, subcylindrical, clavate to broadly clavate
or irregular in outline, rarely lobed, hyaline,
inamyloid, thick-walled; apical setulae 2-4 (-5)
× 1-2 μm, cylindrical to conical, rarely forked
or warted, light yellow, inamyloid, thin-walled.
Pleurocystidia absent. Pileipellis weakly
mottled, composed of a hymeniform layer of
Siccus-type broom cells; main body (10-) 1118 (-27) × 9-15 (-16) μm, turbinate to clavate,
broadly clavate, subglobose or subcylindrical,
seldom lobed, hyaline, inamyloid, thin- to
thick-walled; apical setulae 2-3 (-5) × 0.5-1 μm,
cylindrical to conical, subobtuse, seldom
forked, light yellow to light brown, inamyloid
to weakly dextrinoid, thin-walled. Pileus trama
interwoven; hyphae 2.5-5 μm diam, cylindrical,
hyaline, inamyloid to weakly dextrinoid, thinwalled, non-gelatinous. Lamellar trama regular;
hyphae 3-6 μm diam, cylindrical, hyaline,
inamyloid, thin-walled, non-gelatinous. Stipe
tissue monomitic; cortical hyphae 3.5-5 μm
diam, parallel, cylindrical, smooth, brown,
weakly dextrinoid, thick-walled; medullary
hyphae 3-6 μm diam, parallel, cylindrical,
smooth, light yellow, inamyloid, thick-walled
(1-2 μm). Caulocystidia absent. Clamp
connections present.
Habit, habitat and known distribution:
gregarious on bamboo and other monocotyledonous leaves. Ecuador, Indonesia (Bali,
Java), Malaysia (Selangor), Mexico, Papua
New Guinea, Thailand, United States (Florida).
Material examined: Malaysia, Johore, EndauRompin National Park, 14 Jul. 2005, Yee-Shin Tan,
TYS 509 (KLU-M#95, SFSU). United States, Florida,
Dade Co., Matheson Hammock, 3 Nov. 1942, Singer
F 1456 (Holotype: FH!); Negeri Sembilan, Ulu
Bendul Forest Reserve, 23 Oct. 2004, Yee-Shin Tan,
TYS 369 (KLU-M#93, SFSU); same location, 23 Oct.
2004, Yee-Shin Tan, TYS 370 (KLU-M#96, SFSU);
Selangor, Kanching Forest Reserve, N 03°17. 958’, E
101°37.151’, elev. 110m, 9 Jan. 2005, Yee-Shin Tan,
TYS 438 (KLU-M#94, SFSU); Selangor, Ulu
Gombak, University of Malaya’s Field Study Centre,
16 Oct. 2004, Yee-Shin Tan, TYS 360 (KLU-M#97,
SFSU); same location, 28 Nov. 2004, Yee-Shin Tan,
TYS 385 (KLU-M#98, SFSU).
Notes:
Marasmius
ruforotula
is
characterized by the following features: a
convex-depressed pileus coloured grayish
orange to brownish orange or light brown with
a dark papilla; 8-13 collariate lamellae with
non-marginate edges; and numerous black
rhizomorphs. The Malaysian material differs
from Indonesian specimens in having slightly
larger basidiospores mean (9.6 × 4.6 µm versus
8.9 × 4.3 µm in Java material) and shorter stipe
(3-15 × 0.2 mm versus 7-25 × 0.1-0.3 mm in
Java material; Desjardin et al,. 2000).
Basidiospores size in the holotype specimen is
nearly identical to that of the Indonesian
populations.
A tetrapolar (bifactorial) mating system
was reported for M. ruforotula by Wannathes
et al. (2007) based on Thai material.
45
Fig. 18. Marasmius ruforotula (TYS 438 = KLU-M#94). a. Basidiomes. b. Basidiospores. c. Basidia and basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
Type study of Marasmius ruforotula:
The holotype specimen (United States,
Florida, Dade Co., Matheson Hammock, 3 Nov.
1942, Singer F 1456, FH) consists of several
basidiomes in good condition. As dried: Pileus
convex-umbilicate with a small centralpapilla,
striate or sulcate to the central depression,
glabrous, deep ferruginous overall (lacking a
pale zone surrounding a dark central spot).
Lamellae adnate to a well-developed collarium,
subdistant, broad, pallid, non-marginate. Stipe
filiform, glabrous, shiny, upper half
stramineous, base ochraceous or brown,
insititious, lignicolous.
Basidiospores 7.5-9.5 × 4-5 µm [x = 8.6
± 0.5 × 4.3 ± 0.3 µm, Q = 1.8-2.2, Qm = 2 ± 0.1,
n = 20], ellipsoid or sublacrymoid, hyaline,
inamyloid, smooth. Basidia 20-26 × 5.5-7.5 µm,
clavate, 4-spored. Basidioles cylindrical to
46
fusoid. Cheilocystidia numerous, of Siccustype broom cells; main body 12-16 × 6.5-10
µm, clavate, hyaline, thin-walled; apical setulae
2-5 × 1-2 µm, cylindrical to subconical, obtuse,
thick-walled, hyaline or pale yellow.
Pleurocystidia absent. Pileipellis weakly
mottled, a hymeniform layer of Siccus-type
broom cells; main body 12-20 × 7-12 µm,
cylindrical to clavate or turbinate, seldom
lobed, many pale yellow or pale orange and
thin-walled, many orange or orange-tawny and
thick-walled; apical setulae 2-5 × 1-2 µm, rodlike, conical or irregular in outline, obtuse,
thick-walled and yellowish orange, or solid and
orange-tawny to ferruginous. Pileus trama
interwoven; lamellar trama regular; hyphae 310 µm diam, cylindrical, smooth, nongelatinous, hyaline, inamyloid or weakly
dextrinoid, thin-walled. Stipe tissue monomitic;
cortical hyphae 3-5.5 µm diam, parallel,
Fungal Diversity
cylindrical, smooth, stramineous or tawny,
dextrinoid, thick-walled (0.5-1.2 µm); medullary hyphae similar but hyaline, inamyloid,
thinner-walled. Caulocystidia absent. Clamp
connections present.
Marasmius sect. Globulares Kühner,
Botaniste 25: 100. 1944 (ut Globularineae).
Type species: Marasmius globularis Fr. (= M.
wynneae Berk. & Broome).
14. Marasmius aff. camerunensis Antonín &
Mossebo, Mycotaxon 85: 113. 2003.
(Fig. 19, Plate 2A)
Type: Cameroon, Littoral Province, near
the village of Poola’a, ca. 5 km from
Nkongsamba, 20 Aug. 1998. leg. D.C.
Mossebo M196(1) (BRNM 670732).
Pileus 16-58 mm diam, convex to planoconvex; disc flat or very slightly broadly
depressed in age; margin smooth to striate or
radially wrinkled; surface dry, dull, hygrophanous, glabrous; disc and radial wrinkles
grayish brown (8D3) to brownish orange (7C3)
or grayish orange (6B3), margin pinkish brown.
Context thin (< 1 mm), tough, light brown.
Lamellae broadly adnate, distant to subdistant
(10-16) with 1-2 series of lamellulae, very
broad, pale brownish gray, non-marginate.
Stipe 45-150 × 2-4 mm, central, cylindrical,
terete, equal, tough, pliant to cartilaginous,
hollow, dull, dry, glabrous, non-insititious,
grayish brown to light brown overall, base
covered with yellowish white mycelium. Odor
and taste not distinctive.
Basidiospores (17-) 25-35 (-40) × 5-7 μm
[x = 28.4 ± 6.2 × 5.8 ± 0.7 μm, Q =3.3-6.4, Qm
= 5.0 ± 1.0, n = 29 spores], elongate-lacrymoid
to clavate, smooth, hyaline, inamyloid, thinwalled. Basidia not observed. Basidioles 35-52
× 8-10 μm, fusoid to clavate. Cheilocystidia
common, (17-) 18-26 (-28) × (8-) 10-16 μm,
subglobose to broadly clavate or subcylindrical,
hyaline, inamyloid, thin-walled. Pleurocystidia
absent. Pileipellis hymeniform, not mottled,
composed of clavate to broadly clavate,
pyriform or subglobose cells (11-) 12-20 (-25)
× (6-) 9-13 μm, hyaline, inamyloid, thin-walled.
Pileus trama interwoven; hyphae 4.5-8 μm
diam, cylindrical, light yellow, dextrinoid, thinwalled (skeletals absent), non-gelatinous.
Lamellar trama regular; hyphae 3-5 μm diam,
cylindrical, hyaline, dextrinoid to strongly
dextrinoid, thin-walled, non-gelatinous. Stipe
tissue monomitic; cortical hyphae 4-10 μm
diam, parallel, cylindrical, smooth, yellow,
dextrinoid, thick-walled, non-gelatinous; medullary hyphae 4-7 μm, parallel, cylindrical,
smooth, light yellow, dextrinoid, thick-walled,
non-gelatinous. Caulocystidia absent. Clamp
connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves in lowland forest. Cameroon, Malaysia
(Kedah).
Material examined. Malaysia, Kedah, Langkawi
Island, Lubuk Semilang Forest Reserve, N 06° 21.58.4’,
E 99° 47.29.6’, elev. 59 m, 8 Apr. 2004, Yee-Shin Tan,
KUM 60134 (KLU-M#46, SFSU); same location, 8 Apr.
2004, Yee-Shin Tan, KUM 60135 (KLU-M#47, SFSU).
Notes: Two specimens of this taxon are
known from Malaysia. It is most closely allied
with M. camerunensis from Africa. Distinctive
features of the Malaysian material include: a
convex-depressed, striate pileus 16-58 mm
diam coloured grayish brown to brownish
orange with a pinkish brown margin; subdistant
(10-16), pale brownish grey lamellae; a long
(45-150 mm), glabrous, grayish brown stipe;
variable basidiospores in the range 17-40 × 5-7
µm with mean 28.4 × 5.8 µm; no
pleurocystidia; broadly clavate cheilocystidia
and pileipellis cells in the range 11-30 × 6-16
µm lacking setulae; no caulocystidia; and
growth on dicotyledonous leaves. The
Malaysian material differs from the description
of Antonín (2007) in forming more deeply
pigmented lamellae (not white), and in growing
on leaf debris (not lignicolous); other features
are indistinguishable. Until more material is
collected in Malaysia and compared with
authentic African specimens, the specimens
reported here are tentatively identified as M.
camerunensis.
15. Marasmius purpureostriatus Hongo, J. Jap.
Bot. 33: 344. 1958.
(Fig. 20, Plate 2B)
Type: Japan, Otsu City, 7 May 1957,
Hongo #1609, Herbarium Hongo (Isotype ZT!).
Pileus 5-22 mm diam, convex to planoconvex, disc depressed or subumbilicate,
sometimes with an umbo; margin sulcate to
plicate; surface dry, dull, hygrophanous,
47
Plate 2. Marasmius s.s. in section Globulares, Neosessiles, Leveilleani and Sicci. A: Marasmius aff. camerunensis (x
0.5); B: Marasmius purpureostriatus (x 2); C: Marasmius aff. leoninus (x 1.5); D: Marasmius tenuissimus (x 0.5); E:
Marasmius araucariae var. siccipes (x 0.5); F: Marasmius nummularioides (x1); G: Marasmius leveilleanus (x 0.5); H :
Marasmius iras (x 1); I: Marasmius luteomarginatus (x 3).
glabrous; disc and radial furrows purplish gray
(14E3) to dull violet (17D4) or grayish violet
(18E5), dull yellow (3B3) to yellowish white
(3A2) elsewhere. Context thin (< 1 mm),
purplish white. Lamellae adnate, distant (10-13)
with 1 series of lamellulae, very broad,
yellowish white (3A2), non-marginate. Stipe
17-50 × 1 mm, central, cylindrical, terete, equal,
tough, hollow, dull, dry, non-insititious,
yellowish gray (2B2) at apex, grayish yellow
(4B4-5) at base, base covered with yellowish
white mycelium. Odor and taste not distinctive.
48
Basidiospores (20-) 22-26 (-30) × 4-6 μm
[x = 25 ± 1.9 × 5 ± 0.6 μm, Q =4.2-6.4, Qm =
5.1 ± 0.5, n = 23 spores], elongated-lacrymoid
to narrowly clavate, smooth, hyaline, inamyloid,
thin-walled. Basidia 28-35 × 6 μm, clavate, 4spored. Basidioles 25-30 × 6-7 μm, fusoid to
clavate. Cheilocystidia common, 16-25 (-27) ×
8-12 (-15) μm, subglobose to broadly clavate,
hyaline, inamyloid, thin-walled. Pleurocystidia
absent. Pileipellis hymeniform, not mottled,
composed of clavate to broadly clavate,
Fungal Diversity
pyriform or subglobose cells 13-24 (-35) × 8-16
(-24) μm, hyaline, inamyloid, thin-walled.
Pileus trama interwoven; hyphae 4-10 μm
diam, cylindrical, hyaline, dextrinoid, thinwalled, non-gelatinous. Lamellar trama regular;
hyphae 4-7 μm diam, cylindrical, hyaline,
dextrinoid, thin-walled, non-gelatinous. Stipe
tissue monomitic; cortical hyphae 6-8 μm diam,
parallel, cylindrical, smooth, yellow at apex,
pale brown at base, weakly dextrinoid, thickwalled, non-gelatinous; medullary hyphae 4-9
μm diam, parallel, cylindrical, smooth, hyaline,
inamyloid, thin-walled, non-gelatinous. Caulocystidia absent. Clamp connections present.
Fig. 19. Marasmius aff. camerunensis (KUM 60134 = KLU–M#46). a. Basidiomes. b. Basidiospores. c. Basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
49
Fig. 20. Marasmius purpureostriatus (TYS 328 = KLU-M#92). a. Basidiomes. b. Basidiospores. c. Basidia and
basidioles. d. Cheilocystidia. e. Pileipellis. Bars: a = 20 mm, b-e = 10 µm.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves. Japan, Malaysia (Kuala Lumpur),
Papua New Guinea, Thailand.
Material examined. Malaysia, Kuala Lumpur,
University of Malaya, 21 Jul. 2004, S. Vikineswary,
TYS 328 (KLU-M#92, SFSU).
Notes: Marasmius purpureostriatus is
characterized by forming a striped, plicate
pileus coloured grayish violet to dull violet on
the disc and radial furrows and pale yellowish
white elsewhere, distant (10-13), very broad
lamellae, a glabrous, pallid stipe, long, clavate
basidiospores (22-26 × 4-6 µm), an absence of
pleurocystidia
and
caulocystidia,
and
association with dicotyledonous debris. This
species have been reported previously from
Papau New Guinea (Desjardin & Horak, 1997).
Marasmius
sect.
Neosessiles
Mycologia 50: 104. 1958.
50
Singer,
Type species: Marasmius neosessilis
Singer.
16. Marasmius tenuissimus (Jungh.) Singer,
Fl. Neotrop. Monogr. 17: 258. 1976.
(Fig. 21, Plate 2D)
= Agaricus tenuissimus Jungh., Verh. Batav.
Genootsch. 17: 84. 1838.
= Marasmius rufescens Berk. & Broome, J. Linn.
Soc. Bot. 14: 41. 1873.
= Xerotus tener Berk. & Broome, J. Linn. Soc.
Bot. 14: 45. 1873.
= Marasmius campanella Holterm., Mykol.
Unters. Tropen: 105. 1898.
= Marasmius campanella var. rufescens (Berk. &
Broome) Petch, in Petch & Bisby, Peradeniya Manual. 6:
59. 1950.
Type: Indonesia, Java, near Kebokoening, May
(L).
Pileus (2.5-) 15-40 (-45) mm diam,
irregularly convex in face view, to campanulate
in side view, deeply plicate-folded to rugulosereticulate, pustulate, dull, dry, glabrous overall
Fungal Diversity
or minutely pruinose near attachment point;
disc brownish orange (6-7C7-8), grading to
pale orange (6B4-5) to orangish white (5A2-3)
at the margin, nearly cream-buff on the margin
of large basidiomes. Context thin, white.
Lamellae adnexed, remote, interwoven and
anastomosing to subreticulate, with many
transverse lamellulae of shorter height, broad
(1-4 mm), cream (4A3) to pale orangish white
(5A2-3), non-marginate. Stipe usually absent,
rarely lateral and rudimentary, up to 2 × 1 mm,
cylindrical, pruinose, white with a brown base,
non-insititious. Odor none.
Basidiospores (6-) 7-10 × 3-6 μm [xmr =
7.1-9.4 × 3.3-5.2 μm, xmm = 8.5 ± 1.2 × 4.6 ±
0.8 μm, Q = 1.3-2.4, Qmr = 1.7-2.1, Qmm = 1.9
± 0.2, n = 20 spores per 5 specimens], ellipsoid
to fusoid, smooth, hyaline, inamyloid, thinwalled. Basidia 30 × 4 μm, 4-spored.
Basidioles 29 × 6 μm, fusoid. Cheilocystidia
common, polymorphic, composed of Siccustype broom cells; main body (10-) 11-16 (-17)
× (5-) 6-9 μm, cylindrical to clavate, fusoid or
irregular in outline, rarely lobed, hyaline,
inamyloid, thin- to thick-walled; apical setulae
1-3 × 1 μm, broadly conical to irregularly
cylindrical or knobby, obtuse to subacute,
rarely forked, hyaline, inamyloid, thin-walled.
Pleurocystidia absent. Pileipellis mottled,
composed of a hymeniform layer of two types
of cells: a) Siccus-type broom cells with main
body (6-) 10-20 (-25) × (5-) 7-11 μm, clavate
to broadly clavate, cylindrical or irregular in
outline, rarely lobed, hyaline to yellowish
brown, inamyloid, thick-walled; apical setulae
(1-) 3-5 (-7) × 1-2 μm, broadly conical, obtuse
to subacute, yellow to yellowish brown,
inamyloid to dextrinoid, thin- to thick-walled
(up to 2 µm); b) Globulares-type cells, rare, 913 × 10-18 μm, clavate to subglobose or
irregular in outline, rarely lobed, non-setulose,
hyaline, inamyloid to dextrinoid, thick-walled
(0.5-1.5 µm). Pileus trama interwoven; hyphae
5-7 μm diam, cylindrical, hyaline, dextrinoid,
thin-walled, slightly gelatinous. Lamellar
trama regular; hyphae 2-3 μm diam, cylindrical,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Stipe tissue not observed. Caulocystidia absent.
Clamp connections present.
Habit, habitat and known distribution:
scattered on undetermined liana stem in
primary montane cloud forest, and gregarious
on bamboo leaves and on undetermined
dicotyledonous leaves and twigs. Indonesia
(Java), Malaysia (Kedah, Pahang, Selangor),
Thailand.
Material examined: Malaysia, Kedah, Langkawi
Island, Matchinchang, 1 Sept. 2004, Yee-Shin Tan, TYS
345 (KLU-M#100, SFSU); Pahang Prov., Fraser’s Hill,
Hemmant Trail, N03˚42.925’, E101˚44.306’, elev. 1200
m., 15 Jan. 2004, E. Horak, DED 7659 (KLU-M#102,
SFSU); Selangor, Hulu Langat, Sungai Chongkak Forest
Reserve, 16 May 2005, Yee-Shin Tan, TYS 468 (KLUM#101, SFSU); Selangor, Ulu Gombak, University of
Malaysia’s Field Study Centre, 28 Nov. 2004, Yee-Shin
Tan, TYS 398 (KLU-M#103, SFSU); same location, 5
Jan. 2005, Yee-Shin Tan, TYS 392 (KLU-M#104).
Notes: Marasmius tenuissimus is
characterized by moderately large, deeply
plicate-folded to rugulose-reticulate, pale
orange pileus, adnexed, broad, intervenose
lamellae, sessile basidiomes or with a
rudimentary lateral to eccentric stipe, relatively
short and broad basidiospores, and growth on
liana stems and dicotyledonous leaves. It is the
only commonly encountered Marasmius in
Southeast Asia that lacks a stipe and has
Siccus-type broom cells.
Molecular data
suggest that it is more closely related to species
currently placed in sect. Marasmius subsect.
Sicciformes and sect. Leveilleani, than it is to
members of sect. Sicci.
Marasmius sect. Hygrometrici Kühner,
Botaniste 25:95. 1933.
Type species : Marasmius hygrometricus
(V. Brig.) Sacc. [= Marasmius corbariensis
(Roum.) Sacc. & Traverso].
17. Marasmius micraster Petch, Trans. Brit.
Mycol. Soc. 31: 42. 1947.
(Fig. 22)
Type: Sri Lanka, Kandy District,
Peradeniya, 21 Oct. 1914, Petch 4195 (K).
Pileus 1-2 mm diam, convex to obtusely
conical, smooth when young, becoming striate
to sulcate in age, dull, dry, minutely pruinose,
pale grayish brown overall, in age and when
expanded becoming paler and with a nearly
white margin. Lamellae shallowly adnexed, not
collariate, distant with 0-1 series of lamellulae,
moderately broad (up to 0.5 mm), white. Stipe
10-18 × 0.1 mm, central, wiry, glabrous, shiny,
tough, insititious, dark brown to black;
associated with short, black rhizomorphs.
51
Fig. 21. Marasmius tenuissimus (DED 7659 = KLU-M#102). a. Basidiomes. b. Basidiospores. c. Basidia and basidiole.
d. Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
Basidiospores 9-12 (-13) × 4.5-6.5 (-7)
µm [xmr = 9.9-11.3 × 4.7-6 µm, xmm 10.5 ± 0.7
× 5.2 ± 0.6 µm, Q = 1.6-2.8, Qmr = 1.8-2.2,
Qmm = 2 ± 0.2, n= 17 spores per 5 specimens],
ellipsoid, smooth, hyaline, inamyloid. Basidia
28-35 × 7-10 µm, clavate, 4-spored. Basidioles
32-40 × 8-10 µm, fusoid to clavate.
Pleurocystidia absent. Cheilocystidia composed of two types of cells: a) Rotalis-type
broom cells with main body 33-42 (-60) × 7-12
(-16) µm, subclavate to clavate or subglobose,
hyaline, inamyloid, thin-walled or apically
thick-walled; divergent setulae 1-2 × 0.5-1 µm,
cylindrical, densely setulose over the upper
half of cell, hyaline, inamyloid, thin-walled; b)
non-setulose cells, 30-36 × 7-9 (-12) µm,
clavate to cylindrical or more commonly fusoid,
hyaline, inamyloid, thin-walled. Pileipellis
52
mottled, composed of a hymeniform layer of
Rotalis-type broom cells; main body 17-22 (-30)
× 8-20 (-22) µm, broadly clavate to subglobose,
hyaline to pale orangish brown, inamyloid,
thin-walled; divergent setulae 1-1.5 × 0.5-1 µm,
cylindrical, obtuse, light yellow, inamyloid,
thin-walled. Pileus trama interwoven; hyphae
3-5 µm diam, cylindrical, hyaline, inamyloid,
thin-walled. Lamellar trama regular; hyphae 38 µm diam, cylindrical, hyaline, inamyloid,
thin-walled. Stipe tissue monomitic; cortical
hyphae 5-8 µm diam, parallel, cylindrical,
densely setulose overall, thin- to thick-walled,
brown, inamyloid; setulae thin-walled, hyaline,
inamyloid; medullary hyphae 4-6 µm diam,
parallel, smooth, thin-walled, pale yellow,
inamyloid. Caulocystidia absent. Clamp
connections present.
Fungal Diversity
Fig. 22. Marasmius micraster (TYS 229 = KLU-M#86). a. Basidiomes. b. Basidiospores. c. Basidia and basidiole. d.
Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
Habit, habitat and known distribution:
gregarious on petioles and main veins of
undetermined dicot trees; in Malaysia on a
single species of tree with very large leaves, ca
30-40 cm long × 15-20 cm wide in lowland
forests. Indonesia (Java), Malaysia (Selangor,
Pahang), New Zealand, Singapore, Sri Lanka.
Material examined: Malaysia, Pahang, Cameron
Highlands, Brinchang mountain, jungle walk no.1, N
04˚31.087’, E 101˚22.971’, elev. ca 2210 m, 10 Jan.
2004, Yee-Shin Tan, TYS 229 (KLU-M#86, SFSU);
Selangor, Ulu Gombak, University of Malaya’s Field
Study Centre, N 03˚19.500’, E 101˚45.167’, elev. ca 240
m, 13 Jan. 2004, D.E. Desjardin, DED 7647= TYS 237
(KLU-M#82, SFSU); same location, 27 Aug. 2003, YeeShin Tan, TYS 068 (KLU-M#83, SFSU); same location,
5 Jan. 2005, Yee-Shin Tan, TYS 391 (KLU-M#84,
SFSU); same location, 5 Jan. 2005, Yee-Shin Tan, TYS
393 (KLU-M#85, SFSU).
Notes: Marasmius micraster was
described originally from Sri Lanka and has
been reported recently from New Zealand
(Desjardin & Horak, 1997) and Indonesia
(Desjardin et al., 2000). Distinctive features of
this tiny taxon include a small, pale grayish
brown, sulcate pileus, distant and noncollariate lamellae, a wiry, glabrous stipe
associated with black rhizomorphs, basidiospores in the range of 9-12 × 4.5-6.5 µm,
Rotalis-type cheilocystidia and pileipellis
broom cells, and growth on dicotyledonous
leaves. The Malaysian material differs from
Indonesian specimens in having slightly larger
basidispores with mean 10.5 × 5.2 µm versus 9
× 4.5 µm in Indonesia (Desjardin et al., 2000).
Marasmius sect. Leveilleani Singer, Bull. Jard.
Bot. Bruxelles. 34: 362: 1964.
Type species: Marasmius leveilleanus
(Berk.) Pat.
18. Marasmius leveilleanus (Berk.) Pat., Bull.
Soc. Mycol. Fr. 33: 55. 1917.
(Fig. 23, Plate 2G)
= Heliomyces leveilleanus Berk., Lond. J. Bot. 6:
490. 1847.
53
= Marasmius umbraculum Berk. & Broome, J.
Linn. Soc. Bot. 14: 36. 1873.
Type: Sri Lanka, Hautane Range, July
1844, Gardner 72 (K #99705).
Pileus 10-36 mm diam, convex to
broadly convex, distinctly depressed or with an
umbo; margin striate to sulcate; surface dry,
dull, subvelutinous; reddish brown (8D7) to
brown (7D7), light brown (7D6) or brownish
orange (7C8). Context thin, yellowish white.
Lamellae free, no collarium, subdistant to close
(20-24), pale yellow (3A3), non-marginated.
Stipe 16-80 × 1 mm, central, cylindrical, terete,
dry, dull, glabrous, shiny, insititious, light
brown at apex, dark brown at base; associated
with coarse dark brown rhizomorphs.
Basidiospores (8-) 9-12 (-12.5) × 4-5.5
μm [xmr = 9.6-11.6 × 4.9-5.1 μm, xmm = 10.6 ±
1 × 5 ± 0.1 μm, Q = 1.7-2.9, Qmr = 1.8-2.4,
Qmm = 2.2 ± 0.2, n = 21 spores per 3
specimens], ellipsoid, smooth, hyaline,
inamyloid, thin-walled. Basidia not observed.
Basidioles 18-23 × 5-6 μm, fusoid to clavate.
Cheilocystidia numerous, composed of Siccustype broom cells; main body (10-) 16-30 (-38)
× (5-) 6-10 μm, cylindrical to clavate, rarely
lobed, light yellow, inamyloid, thin- to thickwalled; apical setulae 3-6 (-8) × 1.5-2 (-3) μm,
bluntly or broadly conical to knobby, rarely
forked, hyaline, inamyloid, thin- to thickwalled. Pleurocystidia absent. Pileipellis
mottled, composed of a hymeniform layer of
two types of cells: a) Siccus-type broom cells
with main body (9-) 17-30 (-40) × (6-) 8-10 (11) μm, cylindrical to clavate, broadly clavate
or irregular in outline, rarely lobed, hyaline to
light yellowish brown, inamyloid, thick-walled;
apical setulae 5-8 × 1-3 (-4) μm, cylindrical to
broadly conical or knobby, obtuse, yellowish
brown to tawny, inamyloid, thin- to thickwalled; b) rare, scattered non-setulose cells,
clavate, tawny, apically thick-walled. Pileus
trama interwoven; hyphae 3.5-6 μm diam,
cylindrical, hyaline, inamyloid, thin-walled,
non-gelatinous. Lamellar trama regular;
hyphae 3-7 μm diam, cylindrical, hyaline to
light yellow, inamyloid, thin-walled, nongelatinous. Stipe tissue monomitic; cortical
hyphae 5-7 μm diam, parallel, cylindrical,
smooth, light yellow at stipe apex, pale brown
at base, inamyloid or weakly dextrinoid, thick54
walled; medullary hyphae 5-10 μm diam,
parallel,
cylindrical,
smooth,
hyaline,
inamyloid or weakly dextrinoid, thin-walled.
Caulocystidia absent. Clamp connections
present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves and dead wood. Africa (DR Congo,
Ghana, Ivory Coast, Nigeria, Uganda),
Malaysia (Kedah, Selangor), Mexico, Sri
Lanka, Thailand.
Material examined: Malaysia, Kedah, Langkawi
Island, Lubuk Semilang Forest Reserve, N 6° 21’ 51.9, E
99° 47’ 25.8, 49 m alt., 8 Apr. 2004, Yee-Shin Tan,
KUM 60142 (SFSU); Langkawi Island, Mat ChinChang,
Datai, 7 Apr. 2004, Yee-Shin Tan, KUM 116a (SFSU);
Selangor, Ulu Gombak, University of Malaya’s Field
Study Centre, 28 Nov. 2004, Yee-Shin Tan, TYS 386
(KLU-M#80).
Notes: The monotypic section Leveilleani
was erected around M. leveilleanus and
circumscribed to contain species with the
following unique combination of features:
rather large (10-36 mm diam), deeply
pigmented pilei composed of a hymeniform
layer of a few smooth, thick-walled cells plus
numerous Siccus-type broom cells with few,
broad, knobby, obtuse setulae; free, noncollariate lamellae; a thick and wiry, insititious
stipe associated with rhizomorphs; nondextrinoid hyphae; and an absence of
pleurocystidia. Indeed, the single known
species shows features of both sect. Marasmius
(insititious stipe and rhizomorphs) and sect.
Sicci (large basidiomes with non-collariate
lamellae and Siccus-type broom cells). The
molecular data suggest that M. leveilleanus is
more closely allied with members of sect.
Marasmius than with sect. Sicci. This is the
only known member of the genus Marasmius
with truly free lamellae. In M. leveilleanus
there is a smooth ring of pileus tramal tissue
surrounding the area where the stipe attaches to
the pileus such that the lamellae stop short of
the stipe by about 1 mm. The Malaysian
specimens are lignicolous and are associated
with coarse, thick, wiry, glabrous rhizomorphs
(similar in morphology to the stipes) but the
basidiomes do not arise directly from them.
Rhizomorphs were not reported from Sri
Lankan material, but they were reported from
material collected in Africa (Antonín, 2007). It
will be interesting to see if the New World
Fungal Diversity
Fig. 23. Marasmius leveilleanus (KUM 60142 = KLU-M#79). a. Basidiomes. b. Basidiospores. c. Basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
specimen reported by Singer (1976) from
Mexico represents the same taxon as this
widespread paleotropical M. leveilleanus.
Marasmius sect. Sicci Singer, subsect. Siccini
Singer, ser. Spinulosi (Clémençon) Desjardin,
in Antonín & Noordeloos, Lib. Bot. 8: 179.
1993.
Type species: Marasmius cohaerens (Pers.: Fr.)
Cooke & Quél. [Bas. Agaricus cohaerens Pers.:
Fr.].
= ser. Actinopus Singer pro parte, Fl. Neotrop.
Monogr. 17: 236. 1976.
19. Marasmius nummularioides Desjardin &
Y.S. Tan, Fungal Diversity 25: 200. 2007.
(Fig. 24, Plate 2F)
Type: Malaysia, Johore, Endau Rompin
National Park, Peta Village, 14 July 2005, Y.S.
Tan #508 (KLU-M #7!).
Pileus 5-16 mm diam, convex, with or
without an umbo, disc rugulose, margin smooth
(non-striate); surface dry, dull, minutely
velutinous; reddish brown (8D-E6-8) to brown
(7E6-8) overall when young, margin fading to
brown (6D-E6-7) in age. Lamellae adnate,
crowded (16-18) with 3-4 series of lamellulae,
narrow, yellowish white with brown marginate
edges. Stipe 16-40 × 1 mm, central, cylindrical,
terete, equal, dry, dull, pruinose to hispidulous,
non-insititious with brownish orange basal
tomentum, yellowish orange to brownish
orange overall when young, or with a yellowish
white apex, gradually changing to brownish
orange, light brown or brown in age.
Basidiospores (15-) 16-18 (-19) × (3.5-)
4-4.5 μm [x = 16.7 ± 1.1 × 4.1 ± 0.3 μm, Q =
3.5-4.8, Qm = 4.0 ± 0.3, n = 22 spores],
elongate-lacrymoid to fusoid or clavate,
smooth, hyaline, inamyloid. Basidia not
55
observed. Basidioles 26-30 × 6-8 μm, fusoid to
clavate. Cheilocystidia numerous, composed of
Siccus-type broom cells; main body 14-20 (-28)
× 6-10 μm, cylindrical to clavate or broadly
clavate, hyaline, inamyloid, thin- to thickwalled; apical setulae 4-8 × 0.5-1 μm, crowded,
cylindrical to conical, subacute to obtuse,
rarely forked, light brownish yellow, weakly
dextrinoid, thick-walled. Pleurocystidia absent.
Pileipellis mottled, composed of a hymeniform
layer of Siccus-type broom cells; main body
10-30 × 6-10 (-13) μm, cylindrical to clavate,
broadly clavate or irregular in outline, rarely
lobed, hyaline to pale yellowish brown, weakly
dextrinoid, thick-walled; apical setulae 5-13 (17) × 1-2 μm, cylindrical to conical, rarely
forked, light brownish yellow to brown, weakly
dextrinoid,
thick-walled.
Pileus
trama
subparallel to interwoven; hyphae 4-6 μm diam,
cylindrical, hyaline, strongly dextrinoid, thinwalled, non-gelatinous. Lamellar trama regular;
hyphae 5-8 μm diam, cylindrical, hyaline to
light yellow, dextrinoid to strongly dextrinoid,
thin-walled, non-gelatinous. Stipe tissue
monomitic; cortical hyphae 3.5-5 μm diam,
parallel, cylindrical, smooth, light brown (apex)
to yellowish brown (base), weakly dextrinoid,
thick-walled (-1 μm); medullary hyphae 4-8
μm diam, parallel, cylindrical, smooth, hyaline,
dextrinoid, thin-walled. Caulocystidia numerous, composed of Siccus-type broom cells,
setae-like cells and caulosetae; main body of
broom cells and setae-like cells 14-25 × 8-15
μm, cylindrical to broadly clavate, light yellow
to light brown, inamyloid, thick-walled; apical
setulae 6-60 × 2-4 μm, conical to acicular, light
yellow, inamyloid, thick-walled. Caulosetae
scattered, 50-100 × 3-5 μm, lanceolate, light
brown, inamyloid, thick-walled. Clamp
connections present.
Habit, habitat and known distribution:
solitary and gregarious on undetermined
dicotyledonous leaves. Malaysia (Johore).
Material examined: Malaysia, Johore, Endau
Rompin National Park, on the way to NERC, Peta
Village, Temiang trail, 14 July 2005, Yee-Shin Tan,
TYS 508 (Holotype: KLU-M #7; Isotype: SFSU).
Notes: Marasmius nummularioides is
characterized by relatively small, non-striate,
rugulose pileus coloured brown to reddish
56
brown, by crowded, narrow lamellae with
brown margins, and by a pruinose to
hispidulous stipe with brownish orange basal
mycelium. In addition, the species forms
basidiospores with mean 16.7 × 4.1 μm, has
Siccus-type cheilocystidia and pileipellis
broom cells, and lacks pleurocystidia, hymenial
setae and pileosetae. The most distinctive
microscopic feature is the caulocystidia that are
a combination of Siccus-type broom cells with
very long setulae and narrow setae. The new
species is allied with M. nummularius Berk. &
Broome from Sri Lanka and Indonesia, and
with M. spiculosus Singer and M. atrorubens
(Berk.) Berk., both from South America and
the Caribbean (M. atrorubens is also known
from tropical Africa). Marasmius nummularius
differs from M. nummularioides in forming
smaller basidiospores (mean 13.6 × 4.0 μm)
and in having a stipitipellis covered with very
long, usually simple caulosetae, lacking Siccustype caulocystidia (Desjardin et al., 2000).
Marasmius spiculosus differs in forming a
distinctly sulcate pileus with numerous
pileosetae, and has refractive pleurocystidia.
Interestingly, the stipitipellis is almost identical
to that of M. nummularioides (Singer, 1976;
Pegler, 1983). Marasmius atrorubens differs in
forming a distinctly sulcate pileus with
scattered pileosetae, may or may not have
pleurocystidia, and forms a stipitipellis covered
with simple caulosetae, lacking Siccus-type
broom cells (Singer, 1976; Pegler, 1983; K!).
A bipolar (unifactorial) mating system
was reported for M. nummularioides by Tan et
al. (2007).
Marasmius sect. Sicci Singer, subsect. Siccini
Singer, ser. Atrorubentes Desjardin & Horak,
Biblio. Mycol. 168: 27. 1997.
Typespecies: Agaricus atrorubens Berk.,
London J. Bot. 1: 138. 1842 [≡ Marasmius
atrorubens (Berk.) Berk., Hooker’s J. Bot.
Kew. Gard. Misc. 8: 137. 1856].
20. Marasmius musicolor
Desjardin, sp. nov.
MycoBank: MB512625
Y.S.
Tan &
(Fig. 25)
Etymology: musi- (Latin) = banana; -color (Latin)
= coloured, referring to the banana coloured pileus.
Fungal Diversity
Fig. 24. Marasmius nummularioides (Holotype: TYS 508 = KLU-M#7). a. Basidiomes. b. Basidiospores. c. Basidioles.
d. Cheilocystidia. e. Pileipellis. f. Caulocystidia. Bars: a = 10 mm, b-f = 10 µm.
Pileus 7-21 mm latus, convexus vel planoconvexus, ad apicem rugulosus, ruguloso-striatus,
subvelutinus, pallide luteus vel musicolor. Lamellae
adnatae, densae (12-14), angustae, cremeae, haud
marginatae. Stipes 11-15 × 0.5-1 mm, cylindricus,
pruinosus, haud insititius, apicaliter luteus, basim versus
brunneo-aurantiacus, ad basim tomento pallide aurantioaldido affixus. Basidiosporae 15-18 × 3.5-4 μm, clavatae,
leves, hyalinae, inamyloideae. Basidiola 32-40 × 7 μm,
fusiodeae vel clavatae. Cheilocystidia typi Sicci, 10-22 ×
7-8 μm, cylindracea vel clavata, hyalina; setulae ad
apicem 2-6 × 0.5-1 μm, densae, cylindricae vel conicae,
subacutae vel obtusae, pallide flavae, inamyloideae,
tenui- vel crasse-tunicatae. Pleurocystidia nulla.
Pileipellis hymeniformis, typi Sicci, cellulae 14-19 × 6-
10 μm, cylindricae vel clavatae, hyalinae, inamyloideae,
crasse-tunicatae; setulae ad apicem 4-8 × 0.5-1 μm,
cylindricae vel conicae, hyalinae vel pallide flavae,
pallide dextrinoideae, tenui- vel crasse-tunicatae.
Caulocystidia versiformae, proximus basi, cellulae 55-90
× 4-7 μm, cylindricae vel clavatae, apicaliter ramosa,
pallide flavae, dextrinoideae, tenui-tunicatae apice, basi
crasse-tunicatae. Fibulae presentes. Gregarius, ad folia
putrida plantarum dicotyledonearum. Holotypus:
Malaysia, Selangor, Selayang, Kanching Forest Reserve,
N 03° 17.958’, E 101°37.151’. 8 Jan 2005, collected by
Amy Honan, TYS 417 (Holotypus: KLU-M#87).
Pileus 7-21 mm diam, convex to planoconvex, without an umbo, disc rugulose,
57
Fig. 25. Marasmius musicolor (Holotype: TYS 417 = KLU-M#87). a. Basidiomes. b. Basidiospores. c. Basidioles. d.
Cheilocystidia. e. Pileipellis. f. Caulocystidia. Bars: a = 10 mm, b-f = 10 µm.
margin rugulose-striate; surface dry, dull,
hygrophanous, subvelutinous; light yellow
(3A5, 4A4-5) to banana-coloured overall.
58
Lamellae adnate, close (12-14) with 4 series of
lamellulae, narrow, sometimes forked, pale
yellow, non-marginate. Stipe 11-15 × 0.5-1 mm,
Fungal Diversity
central, cylindrical, terete, equal, dry, dull,
pruinose, non-insititious, pale yellow (3A3) at
apex, brownish orange (5C5. 6C6-7) at base,
with orangish white mycelium.
Basidiospores (13-) 15-18 (-19) × 3.5- 4
μm [x = 16.1 ± 1.7 × 3.9 ± 0.2 μm, Q = 3.7-5.1,
Qm = 4.9 ± 0.5, n = 10 spores], elongatelacrymoid to clavate, smooth, hyaline,
inamyloid. Basidia not observed. Basidioles
32-40 × 7 μm, fusoid to clavate. Cheilocystidia
numerous, composed of Siccus-type broom
cells; main body (6-) 10-22 (-26) × (5-) 7-8 (11) μm, cylindrical to clavate or subclavate,
hyaline, inamyloid, thin- to thick-walled; apical
setulae 2-6 (-7.5) × 0.5-1 μm, close, cylindrical
to conical, subacute to obtuse, rarely forked,
light yellow, inamyloid, thin- to thick-walled.
Pleurocystidia absent. Pileipellis weakly
mottled, composed of a hymeniform layer of
Siccus-type broom cells; main body (10-) 1419 (-20) × (5-) 6-10 (-11) μm, cylindrical to
clavate or broadly clavate, rarely lobed, hyaline,
inamyloid, thick-walled; apical setulae (2-) 4-8
× 0.5-1 μm, cylindrical to conical, rarely forked,
hyaline to light yellow, inamyloid to weakly
dextrinoid, thin- to thick-walled. Pileus trama
interwoven; hyphae 3-4 μm diam, cylindrical,
hyaline, strongly dextrinoid, thin-walled, nongelatinous. Lamellar trama regular; hyphae 3-6
μm diam, cylindrical, hyaline, strongly
dextrinoid, thin-walled, non-gelatinous. Stipe
tissue monomitic; cortical hyphae 3-6 μm diam,
parallel, cylindrical, smooth, brownish yellow,
dextrinoid, thick-walled; medullary hyphae 412 μm diam, parallel, cylindrical, smooth,
hyaline, strongly dextrinoid, thick-walled (up
to 1 μm), non-gelatinous. Caulocystidia
numerous near the base of stipe, 55-90 × 4-7
μm, cylindrical to clavate, branched and often
apically lobed, light yellow, strongly dextrinoid,
thin-walled apically, thick-walled at the base.
Clamp connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
twigs. Malaysia (Selangor).
Material examined: Malaysia, Selangor, Selayang,
Kanching Forest Reserve, N 03° 17.958’, E 101°37.151’.
8 Jan 2005, Amy Honan, TYS 417 (Holotype: KLUM#87; Isotype: SFSU).
Notes:
Marasmius
musicolor
is
characterized by the following features: a
convex to plano-convex, rugulose-striate pileus
7-21 mm diam that is coloured banana yellow
overall; non-marginate, close (12-14) lamellae
with 4 series of lamellulae; a pruinose stipe 1115 mm long with pale yellow apex and
brownish orange base; basidiospores 13-19 ×
3.5-4 µm with mean 16.1 × 3.9 µm; no
pleurocystidia; Siccus-type cheilocystidia with
setulae 2-6 µm long; Siccus-type pileipellis
broom cells with setulae 4-8 µm long;
numerous obtuse, cylindrical caulocystidia that
are often forked or lobed, and an absence of
Siccus-type caulocystidia; and growth on twigs.
In combination, these features are not similar to
any published species, but resemble those
present in M. iras described herein as new.
Marasmius iras differs, however, in forming a
brown pileus, smaller basidiospores (13-16 ×
3-4 µm with mean 14.1 × 3.7 µm), has less
branched or lobate caulocystidia, and grows on
leaves.
21. Marasmius ochropoides Y.S. Tan &
Desjardin, sp. nov.
(Fig. 26)
MycoBank: MB 512627
Etymology: ochropoides = like M. ochropus
Singer.
Pileus 5-6 mm latus, convexus, leves vel
striatulus, subvelutinus, pallide aurantio vel aurantiobrunneus. Lamellae adnatae, densae (12-18), angustae,
cremeae, haud marginatae. Stipes 11-21 × 0.5-1 mm,
cylindricus, pruinosus, haud insititius, apicaliter cremeus,
basim versus pallide aurantiacus, ad basim tomento
cremeo affixus. Basidiosporae 9-12 × 3-4 μm, clavatae
vel ellipsoideae, leves, hyalinae, inamyloideae. Basidia
25-28 × 6-7 μm, clavatae, 4-spora. Cheilocystidia typi
Sicci, 15-25 × 5-6 μm, cylindracea vel clavata, hyalina;
2-4 setulae ad apicem, 10-18 × 2-3 μm, cylindricae vel
conicae, obtusae, hyalinae, inamyloideae, tenui-tunicatae.
Pleurocystidia nulla. Pileipellis hymeniformis, typi Sicci,
cellulae 10-16 × 6-10 μm, cylindricae vel clavatae,
hyalinae, inamyloideae, crasse-tunicatae; setulae ad
apicem 9-18 × 0.5-1 μm, cylindricae vel conicae, fulvae
vel brunneae, inamyloideae, crasse-tunicatae. Caulocystidia 23-100 × 4-7 μm, cylindricae vel clavatae,
obtusae, hyalinae, dextrinoideae, crasse-tunicatae.
Fibulae presentes. Gregarius, ad folia putrida plantarum
dicotyledonearum. Holotypus: Malaysia, Selangor, Ulu
Gombak, University of Malaya’s Field Study Center, 29
Nov 2004, Yee-Shin Tan, TYS 384 (Holotypus: KLUM#89).
59
Fig. 26. Marasmius ochropoides (Holotype: TYS 384 = KLU–M#89). a. Basidiomes. b. Basidiospores. c. Basidia and
basidioles. d. Cheilocystidia. e. Pileipellis. f. Caulocystidia. Bars: a = 10 mm, b-f = 10 µm.
Pileus 5-6 mm diam, convex, without an
umbo, disc smooth, margin smooth to
striatulate; surface dry, dull, hygrophanous,
subvelutinous; light orange (5A4) to brownish
orange (6C7-8), light brown (6D8) or reddish
brown (8D8). Lamellae adnate, close (12-18)
with 2 series of lamellulae, narrow, white to
yellowish white, non-marginate. Stipe 11-21 ×
0.5-1 mm, central, cylindrical, terete, equal, dry,
dull, pruinose, non-insititious, white (4A2) at
the apex, pale orangish white (5A3) at the base,
with yellowish white mycelium.
Basidiospores (8-) 9-12 (-13) × 3-4 μm
[xmr = 10.5-10.6 × 3.6-3.7 μm, xmm = 10.6 ± 0.1
× 3.7 ± 0.1 μm, Q = 2.5-3.7, Qmr = 2.9-3, Qmm
= 2.9 ± 0.1, n = 23 spores per 3 specimens],
clavate to ellipsoid, smooth, hyaline, inamyloid.
Basidia 25-28 × 6-7 μm, clavate, 4-spored.
Basidioles 25-32 × 5-8 μm, fusoid to clavate.
Cheilocystidia numerous, composed of Siccustype broom cells; main body (10-) 15-25 (-30)
× 5-6 (-8) μm, cylindrical to clavate, hyaline,
inamyloid, thin-walled; apical setulae only two
to four per cell, distant, (9-) 10-18 (-20) × 2-3
60
μm, cylindrical to conical, obtuse, rarely forked,
hyaline, inamyloid, thin-walled. Pleurocystidia
absent. Pileipellis weakly mottled, composed
of a hymeniform layer of Siccus-type broom
cells; main body 10-16 (-20) × (4-) 6-10 μm,
cylindrical to clavate or broadly clavate, rarely
lobed, hyaline, inamyloid, thick-walled; apical
setulae (6-) 9-18 (-25) × 0.5-1 (-1.5) μm,
cylindrical to conical, rarely forked, yellow to
brownish yellow, inamyloid, thick-walled.
Pileus trama sub-parallel to interwoven;
hyphae 4-6 μm diam, cylindrical, hyaline,
strongly
dextrinoid,
thin-walled,
nongelatinous. Lamellar trama regular; hyphae 5-9
μm diam, cylindrical, hyaline, dextrinoid to
strongly
dextrinoid,
thin-walled,
nongelatinous. Stipe tissue monomitic; cortical
hyphae 4-7 μm diam, parallel, cylindrical,
smooth, light yellow (apex) to yellow (base),
weakly dextrinoid to dextrinoid, thick-walled (1 μm); medullary hyphae 5-11 μm diam,
parallel,
cylindrical,
smooth,
hyaline,
dextrinoid,
thick-walled,
non-gelatinous.
Caulocystidia common, numerous, 23-100+ ×
Fungal Diversity
4-7 μm diam, cylindrical to clavate, rarely
lobed, obtuse, hyaline, dextrinoid, thick-walled
(-0.5 μm) or apically thin-walled. Clamp
connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves. Malaysia (Selangor, Selayang).
Material
examined:
Malaysia,
Selangor,
University of Malaya’s Field Study Center, N
03˚19.500’, E 101˚45.167’, elev. ca 240 m, 29 Nov 2004,
Yee-Shin Tan, TYS 384 (Holotype: KLU-M#89;
Isotype: SFSU); Selayang, Kanching Forest Reserve, N
03° 17.958’, E 101°37.151’. 8 Jan 2005, Amy Honan,
TYS 416 (KLU-M#88, SFSU); same location, 8 Jan
2005, Yee-Shin Tan, TYS 351 (KLU-M#134, SFSU).
Notes: Marasmius ochropoides is
characterized by a smooth pileus ranging in
colour from pale reddish brown to brownish
orange or orange when fresh but soon fades to
light orange overall; close (12-18) lamellae; a
subglabrous stipe coloured white above and
pale orangish white at the base; basidiospores
9-12 × 3-4 µm with mean 10.6 × 3.7 µm;
unusual cheilocystidia that have only 2-4
conical, hyaline setulae up to 20 µm long; and
scattered, thin- to thick-walled, hyaline, obtuse,
cylindrical caulocystidia. The new species is
phenetically closest to M. ochropus Singer
from Africa, but the latter species differs in
forming typical Siccus-type cheilocystidia with
numerous, short (1-7 µm) setulae, and lacks
caulocystidia (Pegler, 1977; Antonín, 2007). It
is also similar to M. ochroleucus Desjardin & E.
Horak from New Caledonia, but that species
also has typical Siccus-type cheilocystidia with
numerous, shorter (1-8 µm) setulae and has a
distinctly but minutely velutinous stipe with
numerous,
often
acutely
cylindrical
caulocystidia (Desjardin and Horak, 1997).
22. Marasmius iras. Y.S. Tan & Desjardin, sp.
nov.
(Fig. 27, Plate 2H)
MycoBank: MB512628
Etymology: iras (Malaysian) – resembling,
referring to its similarity to Marasmius musicolor.
Pileus 6-18 mm latus, convexus vel planoconvexus, ruguloso-striatus, subvelutinus, brunneus vel
aurantio-brunneus. Lamellae adnatae, densae (14-16),
haud intervenosae, angustae, albidae, haud marginatae.
Stipes 21-45 × 1 mm, cylindricus, pruinosus, haud
insititius, apicaliter stramineus, basim aurantio-brunneus,
ad
basim
tomento
aurantio-brunneo
affixus.
Basidiosporae 13-16 × 3-4 μm , clavatae, leves, hyalinae,
inamyloideae. Cheilocystidia typi Sicci, 12-16 × 4-7 μm,
cylindracea vel clavata, hyalina; setulae ad apicem 4-6
×0.5-1 μm, densae, cylindricae vel conicae, subacutae
vel acutae, fulvae, inamyloideae, tenui- vel crassetunicatae. Pleurocystidia nulla. Pileipellis hymeniformis,
typi Sicci, cellulae 9-12 × 6-8 μm, cylindricae vel
clavatae, pallide fulvae vel brunneae, inamyloideae,
crasse-tunicatae; setulae ad apicem 4-6 × 0.5-1 μm,
cylindricae vel conicae, subacutae, brunneae,
inamyloideae, crasse-tunicatae. Caulocystidia 30-50 × 58 μm, cylindricae vel clavatae, apicaliter diverticulata,
hyalinae vel stramineae, crasse-tunicatae. Fibulae
presentes. Gregarius, ad folia putrida plantarum
dicotyledonearum. Holotypus: Malaysia, Selangor, Ulu
Gombak, University of Malaya’s Field Study Center, 31
May 2005, Yee-Shin Tan, TYS 483 (Holotypus: KLUM#73).
Pileus 6-18 mm diam, convex when
young, becoming broadly convex to planoconvex in age; disc depressed, smooth; margin
rugulose-striate; surface dry, dull, hygrophanous, subvelutinous; brown (6E7) with dark
brown (8E7) disc when young, in age
becoming brownish orange (5B5) to grayish
orange with a brown (6E7) disc. Lamellae
adnate, close to crowded (14-16) with 3-4
series of lamellulae, not intervenose, narrow,
light yellow (3A4) with concolorous or pale
brown edge. Stipe 21-45 × 1 mm, central, terete,
cylindrical, equal, dry, dull, pruinose, noninsititious, yellowish white (4A2) at the apex,
gradually grading to light brown or brownish
orange at the base, with strigose brownish
orange basal mycelium. Odor pleasant.
Basidiospores (10-) 13-16 × 3-4 (-4.5)
μm [x = 14.1 ± 1.6 × 3.7 ± 0.4 μm, Q = 2.6-5.3,
Qm = 3.8 ± 0.6, n = 31 spores], elongatelacrymoid to clavate, smooth, hyaline,
inamyloid, thin-walled. Basidia not observed.
Basidioles (21-) 26-29 (-38) × 5-8 (-9) μm,
fusoid to clavate. Cheilocystidia numerous,
composed of Siccus-type broom cells; main
body (7-) 12-16 (-17) × (3-) 4-7 (-9) μm,
cylindrical to clavate or broadly clavate, rarely
lobed, hyaline, inamyloid, thick-walled; apical
setulae 4-6 (-7) × 0.5-1 μm, crowded,
cylindrical to conical, subacute to acute, rarely
forked, yellow, inamyloid, thin- to thick-walled.
Pleurocystidia absent. Pileipellis weakly
mottled, composed of a hymeniform layer of
Siccus-type broom cells; main body (7-) 9-12 (16) × 6-8 μm, cylindrical to clavate or broadly
clavate, rarely lobed, light yellow to light
brown, inamyloid, thick-walled; apical setulae
61
Fig. 27. Marasmius iras (Holotype: TYS 483 = KLU-M#73). a. Basidiomes. b. Basidiospores. c. Basidioles. d.
Cheilocystidia. e. Pileipellis. f. Caulocystidia. Bars: a = 10 mm, b-f = 10 µm.
4-6 (-7) × 0.5-1 μm.,cylindrical to conical,
rarely forked, subacute, brownish yellow to
brown, inamyloid, thick-walled. Pileus trama
interwoven; hyphae 2-4 μm diam, cylindrical,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Lamellar trama regular; hyphae 3-4 μm diam,
cylindrical, hyaline, strongly dextrinoid, thinwalled, non-gelatinous. Stipe tissue monomitic;
62
cortical hyphae 4-5 μm diam, parallel,
cylindrical, smooth, brownish yellow (apex) to
brown (base), weakly dextrinoid, thick-walled;
medullary hyphae 3-9 μm diam, parallel,
cylindrical, smooth, hyaline, dextrinoid, thinwalled. Caulocystidia scattered in clusters, (25-)
30-50 (-65) × (4-) 5-8 (-9) μm, irregularly
cylindrical to clavate, often apically split or
Fungal Diversity
diverticulate, hyaline to light yellow, inamyloid,
thick-walled. Clamp connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves and twigs. Malaysia (Selangor).
Material examined: Malaysia, Selangor, Ulu
Gombak, University of Malaya’s Field Study Centre, 31
May 2005, Yee-Shin Tan, TYS 483 (Holotype: KLUM#73; Isotype: SFSU).
Notes: Marasmius iras is characterized
by the following suite of features: a convex to
plano-convex pileus 6-18 mm diam with a
depressed, smooth, dark brown disc and
rugulose-striate, dull brown margin; close (1416) lamellae with 3-4 series of lamellulae
coloured yellowish white with concolorous or
pale brown edges; a stipe 20-45 × 1 mm that is
pruinose overall with a pale yellowish white
apex and brownish orange base; basidiospores
12-16 × 3-4.5 µm with mean 14.1 × 3.7 µm; no
pleurocystidia; Siccus-type cheilocystidia and
pileipellis broom cells with setulae 4-6 µm
long;
cylindrical,
obtuse,
non-setulose
caulocystidia and an absence of Siccus-type
broom cells on the stipe; and growth on
dicotyledonous leaves. The new species is
similar to M. convoluticeps Singer from Brazil,
and M. musicolor from Malaysia. Marasmius
convoluticeps differs in forming a convoluted,
gyrose, venose-rugulose pileus coloured bright
ferruginous, a dark brown stipe, broom cell
setulae 4.5-12 µm long, and grows on dead
wood (Singer, 1976). Marasmius musicolor
differs in forming a banana yellow pileus,
slightly longer basidiospores (mean 16.1 × 3.9
µm), more highly branched caulocystidia, and
grows on twigs.
Marasmius sect. Sicci Singer, subsect. Siccini
Singer, ser. Leonini Singer, Fl. Neotrop.
Monogr. 17: 160. 1976.
Type species: Marasmius leoninus Berk.
23. Marasmius aff. leoninus Berk., Hooker’s
J. Bot. 8: 135. 1856.
(Fig. 28, Plate 2 C)
Pileus 18-34 mm diam, convex to
broadly convex; disc depressed, rugulose;
margin striate to sulcate; surface dry, dull,
velutinous; pale orange (5A3) to pale yellow
(4A3) with a brown (6E7) disc. Lamellae
adnate, subdistant to close (13-15) with 3 series
of lamellulae, not intervenose, broad, yellowish
white (4A2), non-marginate. Stipe 22-35 × 12.5 mm, central, terete, equal, dry, dull,
glabrous, non-insititious, yellowish white (4A2)
at apex, dark brown at base, with light brown
basal mycelium. Odor pleasant.
Basidiospores (9-) 10-12 (-14) × 3-4 μm
[x = 11.4 ± 1.1 × 3.8 ± 0.4 μm, Q = 2.4-4, Qm =
3 ± 0.4, n = 26 spores], narrowly cylindrical to
clavate, smooth, hyaline, inamyloid, thinwalled. Basidia 28 × 7 μm, clavate, 4-spored.
Basidioles 25-30 × 5-7 μm, fusoid to clavate.
Cheilocystidia numerous, composed of Siccustype broom cells; main body 8-18 (-20) × (4-)
5-8 (-10) μm, cylindrical to clavate or irregular
in outline, rarely lobed, hyaline, inamyloid,
thin- to thick-walled; apical setulae (8-) 11-16
(-27) × 1-2 (-3) μm, narrowly cylindrical to
conical, rarely forked, light yellow, inamyloid,
thick-walled. Pleurocystidia absent. Pileipellis
mottled, composed of a hymeniform layer of
Siccus-type broom cells; main body 7-12 (-17)
× 5-8 (-10) μm, cylindrical to clavate or
irregular in outline, rarely lobed, hyaline,
inamyloid to weakly dextrinoid, thin- to thickwalled; apical setulae (5-) 6-12 (-17) × 1-1.5 (2) μm, narrowly conical, obtuse, rarely forked,
light yellow, inamyloid, thick-walled. Pileus
trama interwoven; hyphae 5-6 μm diam,
cylindrical, hyaline, strongly dextrinoid, thinwalled, non-gelatinous. Lamellar trama regular;
hyphae 5-8 μm diam, cylindrical, hyaline,
strongly
dextrinoid,
thin-walled,
nongelatinous. Stipe tissue monomitic; cortical
hyphae 5-8 μm diam, parallel, cylindrical,
smooth, light yellow (apex) to yellow (base),
dextrinoid,
thick-walled,
non-gelatinous;
medullary hyphae 3-5 μm, parallel, cylindrical,
smooth, hyaline, inamyloid, thin-walled.
Caulocystidia scattered, 5-26 (-40) × (1-) 2-5
μm, irregularly cylindrical to conical, some
apically forked and setulae-like, plus a few
scattered Siccus-type broom cells, hyaline to
light yellow, inamyloid, thick-walled. Clamp
connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves. Malaysia (Selangor).
Material examined: Malaysia, Selangor, Selayang,
Kanching Forest Reserve, N 03° 17.958’, E 101° 37.151’,
elev. 110 m, 9 Jan 2005, collected by D.E. Desjardin,
TYS 423 (KLU-M#78; SFSU).
63
Fig. 28. Marasmius aff. leoninus (TYS 466 = KLU-M#78). a. Basidiomes. b. Basidiospores. c. Basidia and basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a = 20 mm, b-e = 10 µm.
Notes: The Malaysian species is
characterized by the following features: a large
(18-34 mm diam) sulcate-depressed pileus with
brown disc and pale orangish white margin;
non-marginate, close to subdistant (13-15)
lamellae with 3 series of lamellulae; a glabrous
stipe 22-35 × 1-2.5 mm with yellowish white
apex and dark brown base; basidiospores 10-12
× 3-4 µm with mean 11.4 × 3.8 µm; no
pleurocystidia; Siccus-type cheilocystidia with
few setulae 8-27 × 1-3 µm; Siccus-type
pileipellis broom cells with setulae 6-17 × 11.5 µm; and scattered caulocystidia that are
simple to branched, acute, setulae-like
outgrowths in combination with a few Siccustype broom cells. Only one specimen is known
64
currently from Malaysia. It is closest to M.
leoninus Berk. from Brazil, but the latter
species forms fewer lamellae (7-12), has
slightly smaller basidiospores (8.8-10 × 2.8-3.4
µm), and cheilocystidia with more numerous
and much smaller setulae (3-8 µm long)
(Isotype FH!). Interestingly, the two taxa share
the same stipitipellis anatomy. Until further
specimens of this interesting taxon are
discovered in Malaysia, we tentatively
recognize the material as representing M.
leoninus.
24. Marasmius araucariae var. siccipes
Desjardin, Retn. & E. Horak, Sydowia 52: 173.
2000.
(Fig. 29, Plate 2E)
Fungal Diversity
Fig. 29. Marasmius araucariae var. siccipes (TYS 529 = KLU–M#41). a. Basidiomes. b. Basidiospores. c. Basidioles. d.
Cheilocystidia. e. Pileipellis. f. Caulocystidia. Bars: a = 15 mm, b-f = 10 µm.
Type: Indonesia, Java, West Java, Bogor
Botanical Garden, 12 Jan 1999, A Retnowati
124 (BO!).
Pileus 10-24 mm diam, convex to
campanulate when young, expanding to planoconvex or broadly convex with uplifted margin
at maturity; disc rugulose; margin smooth to
striate; surface dry, dull, velutinous; brownish
orange (6C8, 7C8) to light brown (6D8) with
darker disc. Context thin, orangish white.
Lamellae adnate, subdistant (13-16) with 3-5
series of lamellulae, not intervenose, pale
yellow (4A3) with brownish orange edges.
Stipe 20-50 × 1 mm, central, terete, equal,
tough, solid, dry, dull, glabrous to minutely
pruinose, non-insititious, orange white (5A2) at
the apex, grading to light yellow (4A3), grayish
orange (5B6), brownish orange (6C6-7) to
brown at the base, with grayish yellow (5B3)
basal mycelium. Odor pleasant.
65
Basidiospores (8-) 9-11 (-12) × 3-4 (-5)
μm [xmr = 9.2-10.5 × 3.1-3.6 μm, xmm = 9.6 ±
0.5 × 3.3 ± 0.3 μm, Q = 2.3-4.3, Qmr = 2.4-3.3,
Qmm = 3 ± 0.3, n = 34 spores per 3 specimens],
ellipsoid to fusoid, smooth, hyaline, inamyloid,
thin-walled. Basidia not observed. Basidioles
17-30 × 5-8 μm, fusoid. Cheilocystidia
numerous, composed of Siccus-type broom
cells; main body (5-) 6-18 (-22) × (3-) 5-8 μm,
clavate to broadly clavate, vesiculose or
irregular in outline, rarely lobed, hyaline to
light yellow, inamyloid, thick-walled; apical
setulae (2-) 3-13 × 0.5-1.5 μm, conical, acute,
rarely forked, hyaline to brownish yellow or
brown, inamyloid, thin to thick-walled.
Pleurocystidia absent. Pileipellis mottled,
composed of a hymeniform layer of Siccustype broom cells; main body (6-) 10-22 (-30) ×
5-8 (-10) μm, clavate to broadly clavate or
irregular in outline, rarely lobed, hyaline,
inamyloid, thick-walled; apical setulae 5-8 × 11.5 μm, conical, acute, light yellow to brown,
inamyloid, thick-walled. Pileus trama interwoven; hyphae 4-10 μm diam, cylindrical,
hyaline, weakly dextrinoid to dextrinoid, thinwalled, non-gelatinous. Lamellar trama regular;
hyphae 4-5 μm diam, cylindrical, hyaline to
light yellow, dextrinoid, thin-walled, nongelatinous. Stipe tissue monomitic; cortical
hyphae 3-8 μm diam, parallel, cylindrical,
smooth, brownish yellow to brown, inamyloid,
thick-walled,
non-gelatinous;
medullary
hyphae 3-5 μm, parallel, cylindrical, smooth,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Stipe vesture of two types of caulocystidia: a)
Siccus-type broom cells with main body 15-40
× 6-13 μm, scattered, uncommon, clavate to
cylindrical or irregular in outline, light yellow,
inamyloid, thick-walled; apical setulae 5-6 × 12 μm, cylindrical to conical, yellow to
brownish yellow, inamyloid, thick-walled; b)
non-setulose cells, 12-27 × 3-7 μm, cylindrical
to clavate, obtuse, yellow, inamyloid, thin- to
thick-walled. Clamp connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves. Indonesia (Java), Malaysia (Selangor),
Thailand.
Material examined: Malaysia, Selangor, Hulu
Langat, Sungai Chongkak Forest Reserve, 6 Jan 2006,
Yee-Shin Tan, TYS 529 (KLU-M#41, SFSU); same
66
location, 22 July 2005, Hong-Twu Chan, MY 38 (KLUM#42, SFSU); same location, 16 May 2005, Yee-Shin
Tan, TYS 463 (KLU-M#43, SFSU).
Notes: Distinctive features of this species
include: a smooth to weakly striate pileus 1024 mm diam coloured brown to brownish
orange; subdistant (13-16) lamellae with 3
series of lamellulae; a glabrous to minutely
pruinose stipe; basidiospores in the range 9-11
× 3-4 µm with mean 9.6 × 3.3 µm; Siccus-type
cheilocystidia with acute setulae 3-13 × 0.5-1.5
µm; Siccus-type pileipellis broom cells with
acute setulae 5-8 × 1-1.5 µm; two types of
caulocystidia, viz., smooth clavate cells and
small setulose broom cells; and growth on
dicotyledonous
leaves.
The
Malaysian
population differs from the single specimen
reported from Java only in having slightly
smaller basidiospores (9.6 × 3.3 µm v.s. 11.7 ×
3.8 µm, respectively).
25. Marasmius luteomarginatus Desjardin,
Retn. & E. Horak, Sydowia 52: 166. 2000.
(Fig. 30, Plate 2I)
Type: Indonesia, Java, West Java, Mt.
Halimun National Park, loop trail from
Cikaniki, 9 Jan. 1999, A. Retnowati 110 (BO!).
Pileus 3-9 mm diam, convex to broadly
convex, umbilicate; margin striate to sulcate;
surface dry, dull, minutely velutinous; orange
(6B7-8) when young, in age fading to light
orange (5A5) or grayish orange (5B5-6).
Lamellae adnate, subdistant (7-10) with 2
series of lamellulae, orangish white with
yellow edges. Stipe 2-3.5 × 1 mm, mostly
eccentric when young, in age central,
cylindrical, terete, equal, dry, dull, glabrous,
non-insititious, orangish white at the apex,
brownish orange (6C7-8) at the base, with
grayish orange basal tomentum. Odor none.
Basidiospores (14-) 16-20 × (3.5-) 4-5
μm [x = 17.9 ± 1.8 × 4.2 ± 0.4 μm, Q = 3.5-5.1,
Qm = 4.1 ± 0.9, n = 28 spores], elongatelacrymoid to clavate, hyaline, smooth,
inamyloid. Basidia not observed. Basidioles
25-36 × 7.5-10 μm, fusoid to clavate.
Cheilocystidia abundant, 25-40 × 4.8-8 μm,
irregularly cylindrical to sinuous, often forked
or with a broad lateral outgrowth, hyaline,
inamyloid, thin-walled, with reddish brown
resinous exudates soluble in 3% KOH.
Pleurocystidia absent. Pileipellis weakly
Fungal Diversity
Fig. 30. Marasmius luteomarginatus (TYS 477= KLU-M#3). a. Basidiomes. b. Basidiospores. c. Basidia. d.
Cheilocystidia. e. Pileipellis. f. Caulocystidia. Bars: a = 10 mm, b-f = 10 µm.
mottled to mottled, composed of a hymeniform
layer of Siccus-type broom cells; main body 820 × 8-10 (-13) μm, cylindrical to clavate,
broadly clavate or irregular in outline, rarely
lobed, light yellow, inamyloid to weakly
dextrinoid, thin- to thick-walled; apical setulae
2-6 × 0.5-1 μm, cylindrical to conical, rarely
forked, yellow, weakly dextrinoid, thick-walled.
Pileus trama subparallel; hyphae 4-6 μm diam,
cylindrical, hyaline, dextrinoid, thin-walled,
non-gelatinous. Lamellar trama regular;
hyphae 4-5 μm diam, cylindrical, hyaline to
light yellow, weakly dextrinoid, thin-walled,
non-gelatinous. Stipe tissue monomitic; cortical
hyphae 4-6.5 μm diam, parallel, cylindrical,
smooth, light yellow to light brown, weakly
dextrinoid to dextrinoid, thick-walled (-1 μm);
medullary hyphae 3-4 μm diam, parallel,
cylindrical, smooth, hyaline, weakly dextrinoid,
thin-walled. Caulocystidia scattered, composed
of two types of cells: a) rare Siccus-type broom
cells with main body 6-8 × 5-7 μm, cylindrical
to conical, hyaline, inamyloid, thin-walled;
apical setulae 2-3 × 1 μm, narrowly cylindrical
to clavate, obtuse to subacute, hyaline,
inamyloid, thin-walled; b) non-setulose cells
like the cheilocystidia, 12-22 × 3.2-6.5 μm,
clavate, hyaline, inamyloid, thin-walled. Clamp
connections present.
67
Habit, habitat and known distribution:
gregarious on leaves of palm. Indonesia (Java),
Malaysia (Selangor).
Material examined: Malaysia, Selangor, Pangsun,
Gunung Nuang Forest Reserve, N 03° 12.593’, E 101 °
52.575’, 319 m, 17 May 2005, Yee-Shin Tan, TYS 477
(KLU-M #3, SFSU).
Notes: This is the second known
specimen of M. luteomarginatus and extends
the range from Mt. Halimun, Java northward to
Selangor, Malaysia. The small orange
basidiomes with yellow-marginate lamellae
and short eccentric stipe are distinctive. The
thin-walled, irregularly cylindrical cheilocystidia, unusual in Marasmius, are presumably
exudative because dried specimens have
lamellar edges with dark reddish brown
resinous incrustations.
A bipolar (unifactorial) mating system
was reported for M. luteomarginatus by Tan et
al. (2007) based on Malaysian material.
26. Marasmius jasminodorus Wannathes,
Desjardin & Lumyong, Fungal Diversity 37
(This volume)
(Fig. 31)
Type: Thailand, Chiang Mai Prov., Mok
Fa waterfall on Hwy 1095, 1 August 2003, N.
Wannathes 067 (CMU!).
Pileus 2-11 mm diam, convex to
campanulate; margin smooth; surface dry, dull,
subvelutinous; brownish orange (6C8, 7C7-8)
to orangish red (8B6) with darker disc.
Lamellae adnate, close (14-16) with 2-3 series
of lamellulae, not intervenose, yellowish white
with orange edges. Stipe 16-17 × 0.5-1 mm,
central, terete, equal, tough, dry, dull, glabrous
to pruinose, non-insititious with brownish
orange basal mycelium; yellowish white at
apex and brownish orange at base. Odor
pleasant, floral, like jasmine tea.
Basidiospores (11-) 12-14 (-15) × (3.5-)
4-4.5 (-5) μm [x = 12.7 ± 1 × 4 ± 0.3 μm, Q =
2.7-3.6, Qm = 3.1 ± 0.2, n = 31 spores],
narrowly ellipsoid to clavate, smooth, hyaline,
inamyloid, thin-walled. Basidia not observed.
Basidioles 23-30 × 5-8 μm, fusoid.
Cheilocystidia numerous, composed of Siccustype broom cells; main body (7-) 9-20 (-30) ×
(6-) 7-10 (-11) μm, cylindrical, clavate to
broadly clavate, vesiculose or some with
68
irregular outline, rarely lobed, hyaline,
inamyloid, thick-walled; apical setulae (5-) 610 × 1-1.5 μm, acute to subacute, rarely forked,
light yellow, inamyloid, thin to thick-walled.
Pleurocystidia absent. Pileipellis strongly
mottled, composed of a hymeniform layer of
Siccus-type broom cells; main body (10-) 1122 (-29) × (5-) 6-11 μm, clavate to broadly
clavate or irregular in outline, rarely lobed,
light yellow, inamyloid, thin- to thick-walled;
apical setulae (5-) 6-10 (-12) × 0.5-1 μm, acute
to subacute, brown, inamyloid, thick-walled.
Pileus trama interwoven, hyphae 3-6 μm diam,
cylindrical, hyaline, strongly dextrinoid, thinwalled, non-gelatinous. Lamellar trama regular
to subparallel, hyphae 5-8 μm diam, cylindrical,
hyaline, strongly dextrinoid, thin-walled, nongelatinous. Stipe tissue monomitic; cortical
hyphae 4-9 μm diam, parallel, cylindrical,
smooth, light yellow (apex) to brownish yellow
(base), dextrinoid, thick-walled (1 μm), nongelatinous; medullary hyphae 6-11 μm, parallel,
cylindrical, smooth, hyaline, dextrinoid, thinwalled, non-gelatinous. Caulocystidia scattered,
composed of two types of cells: a) Siccus-type
broom cells similar to the cheilocystidia; b)
non-setulose cells 15-30 × 3-10 μm, common,
cylindrical or ventricose, some apically split or
diverticulate, light yellow, inamyloid, thinwalled. Clamp connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves. Malaysia (Kedah), Thailand.
Material examined: Malaysia, Kedah, Langkawi
Island, Matchinchang Forest Reserve, 1 September 2004,
Yee-Shin Tan, TYS 333 (KLU-M#138, SFSU).
Notes: Marasmius jasminodorus is
characterized by a relatively small, smooth,
brownish orange to orangish red pileus, close,
orange-marginate lamellae, a minutely
pruinose stipe with small, cylindrical
caulocystidia and Siccus-type broom cells,
relatively short basidiospores with mean length
12.7 µm, Siccus-type cheilocystidia and
pileipellis broom cells with acute setulae 6-10
mm long, an absence of pleurocystidia, and
growth on dicotyledonous leaves. The most
distinctive feature is a strong pleasant odor of
jasmine tea.
Fungal Diversity
Fig. 31. Marasmius jasminodorus (TYS 333 = KLU–M#138). a. Basidiomes. b. Basidiospores. c. Basidioles. d.
Cheilocystidia. e. Pileipellis. f. Caulocystidia. Bars: a = 10 mm, b-d = 10 µm.
27. Marasmius angustilamellatus Y.S. Tan &
Desjardin, sp. nov.
(Fig. 32, Plate 4.3A)
MycoBank: MB512629
Etylomology: angusti- (Latin) = narrow; lamellatus (Latin) = lamellae, referring to the very
narrow lamellae.
Pileus 18-50 mm latus, convexus vel planoconvexus, haud striatus, subvelutinus, fulvae-brunneus
vel pallide brunneus ad centrum ater. Lamellae adnatae,
densae (24-40), angustae, pallide brunneae, haud
marginatae. Stipes 27-63 × 1-2 mm, cylindricus, leves,
glabrus, haud insititius, apicaliter fulvae-brunneus,
basim atrobrunneus, ad basim tomento aurantio
strigosoque affixus. Basidiosporae 8-12 × 3.5-4 μm,
ellipsoideae vel subclavatae, hyalinae, inamyloideae.
Cheilocystidia typi Sicci, 10-16 × 7-11 μm, cylindracea
69
Plate 3. Marasmius s.s. in section Sicci. A: Marasmius angustilamellatus (x 0.5); B: Marasmius acerosus (x 1); C:
Marasmius olivascens (x 1); D: Marasmius persicinus (x 0.5); E: Marasmius haematocephalus (x 1); F: Marasmius
florideus (x 1); G: Marasmius abundans var. abundans (x 0.25); H: Marasmius distantifolius (x 2); I: Marasmius
adhaesus (x 0.5); J: Marasmius selangorensis (x 0.5); K: Marasmius abundans var. aurantiacus (x 0.5); L: Marasmius
aratus (x 1).
vel clavata, straminea; setulae ad apicem 3-8 × 1-2 μm,
conicae, subacutae vel obtusae, hyalinae vel stramineae,
inamyloideae, tenui- vel crasse-tunicatae. Pleurocystidia
nulla. Pileipellis hymeniformis, typi Sicci, cellulae 1522 × 8.5-11 μm, cylindricae, clavatae vel irregulare,
stramineae, inamyloideae, crasse-tunicatae; setulae ad
apicem 3-6 × 0.5-1 μm, cylindricae vel conicae,
subacutae, stramineae, inamyloideae, crasse-tunicatae.
Caulocystidia nulla. Fibulae presentes. Solitarius vel
gregarius ad folia putrida plantarum dicotyledonearum.
Holotypus: Malaysia, Selangor, Ulu Gombak,
70
University of Malaya’s Field Study Centre, 5 Sept 2005,
Yee-Shin Tan, TYS 524 (Holotypus: KLU-M#36).
Pileus 18-50 mm diam, convex when
young, becoming broadly convex to planoconvex in age; margin smooth (non-striate);
surface dull, dry, hygrophanous, subvelutinous;
yellowish brown (5E8, F8) to grayish orange
(5B4), brownish orange (5C5) or light brown
(5D5-7) with black disc. Lamellae adnate,
crowded (24-40) with 3-7 series of lamellulae,
Fungal Diversity
Fig. 32. Marasmius angustilamellatus (Holotype: TYS 524 = KLU–M#33). a. Basidiomes. b. Basidiospores. c.
Cheilocystidia. d. Pileipellis. Bars: a = 20 mm, b-d = 10 µm.
very narrow (1 mm), light brown, nonmarginate. Stipe 27-63 × 1-2 mm, central,
terete, equal, cylindrical, tough, wiry, dry, dull,
glabrous, hollow, non-insititious, yellowish
brown (5E5-8) at the apex, dark brown (6F8)
at the base, with strigose light orange basal
mycelium. Odor not distinctive.
Basidiospores 8-12 (-14) × 3.5-4 (-4.5)
μm [x = 9.9 ± 1.7 × 3.8 ± 0.4 μm, Q = 2-3.5,
Qm = 2.6 ± 0.3, n = 15 spores (observed in only
one specimen; other specimens yielded no
spores)], elongate-ellipsoid to subclavate,
smooth, hyaline, inamyloid, thin-walled.
Basidia not observed. Basidioles fusoid.
Lamellar edge sterile, with a broad band of
cheilocystidia; cheilocystidia composed of
Siccus-type broom cells; main body (7-) 10-16
× 7-11 μm, cylindrical to clavate or broadly
clavate, rarely lobed, light yellow, inamyloid,
thick-walled; apical setulae (2-) 3-8 (-10) × 1-2
μm, not crowded, conical, subacute to obtuse,
rarely forked, hyaline to light yellow,
inamyloid, thin to thick-walled. Pleurocystidia
absent. Pileipellis weakly mottled, composed
of a hymeniform layer of Siccus-type broom
cells; main body (10-) 15-22 (-26) × (7-) 8.5-11
(-12) μm, cylindrical to clavate, broadly clavate
or irregular in outline, rarely lobed, light
yellow, inamyloid to weakly dextrinoid, thickwalled; apical setulae (2-) 3-6 × 0.5-1 (1.5) μm,
cylindrical or conical, rarely forked, subacute,
light yellow, inamyloid to weakly dextrinoid,
thick-walled. Pileus trama interwoven; hyphae
2.5-5 μm diam, cylindrical, hyaline, strongly
dextrinoid,
thin-walled,
non-gelatinous.
Lamellar trama regular; hyphae 3-5 μm diam,
71
cylindrical, hyaline to light yellow, strongly
dextrinoid, thin-walled, non-gelatinous. Stipe
tissue monomitic; cortical hyphae 4-6 μm diam,
parallel, cylindrical, smooth, orangish yellow
(apex) to brownish yellow (base), weakly
dextrinoid,
thick-walled,
non-gelatinous;
medullary hyphae 5-10 μm diam, parallel,
cylindrical, smooth, light yellow, strongly
dextrinoid,
thick-walled,
non-gelatinous.
Caulocystidia absent. Clamp connections
present.
Habit, habitat and known distribution:
solitary and gregarious on undetermined
dicotyledonous leaves. Malaysia (Selangor).
Material examined: Malaysia, Selangor, Hulu
Langat, Sungai Chongkak Forest Reserve, 6 Jan 2006,
Dennis Desjardin, TYS 528 (KLU-M#37, SFSU);
Selangor, Ulu Gombak, University of Malaya’s Field
Study Centre, 5 Sept 2005, Yee-Shin Tan, TYS 524
(Holotype: KLU-M#36; Isotype: SFSU); Selangor,
Selayang, Kanching Forest Reserve, N 03° 17.958’, E
101° 37.151’, elev. 110m, 9 Jan 2005, Yee-Shin Tan,
TYS 437 (KLU-M#35, SFSU).
Notes: The new species is distinctive
because of the following characters: a relative
large (18-50 mm diam), smooth pileus
coloured yellowish brown with a black disc;
very crowded and very narrow, non-marginate
lamellae; basidiospores in the range 8-14 x 3.54.5 µm with mean 9.9 × 3.8 µm; short and
broad cheilocystidia (7-16 × 7-11 µm); an
absence of pleurocystidia and caulocystidia;
and growth on dicotyledonous leaves.
Marasmius angustilamellatus is most closely
similar to M. corrugatus (Pat.) P. Sydow from
the Greater Antilles, and M. xestocephalus
Singer and M. xestocephaloides Antonín from
Africa. Marasmius corrugatus differs in
forming a rugulose-corrugate pileus coloured
reddish cinnamon to chestnut brown, and is
New World in origin. Marasmius xestocephalus and X. xestocephaloides both differ in
forming prominent caulocystidia.
28. Marasmius olivascens Y.S. Tan &
Desjardin, sp. nov.
(Fig. 33, Plate 3C)
MycoBank: MB 512630
Etymology: olivascens (Latin) = becoming olive
coloured, referring to the pigmentation of the pileus.
Pileus 5-8 mm latus, campanulatus vel convexus,
papillatus, sulcatus vel plicatus, pruinosus, olivaceobrunneus vel olivaceus. Lamellae adnatae, subdistantes
(12-16), haud intervenosae, pallide stramineae,
olivaceo-marginatae. Stipes 35-56 × 0.5 mm,
cylindricus, glabrus, haud insititius, apicaliter
72
stramineus, basim atrobrunneus, ad basim tomento
aurantio-brunneo strigosoque affixus. Basidiosporae 1114 × 4-5.5 μm, ellipsoideae vel clavateae, leves,
hyalinae, inamyloideae. Basidia 30-38 × 6-8 μm,
clavatae, 4-spora. Cheilocystidia typi Sicci, 15-23 × 5-8
μm, cylindracea vel clavata, straminea; setulae ad
apicem 4-7 × 0.5-1 μm, densae, cylindricae vel conicae,
obtusae vel subacutae, stramineae, inamyloideae, tenuitunicatae. Pleurocystidia nulla. Pileipellis hymeniformis,
typi Sicci, cellulae 10-26 × 5-9 μm, cylindricae,
clavatae vel irregulare, stramineae, inamyloideae,
crasse-tunicatae; setulae ad apicem 5-9 × 0.5-1 μm,
cylindricae vel conicae, subacutae, pallide flavae vel
flavo-brunneae, inamyloideae, tenui- vel crassetunicatae. Caulocystidia nulla. Fibulae presentes.
Gregarius, ad folia putrida plantarum dicotyledonearum.
Holotypus: Malaysia, Selangor, Selayang, Kanching
Forest Reserve, 9 Jan 2005, Yee-Shin Tan, TYS 426
(Holotypus: KLU-M#90).
Pileus 5-18 mm diam, hemispherical to
campanulate or broadly convex with an acute
papilla; margin sulcate to plicate; surface dull,
dry, pruinose; olive brown (4F8) with darker
disc when young, in age becoming olive (2F78) overall. Lamellae adnate, subdistant (12-16)
with 0-1 series of lamellulae, not intervenose,
light yellow (4A2-3) with olive brown edge.
Stipe 35-56 × 0.5 mm, central, terete,
cylindrical, equal, dry, dull, glabrous, noninsititious, yellowish white (4A2) at the apex,
dark brown at the base, with strigose brownish
orange (6C8) to light brown (6D7-8)
mycelium. Odor not distinctive.
Basidiospores 11-14 (-15) × 4-5.5 (-6)
μm [xmr = 12.3-13.1 × 4.6-5.3 μm, xmm = 12.7 ±
0.6 × 5 ± 0.5 μm, Q = 1.9-3.5, Qmr = 1.9-2.8,
Qmm = 2.6 ± 0.4, n = 25 spores per 2
specimens], ellipsoid to clavate, smooth,
hyaline, inamyloid, thin-walled. Basidia 30-38
× 6-8 μm, clavate, 4-spored. Basidioles 30-40 ×
6-9 μm, fusoid to clavate. Cheilocystidia
numerous, composed of Siccus-type broom
cells; main body (12-) 15-23 (-27) × (4-) 5-8 (10) μm, cylindrical to clavate or broadly
clavate, rarely lobed, light yellow, inamyloid,
thin- to thick-walled; apical setulae (3-) 4-7 ×
0.5-1 (-1.5) μm, crowded, narrowly cylindrical
to conical, obtuse to subacute, rarely forked,
yellow, inamyloid, thin-walled. Pleurocystidia
absent. Pileipellis strongly mottled, composed
of a hymeniform layer of Siccus-type broom
cells; main body (8-) 10-26 (-33) × (4-) 5-9 (12) μm, cylindrical to clavate, broadly clavate
or irregular in outline, often lobed, light yellow,
Fungal Diversity
Fig. 33. Marasmius olivascens (Holotype: TYS 426 = KLU-M#90). a. Basidiomes. b. Basidiospores. c. Basidia and
basidiole. d. Cheilocystidia. e. Pileipellis. Bars: a = 20 mm, b-e = 10 µm.
inamyloid, thick-walled; apical setulae 5-9 (-10)
× 0.5-1 μm, cylindrical to conical, rarely forked,
subacute, light yellow to brownish yellow or
golden brown, inamyloid to weakly dextrinoid,
thin- to thick-walled. Pileus trama interwoven;
hyphae 5-9 μm diam, cylindrical, hyaline,
strongly
dextrinoid,
thin-walled,
nongelatinous. Lamellar trama regular; hyphae 410 μm diam, cylindrical, hyaline, dextrinoid,
thin-walled, non-gelatinous. Stipe tissue
monomitic; cortical hyphae 3-5 μm diam,
parallel, cylindrical, smooth, yellowish brown
(apex) to brown (base), inamyloid to weakly
dextrinoid,
thick-walled,
non-gelatinous;
medullary hyphae 6-9 μm, parallel, cylindrical,
smooth, hyaline, strongly dextrinoid, thickwalled, non-gelatinous. Caulocystidia absent.
Clamp connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves and twigs. Malaysia (Selangor).
Material examined: Malaysia, Selangor, Selayang,
Kanching Forest Reserve, 9 Jan. 2005, Yee-Shin Tan,
TYS 426 (Holotype: KLU-M#90; Isotype: SFSU); same
location, 9 Jan. 2005, Dennis Desjardin, TYS 424 (KLUM#91, SFSU).
Notes: Diagnostic features of M.
olivascens include: a plicate pileus 5-18 mm
diam that is coloured olive overall; distant (1273
16) lamellae with 0-1 series of lamellulae that
have olive edges; basidiospores 11-15 x 4-6
µm with mean 12.7 x 5 µm; Siccus-type
cheilocystidia with narrowly conical, acute
setulae 3-7 µm long; Siccus-type pileipellis
broom cells with narrowly conical, acute
setulae 5-9 µm long; no pleurocystidia and no
caulocystidia; and growth on dicotyledonous
leaves. The new species is closest to M.
trinitatis Dennis from the New World tropics
and Papua New Guinea. The latter species
differs in forming close lamellae with 2-3
series of lamellulae, smaller basidiospores in
the range 8.5-12.5 x 3-4 µm with mean about
10.5 x 3.3 µm, and grows mainly on woody
debris (Singer, 1976; Desjardin and Horak,
1997).
29. Marasmius florideus Berk. & Broome, J.
Linn. Soc. Bot. 14: 39. 1873.
(Fig. 34, Plate 3F)
Type: Sri Lanka, Kandy District,
Peradeniya, on dead wood, Sept 1897,
Thwaites 204 p.p. (K).
Pileus 3-6 mm diam, convex to broadly
convex, umbonate; margin smooth to striate;
surface dull, dry, velutinous; reddish brown
(8D7-8, 8E7-8) to brown (7E7-8). Lamellae
adnate, close to subdistant (10-12) with 2-3
series lamellulae, not intervenose, pale yellow
(4A3) with brown (7E8) edges. Stipe 16-33 ×
0.5-1 mm, central, terete, equal, tough, dry,
dull, glabrous, non-insititious, orangish white
at the apex, gradually becoming orange, to
grayish orange (6B5-6), golden yellow (5B7-8)
or brown (6E8) at the base, with strigose
grayish orange basal mycelium. Odor not
distinctive.
Basidiospores (8-) 9.6-12 (-13) × 3-4 μm
[x = 10.4 ± 2.2 × 3.5 ± 0.4 μm, Q = 2-3.6, Qm =
3 ± 0.4, n = 22 spores], narrowly ellipsoid to
fusoid, clavate, smooth, hyaline, inamyloid,
thin-walled. Basidia not observed. Basidioles
17.6-19.2 × 4-4.8 μm, fusoid to clavate.
Cheilocystidia numerous, composed of Siccustype broom cells; main body (4.8-) 7.3-11.5 (12.8) × 4.8-6.4 (-8) μm, cylindrical to clavate,
broadly clavate or irregular in outline, rarely
lobed, hyaline, inamyloid, thick-walled; apical
setulae (2-) 3.2-7 (-10) × 1-1.5 μm, narrowly
cylindrical to conical, rarely forked, yellow,
inamyloid, thick-walled. Pleurocystidia absent.
74
Pileipellis weakly mottled, composed of a
hymeniform layer of Siccus-type broom cells;
main body 6.4-12.8 × 4.2-6.4 μm, cylindrical to
clavate, broadly clavate or irregular in outline,
rarely lobed, hyaline to pale yellow, inamyloid,
thick-walled; apical setulae (2-) 5-7 × 1 μm,
narrowly cylindrical to conical, rarely forked,
subacute, yellow to yellowish brown,
inamyloid,
thick-walled.
Pileus
trama
interwoven; hyphae 5.4-6.4 μm diam,
cylindrical, hyaline, strongly dextrinoid, thinwalled, non-gelatinous. Lamellar trama regular;
hyphae 5.1-6.4 μm diam, cylindrical, hyaline,
strongly
dextrinoid,
thin-walled,
nongelatinous. Stipe tissue monomitic; cortical
hyphae 3.2-4.8 μm diam, parallel, cylindrical,
smooth, light yellow to yellow, weakly
dextrinoid to dextrinoid, thick-walled, nongelatinous; medullary hyphae 4-5.4 μm diam,
parallel,
cylindrical,
smooth,
hyaline,
inamyloid to weakly dextrinoid, thick-walled,
non-gelatinous. Caulocystidia absent. Clamp
connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves. Malaysia (Selangor), Sri Lanka.
Material examined: Malaysia, Selangor, Pangsun,
Mount Nuang Forest Reserve, N 03° 12.593’, E 101 °
52.575’, 319 m, 17 May 2005, Yee-Shin Tan, TYS 480
(SFSU).
Notes: Marasmius florideus Berk. &
Broome, described from Sri Lanka, is
characterized by a smooth to striate, conicalsubumbonate, dark reddish brown pileus;
subdistant (10-12) lamellae with reddish brown
edges; a glabrous stipe lacking caulocystidia;
basidiospores in the range 8-12 x 3-4.5 µm
(mean 10.4 x 3.5 µm); no pleurocystidia; and
cheilocystidia and pileipellis broom cells with
narrowly conical, acute setulae 3-10 µm long.
30. Marasmius acerosus Y.S. Tan &
Desjardin, Fungal Diversity 25: 192. 2007.
(Fig. 35, Plate 3B)
Type: Malaysia, Selangor, Sungai
Chongkak Forest Reserve, 16 May 2005, Y.S.
Tan #458 (KLU-M #2!).
Pileus 4-15 mm diam, conical to convex
with a small umbo when young, becoming
broadly convex to plano-convex in age; margin
smooth to weakly striatulate in age; surface dry,
dull, glabrous; grayish yellow (4B4) with
Fungal Diversity
Fig. 34. Marasmius florideus (TYS 480). a. Basidiomes. b. Basidiospores. c. Basidioles. d. Cheilocystidia. e. Pileipellis.
Bars: a = 10 mm, b-e = 10 µm.
brownish orange (5C6) disc when young, in
age becoming grayish yellow (5B5) to golden
brown with a brown (6E7) disc, or grayish
yellow (4B6) with a brownish orange (5B5)
disc. Lamellae adnate, crowded (16-25) with 34 series of lamellulae, light yellow (3A4), nonmarginate. Stipe 15-34 × 0.5 mm, central, terete,
equal, dry, dull, glabrous, non-insititious,
yellowish white (4A2) at the apex, gradually
grading to brownish orange (5C6) or light
brown (6D7-8) at the base, with strigose pale
orange (5A3) basal mycelium. Odor pleasant.
Basidiospores 13-15 (-15.5) × 4-4.5 (-5)
μm [xmr = 14.3-14.4 × 4.4-4.7 μm, xmm = 14.4 ±
0.1 × 4.5 ± 0.2 μm, Q = 2.9-3.8, Qmr = 3.1-3.3,
Qmm = 3.2 ± 0.2, n = 18 spores per 2
specimens], elongate-lacrymoid to clavate,
smooth, hyaline, inamyloid, thin-walled.
Basidia not observed. Basidioles 23-36 × 6-8
μm, fusoid to clavate. Lamellar edge sterile,
with a broad band of cheilocystidia;
cheilocystidia numerous, composed of Siccustype broom cells; main body (8-)12-24 × 6-10
μm, cylindrical to clavate, broadly clavate or
irregular in outline, rarely lobed, hyaline,
inamyloid, thick-walled; apical setulae 2-10 ×
0.5-1 μm, crowded, conical to acerose,
subacute to acute, rarely forked, light yellow,
inamyloid, thick-walled. Pleurocystidia absent.
Pileipellis weakly mottled, composed of a
hymeniform layer of Siccus-type broom cells;
main body (8-) 10-25 × 6-13 μm, cylindrical to
clavate, broadly clavate or irregular in outline,
rarely lobed, light yellow, inamyloid, thickwalled; apical setulae 6-13 × 0.5-1 μm, conical
to acerose, rarely forked, acute, yellow to
yellowish brown, inamyloid, thick-walled.
Pileus trama interwoven; hyphae 4-9 μm diam,
cylindrical, hyaline, weakly dextrinoid to
dextrinoid,
thin-walled,
non-gelatinous.
Lamellar trama regular; hyphae 3.5-5 μm diam,
cylindrical, hyaline, dextrinoid to strongly
dextrinoid, thin-walled, non-gelatinous. Stipe
tissue monomitic; cortical hyphae 4.5-6 μm
diam, parallel, cylindrical, smooth, yellow
(apex) to brownish yellow (base), inamyloid to
weakly dextrinoid, thick-walled (-1 μm), nongelatinous; medullary hyphae 5-9 μm diam,
75
Fig. 35. Marasmius acerosus (Holotype: TYS 458 = KLU-M#2). a. Basidiomes. b. Basidiospores. c. Basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
parallel, cylindrical, smooth, hyaline to light
yellow, dextrinoid, thin-walled. Caulocystidia
absent. Clamp connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves and twigs. Malaysia (Selangor).
Material
examined:
Malaysia,
Selangor,
Kanching Forest Reserve, N 03°171.958’, E 101° 37.15’,
110 m elev., 9 Jan 2005, Yee-Shin Tan, TYS 427 (KLUM#29, SFSU); Selangor, Sungai Chongkak Forest
Reserve, 16 May 2005, Yee-Shin Tan, TYS 458
(Holotype: KLU-M #2; Isotype: SFSU).
Notes:
Marasmius
acerosus
is
characterized by the following features: a
smooth to weakly striatulate pileus coloured
brownish orange on the disc and grayish
yellow on the margin; crowded lamellae (16-
76
25 with 3-4 series of lamellulae) with nonmarginate edges; basidiospores in the range
13-15 × 4-4.5 μm with mean 14.4 × 4.5 μm;
Siccus-type cheilocystidia and pileipellis
broom cells with needle-like setulae up to 10
µm long or longer; and growth on
dicotyledonous leaves. The new species is
similar to M. abundans Corner and its varieties
from Malaysia, and to M. berteroi (Lév.)
Murrill from Java and the neotropics.
Marasmius abundans differs in forming fewer
lamellae (ca 14-19), slightly longer basidiospores (mean length 15.4-16.2 μm), and
cheilocystidia with cylindrical to subconical,
obtuse setulae only up to 7 µm long (Corner,
1996; type studies and this paper). Marasmius
Fungal Diversity
Fig. 36 Marasmius bambusiniformis (TYS 511). a. Basidiomes. b. Basidiospores. c. Basidioles. d. Cheilocystidia. e.
Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
berteroi differs in forming plicate pilei,
subdistant lamellae (11-16), and cheilocystidia
with obtuse to subacute setulae up to 5 μm long
(Desjardin et al., 2000).
A bipolar (unifactorial) mating system
was reported for M. acerosus by Tan et al.
(2007).
31. Marasmius bambusiniformis Singer, Fl.
Neotrop. Monogr. 17: 167. 1976.
(Fig. 36)
Type: Ecuador, Napo, Lago Agrio, 16
May 1973, Singer B7480 (F!).
Pileus 1.5-5 mm diam, convex to
obtusely conical; margin smooth when young,
striate to sulcate in age; surface dull, dry, subvelutinous; orange (6A7, 6B7-8) to reddish
orange (7B8). Lamellae adnate, subdistant (1213) with 0-1 series of lamellulae, not
intervenose, yellowish white, non-marginate.
Stipe 10-30 × 0.5-1 mm, central, terete, equal,
tough, dry, dull, glabrous, non-insititious,
yellowish white at the apex, grading to
brownish orange or reddish brown (6E8) at the
base, with strigose grayish orange basal
mycelium. Odor not distinctive.
Basidiospores (15-) 16-18 (-19) × 3-3.5
(-4) μm [x = 16.9 ± 1.3 × 3.4 ± 0.5 μm, Q =
3.8-6.3, Qm = 5 ± 0.6, n = 27 spores], narrowly
ellipsoid to fusoid or clavate, smooth, hyaline,
inamyloid, thin-walled. Basidia not observed.
Basidioles 22-26 × 6-7 μm, fusoid to clavate.
Cheilocystidia numerous, composed of Siccustype broom cells; main body 13-16 × 5-7 μm,
cylindrical to clavate, broadly clavate or
irregular in outline, rarely lobed, light yellow,
inamyloid, thick-walled; apical setulae 3-5.5 ×
0.5-1 μm, narrowly cylindrical to conical,
rarely forked, light yellow, inamyloid, thin- to
thick-walled. Pleurocystidia absent. Pileipellis
mottled, composed of a hymeniform layer of
Siccus-type broom cells; main body 10-20 × 38 (-10) μm, cylindrical to clavate, broadly
clavate or irregular in outline, rarely lobed,
hyaline, inamyloid, thick-walled; apical setulae
4-6 (-7) × 0.5-1 μm, narrowly cylindrical to
conical, rarely forked, subacute, hyaline to
77
brownish yellow or dull yellow, inamyloid,
thick-walled. Pileus trama interwoven; hyphae
4-5 μm, cylindrical, hyaline, weakly dextrinoid,
thin-walled, non-gelatinous. Lamellar trama
regular; hyphae 3-5 μm diam, cylindrical,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Stipe tissue monomitic; cortical hyphae 3-5 μm
diam, parallel, cylindrical, smooth, brownish
yellow,
inamyloid,
thick-walled,
nongelatinous; medullary hyphae 4-7 μm diam,
parallel, cylindrical, smooth, hyaline, weakly
dextrinoid,
thick-walled,
non-gelatinous.
Caulocystidia absent. Clamp connections
present.
Habit, habitat and known distribution:
solitary and gregarious on undetermined
dicotyledonous leaves. South America (Brazil,
Ecuador), Malaysia (Johore), Papua New
Guinea, Unitd States (Florida).
Material examined: Malaysia, Johore, Endau
Rompin National Park, way to NERC, Peta village,
Temiang trail, 14 July 2005, Yee-Shin Tan, TYS 511
(SFSU).
Notes: Marasmius bambusiniformis has
diagnostic features that include a small,
obtusely conical to convex, smooth to striate,
orange to reddish orange pileus, subdistant,
narrow lamellae, a glabrous stipe lacking
caulocystidia, moderately long basidiospores
(16-18 × 3.3.5 µm with mean 16.9 × 3.4 μm),
and growth on dicotyledonous leaves. The
Malaysian material matches nicely specimens
reported from Papua New Guinea (Desjardin
and Horak, 1997), and differs subtly from
South American material in having more
lamellae (12-13 versus 7-10 in the holotype @
F!).
32. Marasmius abundans Corner var.
abundans, Beih. Nova Hedwigia 111: 22. 1996.
(Fig. 37 & 38, Plate 3G)
Type: Singapore, Gardens Jungle, 15 Oct.
1929, Corner s.n. (E #205879).
Pileus 8-17 mm diam, obtusely conical to
convex or broadly convex; margin smooth
(non-striate); surface dry, dull, hygrophanous,
minutely velutinous; grayish orange (5B6) to
golden yellow (5B7), orange (5A7), light
orange (5A5), brownish yellow (5C7-8) or
ferruginous. Lamellae adnate, close (14-17)
with 4-5 series of lamellulae, yellowish white,
non-marginate. Stipe 32-45 × 0.5-1 mm, central,
78
cylindrical, terete, equal, dry, dull, glabrous,
non-insititious, yellowish white at the apex,
gradually changing to orange, deep orange,
yellowish brown (5E8) or brownish red (8C7-8)
at the base, with brownish orange basal
mycelium.
Basidiospores 12-18 (-20) × 4-5 μm [xmr
= 15.8-16.6 × 4.1-4.6 μm, xmm = 16.2 ± 0.6 ×
4.4 ± 0.4 μm, Q = 3.8-4.5, Qmr = 3.4-4.1, Qmm =
3.8 ± 0.5, n = 17 spores per 9 specimens],
elongate-lacrymoid to clavate, smooth, hyaline,
inamyloid. Basidia 24-28 × 6-10 μm, 4-spored.
Basidioles 26-28 (-32) × 6-7 μm, fusoid to
clavate. Cheilocystidia numerous, lamellar
edge sterile, composed of Siccus-type broom
cells; main body (10-) 11-20 (-22) × 6-10 (-11)
μm, cylindrical to clavate or broadly clavate,
hyaline, inamyloid, thin-walled; apical setulae
(2-) 2.5-4 (-5) × 0.5-1 μm, crowded, cylindrical
to conical, subacute to obtuse, light yellow to
yellow, inamyloid, thick-walled. Pleurocystidia
absent. Pileipellis weakly mottled, composed
of a hymeniform layer of Siccus-type broom
cells; main body (7-) 10-13 (-16) × (5-) 6-11 (12) μm, cylindrical to clavate or subglobose,
hyaline, inamyloid, thick-walled; apical setulae
3-7 (-8) × 0.5-1 μm, cylindrical to conical,
yellow, inamyloid, thick-walled. Pileus trama
interwoven; hyphae 5-10 μm diam, cylindrical,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Lamellar trama regular; hyphae 3.5-6 μm diam,
cylindrical, hyaline, dextrinoid, thin-walled,
non-gelatinous. Stipe tissue monomitic; cortical
hyphae 4-5 μm diam, parallel, cylindrical,
smooth, light yellow (apex) to yellow (base),
weakly
dextrinoid,
thick-walled,
nongelatinous; medullary hyphae 3-8 μm diam,
parallel,
cylindrical,
smooth,
hyaline,
dextrinoid, thick-walled. Caulocystidia absent.
Clamp connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves. Malaysia (Selangor), Singapore.
Material examined: Malaysia, Johore, Endau
Rompin National Park, NERC, Peta Village, on way to
jetty, 14 July 2005, Yee-Shin Tan, TYS 513 (KLUM#28, SFSU); Kedah, Langkawi Island, Matchinchang
Forest Reserve, 1 Sept. 2004, Yee-Shin Tan, TYS
344(KLU-M#79, SFSU); Selangor, Sungai Chongkak
Forest Reserve, N 03°12.705’, E 101°50.472’, elev. 122
m, 16 May 2005, Yee-Shin Tan, TYS 460
Fungal Diversity
Fig. 37 Marasmius abundans var. abundans (TYS 460 = KLU-M#27). a. Basidiomes. b. Basidiospores. c. Basidia and
basidioles. d. Cheilocystidia. e. Pileipellis. Bars: a = 20 mm, b-e = 10 µm.
(KLU-M#27, SFSU), same location, 1 Jan. 2006, YeeShin Tan, TYS 530 (KLU-M#81, SFSU); Selangor,
Selayang, Kanching Forest Reserve, N 03°17.958’, E
101°37.151’, elev. 110 m, 9 Jan 2005, Yee-Shin Tan,
TYS 425 (KLU-M#24, SFSU); Singapore, Gardens
Jungle, 15 Oct. 1929, Corner s.n. (Holotype: E #205879).
Notes: Marasmius abundans var.
abundans has only been reported from Malesia
(Corner, 1996). This species is distinguished by
a moderately large, grayish orange to light
orange, smooth pileus, close (14-17) lamellae,
a glabrous stipe, and relatively large
basidiospores (12-18 × 4-4.5 µm). It has been
reported as very common on the Malay
Peninsula, hence the choice of epithets (Corner,
1996). The difference between M. abundans
var. abundans and M. abundans var.
aurantiacus is indicated in the key.
Type Study of Marasmius abundans var.
abundans:
The holotype specimen (Singapore,
Gardens Jungle, 15 Oct. 1929, Corner s.n., E
#205879) is a mixture of two species,
comprising approx. 8 basidiomes, whole or
fragmented, in fair condition, attached to dicot
leaves: Species 1, herein recognized as the
holotype specimen. As dried: Pileus 5-7 mm
diam, convex, margin even and not striate or
very short-striatulate, ferruginous to reddish
brown. Lamellae adnexed, close, narrow. Stipe
25-30 × 1 mm, terete, cylindrical, glabrous,
non-insititious, with rusty basal mycelium.
Basidiospores 11-14 × 3.8-4.5 µm, subfusoid
to clavate, smooth, hyaline, inamyloid, thinwalled. Basidia not observed. Basidioles
subfusoid, sometimes mucronate. Cheilocystidia abundant, edge sterile, composed of
Siccus-type broom cells; main body 12-16 × 510 µm, subcylindrical to clavate, rarely lobed,
hyaline, thin-walled; apical setulae 3.2-6.5 × 11.5 µm, conical, acute, thin- to thick-walled,
hyaline
to
pale
yellow,
inamyloid.
Pleurocystidia absent. Pileipellis not mottled, a
hymeniform layer of Siccus-type broom cells;
main body 9.5-15 × 5-8 µm, subcylindrical to
79
Fig. 38. Marasmius abundans var. abundans (Holotype: E#205879). a. Basidiospores. b. Cheilocystidia. c. Pileipellis.
Bar = 10 µm.
clavate, basally hyaline and thin-walled,
apically yellowish brown and thick-walled;
apical setulae 3-6.5 × 1-1.5 µm, conical, acute,
thin-walled to thick-walled, yellowish brown,
weakly dextrinoid. Stipe tissue monomitic;
cortical hyphae parallel, cylindrical, smooth,
pale tawny, dextrinoid, thin-walled; medullary
hyphae similar but hyaline. Caulocystidia
absent. Clamp connections present.
Species 2: similar to species one except
that the pileus is striate-sulcate to the disc, and
the lamellae are distant and broad. This
portion of the holotype specimen will not be
considered here.
33. Marasmius abundans var. aurantiacus
Corner, Beih. Nova Hedwigia 111: 23. 1996.
(Fig. 39, Plate 3K)
Type: Singapore, Botanic Garden, Corner
s.n., 15 Dec. 1941 (E).
Pileus 3-30 mm diam, paraboloid to
broadly obtusely conical or broadly convex;
margin smooth (non-striate); surface dry, dull,
hygrophanous, minutely velutinous; orange
(5A6, 6B7-8) to grayish orange (5B5-6, 6B4-6)
or golden yellow (5B7) with deep orange
around the disc when young, becoming deep
orange (5B8, 6A8) overall at maturity.
Lamellae adnate, close (14-19) with 3-5 series
80
of lamellulae, orangish white, non-marginate.
Stipe 11-67 × 0.5-1 mm, central, cylindrical,
terete, equal, dry, dull, glabrous, non-insititious,
yellowish white at the apex, gradually
changing to orange, grayish orange, brownish
orange, light brown or reddish brown on the
base, with brownish orange basal mycelium.
Basidiospores (11-) 13-17 (-20) × (3-) 45 (-6) μm [xmr = 14.3-16.6 × 4.1-4.7 μm, xmm =
15.4 ± 0.9 × 4.4 ± 0.3 μm, Q = 2.9-4.5, Qmr =
3.3-4.1, Qmm = 3.5 ± 0.3, n = 25 spores per 4
collections], elongate-lacrymoid to clavate,
smooth, hyaline, inamyloid. Basidia 20-26 × 510 μm, 2-spored and 4-spored. Basidioles 2328 (-32) × 6-8 μm, fusoid to clavate.
Cheilocystidia numerous, lamellar edge sterile,
composed of Siccus-type broom cells; main
body (8-) 13-20 (-23) × 5-12 μm, cylindrical to
clavate or broadly clavate, hyaline, inamyloid,
thin-walled; apical setulae 2-5 (-7) × 0.5-1 μm,
crowded, cylindrical to conical, subacute to
obtuse, light yellow, inamyloid, thick-walled.
Pleurocystidia absent. Pileipellis weakly
mottled, composed of a hymeniform layer of
Siccus-type broom cells; main body (5-) 7-17 (22) × (4-) 5-13 μm, cylindrical to clavate or
subglobose, hyaline, inamyloid, thick-walled;
apical setulae 3-7 (-8) × 0.5-1 μm, cylindrical
Fungal Diversity
Fig. 39. Marasmius abundans var. aurantiacus (TYS 515 = KLU–M #1). a. Basidiomes. b. Basidiospores. c. Basidioles.
d. Cheilocystidia. e. Pileipellis. Bars: a = 20 mm, b-e = 10 µm.
to conical, light yellow to tawny, weakly
dextrinoid, thick-walled. Pileus trama interwoven; hyphae 4-7 μm diam, cylindrical,
hyaline, strongly dextrinoid, thin-walled, nongelatinous. Lamellar trama regular; hyphae 3-6
μm diam, cylindrical, hyaline, dextrinoid to
strongly
dextrinoid,
thin-walled,
nongelatinous. Stipe tissue monomitic; cortical
hyphae 4-6 μm diam, parallel, cylindrical,
smooth, yellow (apex) to brownish yellow
(base), weakly dextrinoid to dextrinoid, thickwalled; medullary hyphae parallel, cylindrical,
smooth, hyaline, dextrinoid, thin-walled.
Caulocystidia absent. Clamp connections
present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves. Malaysia (Johore, Selangor), Singapore.
Material examined: Malaysia, Johore, Endau
Rompin National Park, NERC, Peta Village, on way to
jetty, 14 July 2005, TYS 515 (KLU-M #1, SFSU);
Selangor, Sungai Chongkak Forest Reserve, N
03°12.705’, E 101°50.472’, elev. 122 m, 16 May 2005,
Yee-Shin Tan, TYS 455 (KLU-M#25, SFSU); same
location, 16 May 2005, Yee-Shin Tan, TYS 456 (KLUM#26, SFSU); Selangor, Ulu Gombak, University of
Malaya’s Field Study Center, 28 July 2003, Yee-Shin
Tan, TYS 035 (KLU-M#23, SFSU).
Notes: The specimen matches quite
closely to Corner’s protologue (1996), differing
only in basidiospore size. The Malaysian
specimens have basidiospores with means in
the 14.3-16.6 × 4.1-4.7 μm, whereas the
protologue documents smaller basidiospores in
the range 11-13 × 3.5-4 μm. It is quite close to
M. fulviceps Berk., described from Sri Lanka,
but that species reputedly grows on wood and
has spores 10-12.5 × 4-5 μm (Pegler, 1986).
A bipolar (unifactorial) mating system
was reported for M. abundans var. aurantiacus
by Tan et al. (2007) based on Malaysian
material.
81
Fig. 40. Marasmius abundans var. campanulatus (Holotype: E#205853). a. Basidiospores. b. Basidioles. c.
Cheilocystidia. d. Pileipellis. Bars: a-d = 7 µm.
34. Marasmius abundans var. campanulatus
Corner, Beih. Nova Hedwigia 111: 23. 1996.
(Fig. 40)
This variety of M. abundans was
described by Corner from material collected in
Singapore and has not yet been found in
Malaysia although the species is expected to
occur here.
Type study of Marasmius abundans var.
campanulatus:
Type: Singapore, Botanic Gardens, 23
March 1943, Corner s.n. (E #205853);
annotated as the type by Corner on the
specimen deposited at E, and he noted that a
painting of this specimen was made.
The holotype specimen consists of
approx. 20 basidiomes, loose or attached to
dicot leaves and twigs, most in good condition
but a few infected by a mold. As dried: Pileus
82
3-7 mm diam, convex, sometimes with a small
rounded umbo, margin even, not striate,
glabrous, ferruginous to reddish brown.
Lamellae adnexed, close to crowded (17-20)
with 3-4 series of lamellulae, narrow, dark
brown. Stipe 5-20 × 1 mm, central, cylindrical,
glabrous,
non-insititious,
arising
from
orangish-tawny basal mycelium, brown; no
rhizomorphs.
Basidiospores 9.6-11.8 × 3.5-4.2 µm [x =
10.7 × 3.9 µm], ellipsoid to subfusoid, smooth,
hyaline, inamyloid, thin-walled. Basidia not
observed.
Basidioles
subfusoid,
often
mucronate. Cheilocystidia abundant, edge
sterile, composed of Siccus-type broom cells;
main body 11-15 × 4.5-6.5 µm, clavate to
broadly clavate, hyaline, thin-walled; apical
setulae 2.5-6.5 (-10) × 1.3-2 µm, conical, acute,
hyaline, thin-walled. Pleurocystidia absent.
Pileipellis not mottled, composed of a
hymeniform layer of Siccus-type broom cells;
Fungal Diversity
main body 10-18 × 3.5-8 µm, cylindrical to
clavate, hyaline, thin-walled, inamyloid; apical
setulae 2.5-8 × 1-1.5 µm, conical, acute,
hyaline to pale yellowish brown, weakly
dextrinoid, thin-walled to slightly thick-walled.
Stipe tissue monomitic; cortical hyphae 3-6.5 (8) µm diam, cylindrical, parallel, pale
yellowish brown to brown, firm-walled,
dextrinoid; medullary hyphae similar but
hyaline. Caulocystidia absent. Clamp connections present.
Notes: There is confusion concerning
which specimen represents the type of this
variety. Corner annotated this specimen from
the Singapore Botanic Gardens (23 March
1943) as type and he noted on the packet that a
painting was made; but in the protologue he
reported a specimen from Bukit Timah (23 July
1943) as the type, and did not note that a
painting was made of this latter specimen.
However, the minimal description and the
painting (Plate 6b) seem to match better the
Bukit Timah specimen than the Botanic Garden
specimen that was annotated as the type.
Unfortunately, these two specimens are not
conspecific.
Specimen Bukit Timah, 23 July 1943.
This specimen matches the protologue,
including forming a striate pileus and broad,
distant lamellae. The three basidiomes in the
colour painting (Plate 6b) match the
basidiomes in the specimen packet, although
there is no note on the packet that a painting
was made of this specimen. This specimen is
not conspecific with the specimen from the
Singapore Botanic Gardens that Corner
annotated as type! In addition, this specimen
does not appear to be conspecific with the type
of M. abundans var. abundans. This specimen
is very moldy and no pertinent microscopic
features were retrieved.
35. Marasmius persicinus Desjardin, Retn. &
E. Horak, Sydowia 52: 164. 2000.
(Fig. 41, Plate 3D)
Type: Indonesia, Java, West Java, Mt.
Halimun National Park, loop trail from
Cikaniki, 14 Jan 1998, D.E. Desjardin 6793
(BO!).
Pileus 3-10 mm diam, paraboloid to
obtusely conical, expanding in age to broadly
convex with an umbo; margin sulcate to plicate;
surface dry, dull, minutely velutinous; orange
(5A6-7, 6B7-8) with light orange (5A5) disc,
seldom brownish orange (6C7-8). Lamellae
adnate, subdistant to close (8-16) with no
lamellulae, orangish white (11A2) with grayish
orange edge. Stipe 10-30 × 0.2-0.5 mm, central,
cylindrical, terete, equal, dry, dull, glabrous,
non-insititious, orangish white at the apex
grading to orange and dark brown at the base,
with brownish orange basal mycelium.
Basidiospores (14-) 16-21 × 3.5-5 μm
[xmr = 17.8-18.3 × 3.6-4 μm, xmm = 18.1 ± 0.4 ×
3.8 ± 0.3 μm, Q = 3.1-6, Qmr = 4.6-5.1, Qmm =
4.9 ± 0.3, n = 38 spores per 2 specimens],
elongate-lacrymoid to clavate, smooth, hyaline,
inamyloid. Basidia 20-25 × 6-8 μm, clavate, 4spored. Basidioles 20-23 × 7-9 μm, fusoid to
clavate. Cheilocystidia numerous, composed of
Siccus-type broom cells; main body 10-18 (-20)
× 6-10 (-11) μm, clavate to broadly clavate,
rarely lobed, hyaline to pale yellow, inamyloid,
thick-walled; apical setulae 2-5 × 0.5-1 μm,
crowded, subcylindrical to conical, subacute to
obtuse, rarely forked, yellow, inamyloid, thinto thick-walled. Pleurocystidia absent.
Pileipellis weakly mottled, composed of a
hymeniform layer of Siccus-type broom cells;
main body 8-15 (-17) × 6-12 μm, cylindrical to
clavate or irregular in outline, hyaline,
inamyloid, thin- to thick-walled; apical setulae
3-6 × 0.5-1 μm, cylindrical to conical, pale
yellow, weakly dextrinoid, thin-walled. Pileus
trama interwoven; hyphae 7-11 μm diam,
cylindrical, hyaline to pale yellow, dextrinoid,
thin-walled, non-gelatinous. Lamellar trama
regular; hyphae 3.5-7 μm diam, cylindrical,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Stipe tissue monomitic; cortical hyphae 5-7 μm
diam, parallel, cylindrical, smooth, yellowish
brown, weakly dextrinoid, thick-walled, nongelatinous;
medullary
hyphae
parallel,
cylindrical, 4-7 μm, smooth, hyaline, strongly
dextrinoid,
thin-walled,
non-gelatinous.
Caulocystidia absent. Clamp connections
present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves and twigs. Indonesia (Java), Malaysia
(Selangor).
83
Fig.41. Marasmius persicinus (TYS 474 = KLU-M#124). a. Basidiomes. b. Basidiospores. c. Basidia and basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
Material examined: Malaysia, Selangor, Pangsun,
Mount. Nuang Forest Reserve, N 03° 12.593’, E 101 °
52.575’, 319 m, 17 May 2005, Yee-Shin Tan, TYS 474
(KLU-M#124, SFSU); same location, 17 May 2005,
Hong-Twu Chan, TYS 481 (KLU-M#125, SFSU).
Notes: Marasmius persicinus, described
from Java, is most closely allied with M.
corneri (syn. M. incarnatus) described from
Singapore and recently reported from Thailand
(Wannathes et al., 2007). The two species
differ primarily in pileus size, number of
lamellae, and basidiospore size. Marasmius
persicinus has pilei 3-10 mm diam, 8-16
lamellae, and basidiospores with mean range
17.7-18.3 × 3.6-4 µm. Marasmius corneri has
pilei 10-43 mm diam, 12-18 lamellae, and
basidiospores with mean range 19-21.8 × 4-4.5
µm.
36. Marasmius corneri Wannathes, Desjardin
& S. Lumyong, Mycol. Res. 111: 992. 2007.
(Synonym: Marasmius incarnatus Corner,
Beih. Nova Hedwigia 111: 60. 1996 [non
Marasmius incarnatus Quél., Enchir. Fung.:
142. 1886].
(Fig. 42)
No specimens of M. corneri have yet
84
been collected in Malaysia although the species
is suspected to occur here. Corner described M.
incarnatus from Singapore, but since this
epithet is a homonym (non Marasmius
incarnatus Quél.), Wannathes et al. (2007)
proposed the new name M. corneri for the
species and reported a bipolar (unifactorial)
mating system based on specimens from
Thailand.
Type study of Marasmius incarnatus:
The holotype specimen (Singapore,
Gardens Jungle, 8 Aug. 1940, Corner s.n., E
#206722) consists of approx. 16 basidiomes in
good condition, loose or attached to leaves of
misc. dicots. As dried: Pileus 4-15 mm diam,
obtusely conical, with a small umbo, sulcate,
minutely subvelutinous, orangish brown (not
pale). Lamellae adnexed, non-collariate, distant,
usually with no lamellulae, broad, pale
orangish brown, edges concolorous. Stipe 1825 × 0.5-1 mm, central, terete, cylindrical, noninsititious, arising from orangish cream or
brownish orange mycelium; pale brown overall;
rhizomorphs absent.
Fungal Diversity
Fig.42. Marasmius incarnatus (Holotype: E#206722). a. Basidiospores. b. Cheilocystidia. c. Pileipellis. Bar = 10 µm.
Basidiospores 19-24 × 3.5-4.5 µm,
elongate-clavate, sometimes curved, smooth,
hyaline, inamyloid, thin-walled, typically with
a thick refractive region at the apiculus.
Basidia not observed. Basidioles 22-27 × 5.5-7
µm, subfusoid. Cheilocystidia abundant, edge
mostly sterile, composed of Siccus-type broom
cells; main body 10-17 × 4.5-8.5 µm, clavate to
subcylindrical, hyaline, inamyloid, thin-walled;
apical setulae 2-4 (-6.5) × 0.5-1.8 µm, conical
to irregular in outline, often wavy or with a few
nodules, obtuse to subacute, pale yellow,
inamyloid, thick-walled. Pleurocystidia absent.
Pileipellis mottled, composed of a hymeniform
layer of Siccus-type broom cells; main body 816 × 5.5-9.5 µm, subcylindrical to clavate or
irregular in outline, sometimes lobed, hyaline,
inamyloid, thin-walled or apically thick-walled;
apical setulae 3-8 × 0.5-1.8 µm, narrowly
conical, subacute, yellow to golden, weakly
dextrinoid, thick-walled; some cells with
longer, more thickly walled and darker
pigmented setulae. Pileus trama weakly
dextrinoid; lamellar trama dextrinoid. Stipe
tissue monomitic; cortical hyphae 2-5 µm diam,
parallel, cylindrical, golden, strongly dextrinoid; medullary hyphae hyaline, dextrinoid.
Caulocystidia absent. Clamp connections
present.
Notes: Marasmius corneri is phenetically
very close to M. persicinus. See there for a
comparison.
37. Marasmius selangorensis Y.S. Tan &
Desjardin, Fungal Diversity 25: 196. 2007.
(Fig. 43, Plate 3J)
Type: Malaysia, Selangor, Hulu Langat,
Sungai Chongkak Forest Reserve, 15 May
2005, Y.S. Tan #453 (KLU-M #4!).
Pileus 10-23 mm diam, paraboloid to
obtusely conical, convex or broadly convex
and umbonate; disc wrinkled; margin sulcate to
plicate; surface dry, dull, minutely velutinous;
brown (6E8) to dark brown (6F8) with reddish
tones overall when young, gradually becoming
yellowish brown (5D5-8, 5E8), light brown
(6D5-6), grayish orange (5B4-5), brownish
orange (6C5), pinkish brown, or golden brown
(5D7) on the margin with a dark brown disc.
Lamellae adnate, distant (12-18) with no
lamellulae, narrow, orangish white with brown
edges. Stipe 19-43 × 0.5-1 mm, central, terete,
equal, tough, dry, dull, glabrous, non-insititious,
base with strigose, yellowish white mycelium;
when young apex yellowish white to pale
orange with a deep orange base, in age
becoming yellowish white at apex, gradually
grading to orange, brownish orange, brown or
dark brown at the base. Odor pleasant.
Basidiospores (17-) 18-24 (-25) × (3.5-)
4-5 μm [xmr = 19.2-21.3 × 4-4.3 μm, xmm = 20.9
± 1.0 × 4.1 ± 0.2 μm, Q = 4-6.3, Qmr = 4.9-5.3,
Qmm = 5.0 ± 0.1, n = 30-40 spores per 3
collections], fusoid to clavate, smooth, hyaline,
85
Fig. 43. Marasmius selangorensis (Holotype: TYS 453 = KLU-M #4). a. Basidiomes. b. Basidiospores. c.
Cheilocystidia. d. Pileipellis. Bars: a = 10 mm, b-d = 10 µm.
inamyloid, thin-walled. Basidia not observed.
Basidioles 22-30 × 6-8 μm, fusoid to clavate.
Cheilocystidia numerous, composed of Siccustype broom cells; main body (7-) 10-16 (-20) ×
6-10 (-13) μm, subcylindrical to clavate,
broadly clavate or irregular in outline, seldom
lobed, hyaline, inamyloid, thick-walled; apical
setulae 4-12 × 1-2 μm, crowded, cylindrical to
conical, subacute to obtuse, sometimes forked,
light yellow to yellow or light brown,
inamyloid, thick-walled. Pleurocystidia absent.
Pileipellis mottled, composed of a hymeniform
layer of Siccus-type broom cells; main body 720 × 5-8 μm, cylindrical to clavate or
subclavate, sometimes lobed, light yellow,
inamyloid, thick-walled; apical setulae 7-15 ×
1-4 μm, numerous, cylindrical to conical,
86
rarely forked, yellow to brownish yellow or
brown, inamyloid, thick-walled. Pileus trama
interwoven; hyphae 4-8 μm diam, hyaline,
dextrinoid to strongly dextrinoid, thin-walled,
non-gelatinous. Lamellar trama regular;
hyphae 4-8 μm diam, cylindrical, hyaline to
light yellow, dextrinoid to strongly dextrinoid,
thin-walled, non-gelatinous. Stipe tissue
monomitic; cortical hyphae 3-9 μm diam,
parallel, cylindrical, smooth, yellow at apex,
brownish yellow at base, weakly dextrinoid to
dextrinoid, thick-walled (0.5-1 μm); medullary
hyphae 4-10 μm diam, parallel, cylindrical,
smooth, light yellow, dextrinoid, thin-walled or
with walls up to 0.5 μm thick. Caulocystidia
absent. Clamp connections present.
Fungal Diversity
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves. Malaysia (Selangor, Selayang).
38. Marasmius kanchingensis Y.S. Tan &
Desjardin, sp. nov.
(Fig. 44)
MycoBank: MB 512631
Material examined: Malaysia, Selangor, Hulu
Langat, Sungai Chongkak Forest Reserve, 15 May 2005,
Yee-Shin Tan, TYS 453 (Holotype: KLU-M #4; Isotype:
SFSU); same location, 15 May 2005, Yee-Shin Tan,
TYS 454 (KLU-M #5, SFSU); same location, 16 May
2005, Yee-Shin Tan, TYS 462 (KLU-M #6, SFSU);
Selayang, Kanching Forest Reserve, 9 Jan 2005, YeeShin Tan, TYS 435 (KLU-M#99, SFSU).
Etymology: kanching- = Kanching, -ensis =
origin, referring to the Kanching Forest Reserve where
the holotype was collected.
Pileus 2-12 mm latus, conicus vel convexus,
umbonatus, sulcatus, velutinus, aurantio-brunneus vel
atrobrunneus. Lamellae adnatae, subdistantes (14-16),
haud intervenosae, aurantio-albae, brunneo-marginatae.
Stipes 11-35 × 0.5 mm, cylindricus, glabrus, haud
insititius, apicaliter pallide aurantiacus, basim
atrobunneus, ad basim tomento pallide brunneo
strigosoque affixus. Basidiosporae 17-22 × 3.5-5.5 μm,
fusiodeae vel clavateae, leves, hyalinae, inamyloideae.
Basidia 31-33 × 8-9 μm, clavatae, 4-spora.
Cheilocystidia typi Sicci, 10-16 × 4-8 μm, clavata vel
late clavata, straminea, crasse-tunicata; setulae ad
apicem 6-12 × 1-1.5 μm, angusti-cylindricae vel conicae,
subacutae, flavae, inamyloideae, crasse-tunicatae.
Pleurocystidia nulla. Pileipellis hymeniformis, typi Sicci,
cellulae 10-16 × 6-11 μm, cylindricae vel late clavatae,
stramineae, inamyloideae, crasse-tunicatae; setulae ad
apicem 5-12 × 0.5-1.5 μm, cylindricae vel conicae,
subacutae, flavae vel brunneo-flavae, inamyloideae,
crasse-tunicatae. Caulocysitida nulla. Fibulae presentes.
Gregarius ad folia putrida plantarum dicotyledonearum.
Holotypus: Malaysia, Selangor, Selayang, Kanching
Forest Reserve, 8 Jan 2005, Yee-Shin Tan, TYS 415
(Holotypus: KLU-M#74).
Notes: Marasmius selangorensis is
characterized by the following features: a
brown to light brown or pinkish brown, sulcate
pileus with wrinkled disc; distant (12-18),
orange-white, narrow lamellae with brown
edges; a brownish orange to reddish brown,
glabrous stipe 20-40 mm long, arising from
yellowish white basal mycelium; clavate
basidiospores in the range 18-24 × 4-5 μm
(mean 20.9 × 4.1 μm); Siccus-type
cheilocystidia and pileipellis broom cells with
setulae up to 12 μm long; an absence of
pleurocystidia and caulocystidia; and growth
on
dicotyledonous
leaves.
Marasmius
selangorensis is allied with M. adhaesus and M.
aciebrunneus from Malaysia, and with M.
sierraleonis Beeli and M. carcharus Singer
from Africa. Marasmius adhaesus differs in
forming a pileus with olivaceous tones
(fuliginous olive, brownish olive) and has
longer basidiospores in the range 25-30 × 3-5
μm
(Corner,
1996;
E!).
Marasmius
aciebrunneus differs primarily in forming pilei
with ferruginous or orange and yellow colours,
and has slightly longer basidiospores (22-26 ×
4-5 μm; Corner, 1996; E!). Marasmius
sierraleonis differs in forming dull yellow to
rusty brown pilei, has a longer stipe (35-70
mm), smaller basidiospores (15.5-20 × 3.5-4.7
μm with mean 17.5 × 4 μm) and growth on
twigs (Pegler, 1977; Antonín, 2007).
Marasmius carcharus differs in forming much
broader lamellae (6-7 mm) that are nonmarginate, has a longer stipe (30-70 mm), and
smaller basidiospores (16-23.5 × 2.3-3.7 μm
with mean 17 × 3 μm; Singer, 1965; Pegler,
1977).
A bipolar (unifactorial) mating system
was reported for M. selangorensis by Tan et al.
(2007).
Pileus 2-12 mm diam, paraboloid to
obtusely conical or convex, umbonate; margin
sulcate; surface dull, dry, velutinous; grayish
orange (6B6) to brownish orange (5C6-7) when
young, dark brown (7F8) to brown (6E8) with
pale grayish orange (6B4) stripes margin in age.
Lamellae narrowly adnate, subdistant (14-16)
with no lamellulae, not intervenose, orangish
white (6A2) with brown edge. Stipe 11-35 ×
0.5 mm, central, terete, cylindrical, equal,
tough, dry, dull, glabrous, non-insititious,
orangish white (5A2) at apex, dark brown
(7F7-8) at base, with strigose, light brown
basal mycelium. Odor not distinctive.
Basidiospores (16-) 17-22 × 3.5-5.5 μm
[xmr = 18.5-20.5 × 4 μm, xmm = 19.5 ± 1.4 × 4 ±
0.0 μm, Q = 3.8-5.3, Qmr = 4.7-5.2, Qmm = 5 ±
0.3, n = 25 spores per 2 specimens], narrowly
elongated to fusoid or clavate, smooth, hyaline,
inamyloid, thin-walled. Basidia 31-33 × 8-9
μm, clavate, 4-spored. Basidioles 28-29 × 6-7
μm, fusoid to clavate. Cheilocystidia numerous,
composed of Siccus-type broom cells; main
body (8-) 10-16 (-22) × 4-8 (-10) μm, clavate
87
Fig.44. Marasmius kanchingensis (Holotype: TYS 415 = KLU-M#74). a. Basidiomes. b. Basidiospores. c. Basidia and
basidiole. d. Cheilocystidia. e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
to broadly clavate or irregular in outline, rarely
lobed, light yellow, inamyloid, thick-walled;
apical setulae (4-) 6-12 (-15) × 1-1.5 μm,
narrowly cylindrical to conical, subacute,
rarely forked, yellow, inamyloid, thick-walled.
Pleurocystidia absent. Pileipellis mottled,
composed of a hymeniform layer of Siccustype broom cells; main body (8-) 10-16 (-17) ×
(5-) 6-11 (-13) μm, cylindrical to clavate,
broadly clavate or irregular in outline, lobed,
light yellow, inamyloid, thick-walled; apical
setulae (3-) 5-12 (-23) × 0.5-1.5 (-2) μm,
narrowly cylindrical to conical, subacute,
rarely forked, yellow to brownish yellow,
inamyloid,
thick-walled.
Pileus
trama
interwoven; hyphae 2.5-5 µm diam, cylindrical,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Lamellar trama regular; hyphae 5-11 μm diam,
cylindrical, hyaline, strongly dextrinoid, thinwalled, non-gelatinous. Stipe tissue monomitic;
88
cortical hyphae 6-9 μm diam, parallel,
cylindrical, smooth, brownish yellow (apex) to
brown (base), weakly dextrinoid, thick-walled,
non-gelatinous; medullary hyphae 3-10 μm
diam, parallel, cylindrical, smooth, hyaline,
dextrinoid,
thick-walled,
non-gelatinous.
Caulocystidia absent. Clamp connections
present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves. Malaysia (Selangor).
Material examined: Malaysia, Selangor, Selayang,
Kanching Forest Reserve, N 03° 17.958’, E 101° 37.151’,
elev. 110m, 8 Jan 2005, Yee-Shin Tan, TYS 415
(Holotype: KLU-M#74; Isotype: SFSU); same location,
9 Jan, 2005, Dennis Desjardin, TYS 428 (KLU-M#75,
SFSU).
Notes: Distinctive features of M.
kanchingensis include: a small (< 12 mm diam),
sulcate and striped pileus coloured brown to
brownish orange with pale greyish orange
Fungal Diversity
sulcae; subdistant (10-16) lamellae with brown
edges; a glabrous stipe lacking caulocystidia;
basidiospores 17-22 × 3.5-5.5 with mean 19.5
× 4 µm; narrowly conical, acute setulae 5-12
µm long (or longer) on Siccus-type broom cells;
and growth on dicotyledonous leaves. The new
species is similar to M. striaepileus Antonin
from Burundi, M. phaeus Berk. & M. A. Curtis
from Cuba, M. bambusiniformis and M.
selangorensis. Marasmius striaepileus differs
in forming larger (15-30 mm diam) pilei,
slightly smaller basidiospores (15.5-18.5 × 3.55 µm) and is lignicolous (Antonín, 2007).
Marasmius phaeus differs in forming broom
cells with shorter setulae (5-8.5 µm), is
lignicolous and New World in origin (Singer,
1976). Marasmius bambusiniformis differs in
forming a paler, non-striped pileus, slightly
smaller basidiospores (15-18 × 3-4.5 µm), and
broom cells with much smaller setulae (2-5 µm
long). Marasmius selangorensis differs in
forming a pileus with more pink to reddish
brown tones and larger basidiospores (18-24 ×
4-5 µm with mean 21 × 4 µm). In the
phylogenetic tree, M. kanchingensis and M.
selangorensis are sister taxa with 1.0 PP
support, but M. kanchingensis is on a long
branch.
39.Marasmius aratus Massee, Bull. Misc.
Inform. 10: 358. 1914. nom. illegit. non
Marasmius aratus W.G. Sm., Lond. J. Bot. 11:
66. 1873.
(Fig. 45 & 46, Plate 3L)
Type: Singapore, E. M. Burkill #113 (K).
Pileus 4-13 (-18) mm diam, convex to
obtusely conical or broadly convex with a
small umbo; margin sulcate to plicate; surface
dry, dull, hygrophanous, minutely velutinous;
blood red to deep brownish purple, drying deep
reddish brown with a hint of burgundy.
Lamellae adnate to subfree, distant (10-16)
with no lamellulae, broad, yellowish white,
non-marginate. Stipe 6-17 × 0.2-0.5 mm,
central, cylindrical, terete, equal, dry, dull,
glabrous, non-insititious, yellowish white to
pink at the apex, dark brown on the base,
arising from white mycelium and accompanied
by yellowish white, wiry rhizomorphs.
Basidiospores 22-32 × 3.5-5 μm [xmr =
26.4-29.8 × 3.7-4.6 μm, xmm = 27 ± 0.8 × 4.1 ±
0.6 μm, Q = 5.4-8.3, Qmr = 5.8-6.8, Qmm = 6.4 ±
0.8, n = 25-27 spores per 4 specimens],
elongate-lacrymoid to clavate, sometimes
curved, smooth, hyaline, inamyloid. Basidia
not observed. Basidioles 32-40 × 7 μm, fusoid
to clavate. Cheilocystidia numerous, composed
of Siccus-type broom cells; main body (10-)
14-20 (-22) × 5-10 μm, subcylindrical to
clavate or broadly clavate, rarely lobed, hyaline,
inamyloid, thin-walled; apical setulae (4-) 6-10
× 0.5-1.5 μm, crowded, cylindrical to conical,
acute to subacute, rarely forked, pale yellow to
yellow, dextrinoid, thin- to thick-walled.
Pleurocystidia absent. Pileipellis strongly
mottled, composed of a hymeniform layer of
Siccus-type broom cells; main body (8-) 12-30
× 5-11 μm, cylindrical to clavate or irregular in
outline, hyaline to pale yellow, inamyloid, thinto thick-walled; apical setulae (2.5-) 4-8 × 0.51.5 μm, crowded, cylindrical to conical,
hyaline to tawny brown, weakly dextrinoid,
thick-walled. Pileus trama interwoven; hyphae
5-6 μm diam, cylindrical, hyaline, inamyloid to
dextrinoid,
thin-walled,
non-gelatinous.
Lamellar trama regular; hyphae 5-8 μm diam,
cylindrical, hyaline, dextrinoid, thin-walled,
non-gelatinous. Stipe tissue monomitic; cortical
hyphae 3-7 μm diam, parallel, cylindrical,
smooth, yellow (apex) to dark brown (base),
dextrinoid, thick-walled; medullary hyphae
parallel, cylindrical, 5-9 μm, smooth, pale
yellow at apex, yellow at base, weakly
dextrinoid
to
dextrinoid,
thin-walled.
Caulocystidia absent. Clamp connections
present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves and twigs. Malaysia (Selangor),
Singapore.
Material
examined:
Malaysia,
Selangor,
University of Malaya’s Field Study Centre, 20 Apr 2003
Yee-Shin Tan, KUM 152 (KLU-M#38, SFSU); Selangor,
Selayang, Kanching Forest Reserve, N 03°17. 958’, E
101°37. 151’, 8 Jan 2005, Yee-Shin Tan, TYS 409
(KLU-M#40). Singapore, Reservoir Jungle, 5 Aug. 1940,
Corner s.n. (E #205864); Singapore, Botanic Garden, 9
Aug. 1940, Corner s.n. (E #205859).
Notes: Our concept of M. aratus follows
that of Corner (1996) based on topotypical
material that he collected in Singapore.
Marasmius aratus is distinguished by a small,
blood red, plicate pileus, distant (10-16)
lamellae, a glabrous stipe that arises from white
89
Fig. 45. Marasmius aratus (KUM 152 = KLU–M#38). a. Basidiomes. b. Basidiospores. c. Basidioles. d. Cheilocystidia.
e. Pileipellis. Bars: a = 10 mm, b-e = 10 µm.
Fig. 46. Marasmius aratus (E#205859). a. Basidiospores. b. Cheilocystidia. c. Pileipellis. Bar = 10 µm.
mycelium and is associated with yellowish
white, wiry rhizomorphs, and very long
basidiospores in the range 22-32 × 3.5-5 µm
with mean 27 × 4.1 µm. In the field, M. aratus
may be mistaken for the typical red form of M.
haematocephalus commonly enclountered in
Southeast Asia, but the latter differs in forming
90
smaller basidiospores with mean range 16.8-22
× 4-4.3 µm and has prominent pleurocystidia.
40. Marasmius aciebrunneus Corner, Beih.
Nova Hedwigia 111: 24. 1996. (Fig. 47 & 48)
Type: Singapore, Reservoir Jungle, 5
August 1940, Corner s.n. (E #205862!).
Fungal Diversity
Pileus 7-20 mm diam, obtusely conical to
convex or broadly convex; disc rugolose;
margin striate to sulcate; surface dry, dull,
velutinous; brown (6-7E6-8) to ferruginous
with darker disc. Lamellae adnate, subdistant
(11-14) with no lamellulae, grayish orange
(5B3) with brown edge. Stipe 15-22 × 0.5-1
mm, central, cylindrical, terete, equal, dry, dull,
glabrous, non-insititious, yellowish white at the
apex, gradually change to brownish orange or
dark brown on the base, with pale orange
strigose basal mycelium; associated with white,
wiry rhizomorphs.
Basidiospores (19-) 20-27 (-30) × (3.2-)
4-5 μm [xmr = 23.4-26.9 × 4.1-4.8 μm, xmm =
24.8 ± 1.9 × 4.3 ± 0.6 μm , Q = 4.7-6.6, Qmr =
5.7-6, Qmm = 5.8 ± 0.2, n = 23 spores per 4
specimens], elongate-lacrimoid to clavate,
smooth, hyaline, inamyloid. Basidia 23-27 × 56 μm, clavate, 4-spored. Basidioles 28-32 × 6
μm, fusoid to clavate. Cheilocystidia numerous,
composed of Siccus-type broom cells; main
body (6-) 7-13 (-21) × 5-8 (-10) μm, clavate to
broadly clavate, subcylindrical or irregular in
outline, rarely lobed, hyaline to light yellow,
inamyloid, thick-walled; apical setulae (2-) 320 (-12) × 0.5-1 μm, crowded, cylindrical to
conical, acute to subacute, rarely forked, light
brownish yellow to yellow, inamyloid, thin- to
thick-walled. Pleurocystidia absent. Pileipellis
mottled, composed of a hymeniform layer of
Siccus-type broom cells; main body (6-) 7-13 (19) × 5-9 (-11) μm, cylindrical to clavate or
broadly clavate, hyaline to pale yellow,
inamyloid, thick-walled; apical setulae (3-) 4-6
× (0.3-) 0.5-1 μm, crowded, cylindrical to
conical, light yellow to brownish yellow, acute
to subacute inamyloid, thin- to thick-walled.
Pileus trama interwoven; hyphae 5-9 μm diam,
cylindrical, light yellow, strongly dextrinoid,
thin-walled, non-gelatinous. Lamellar trama
regular; hyphae 4-6 μm diam, cylindrical,
hyaline, weakly dextrinoid to dextrinoid, thinwalled, non-gelatinous. Stipe tissue monomitic;
cortical hyphae 6-8 μm diam, parallel,
cylindrical, smooth, dull yellow (apex) to
brownish yellow (base), weakly dextrinoid,
thick-walled; medullary hyphae parallel,
cylindrical, 3-8 μm, smooth, hyaline to light
yellow, dextrinoid, thin-walled, slightly
gelatinous. Caulocystidia absent. Clamp
connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves and twigs. Malaysia (Selangor),
Singapore.
Material examined: Malaysia, Selangor, Hulu
Langat, 16 May 2005, Yee-Shin Tan, TYS 457 (KLUM#30, SFSU); Selangor, University of Malaya’s Field
Study Centre, 28 July 2003, Yee-Shin Tan, TYS 027
(KLU-M#31, SFSU); same location, 28 July 2003, YeeShin Tan, TYS 036 (KLU-M#32, SFSU). Singapore,
Reservoir Jungle, 5 August 1940, Corner s.n. (Holotype:
E #205862).
Notes: Diagnostic features of M.
aciebrunneus include: a moderately large,
obtusely conical to convex, brown to
ferruginous, sulcate pileus; subdistant (11-14)
lamellae with brown-edges; a glabrous stipes
associated with white, wiry rhizomorphs; long
basidiospores in the range 20-27 × 4-5 μm with
mean 24.8 × 4.3 μm; and an absence of
pleurocystidia.
Morphologically,
M.
aciebrunneus is similar to M. adhaesus, but the
latter differs subtly in forming pilei with olive
tones (lacking ferruginous tones), has more
lamellae (10-20), and slightly longer
basidiospores in the range 25-30 × 3-5 µm with
mean 27.9 × 4.4 µm. Interestingly, in the ITS
derived phylogeny, the two species are placed
in distant clades, with M. aciebrunneus sister
of M. aratus (94% BS, 1.0 PP) and M.
adhaesus sister of M. bambusiniformis (71%
BS, 0.98 PP).
Type Study of Marasmius aciebrunneus:
The holotype specimen (Singapore,
Reservoir Jungle, 5 August 1940, Corner s.n.,
E #205862) consists of approx. 10 basidiomes
in good condition, loose or attached to dicot
leaves. As dried: Pileus 5-13 mm diam,
campanulate to plano-convex, disc strongly
rugulose, plicate, minutely subvelutinous, dark
ferruginous. Lamellae adnexed, non-collariate,
distant with 0-1 series of lamellulae, broad,
cream, edges nearly concolorous. Stipe 15-25 ×
1 mm, central, terete, cylindrical, glabrous,
non-insititious, pale brown overall, arising
from white to pale yellowish white basal
mycelium.
91
Fig. 47. Marasmius aciebrunneus (TYS 457 = KLU-M#30). a. Basidiomes. b. Basidiospores. c. Basidia and basidioles.
d. Cheilocystidia. e. Pileipellis. Bars: a = 20 mm, b-e = 10 µm.
Fig. 48. Marasmius aciebrunneus (Holotype: E#205862). a. Basidiospores. b. Basidioles. c. Cheilocystidia. d.
Pileipellis. Bar = 10 µm.
92
Fungal Diversity
Basidiospores 20-26 × 4-5 µm, elongateclavate to subfusoid, sometimes curved,
smooth, hyaline, inamyloid, thin-walled.
Basidia 4-spored, clavate, few seen. Basidioles
28-36 × 5.5-9.5 µm, subfusoid. Cheilocystidia
abundant, edge mostly sterile, of Siccus-type
broom cells; main body 12-20 × 5-8 mm,
clavate to subcylindrical, hyaline, inamyloid,
thin-walled; apical setulae 2.5-5 (-6) × 1-1.5
µm, narrowly conical, subacute, pale golden,
inamyloid to weakly dextrinoid, thin- to thickwalled. Pleurocystidia absent (no refractive
cells seen). Pileipellis not mottled or weakly
mottled, a hymeniform layer of Siccus-type
broom cells; main body 9-14 × 6-8 µm, clavate
to pyriform or irregular in outline, hyaline,
inamyloid, thin-walled or apically thick-walled;
apical setulae 2.5-6.5 (-9) × 1-1.5 µm, narrowly
conical, subacute, yellowish brown, weakly
dextrinoid, thick-walled. Pileus trama and
lamellar trama inamyloid to very weakly
dextrinoid. Stipe tissue monomitic; cortical
hyphae 2-5 µm diam, parallel, cylindrical, pale
reddish brown, strongly dextrinoid; medullary
hyphae similar but broader and hyaline.
Caulocystidia absent. Clamp connections
present.
41. Marasmius adhaesus Corner, Beih. Nova
Hedwigia 111: 27. 1996.
(Fig. 49 & 50, Plate 3I)
Type: Singapore, Garden's Jungle, 8 Aug.
1940, Corner s.n. (E #205880!).
Pileus 6-40 mm diam, paraboloid to
truncately conical, convex or broadly convex,
umbonate; margin sulcate to plicate; surface
dry, dull, hygrophanous, minutely velutinous;
grayish orange (5B4) to light brown (6D5-8),
brownish olive (4E4-6) or brown (6E6-7) with
a darker disc. Context thin, yellowish white.
Lamellae narrowly adnate, subdistant (10-20)
with no lamellulae, orangish white with light
brown edges. Stipe 12-55 × 0.5-1 mm, central,
cylindrical, terete, equal, dry, dull, glabrous,
non-insititious, pale orange (5A3) at the apex,
gradually grading to yellow, brownish orange
or dark brown at the base, with brownish
orange basal mycelium.
Basidiospores (21-) 25-30 (-31) × 3-5 μm
[xmr = 27.6-28.2 × 4-4.9 μm, xmm = 27.9 ± 0.8 ×
4.4 ± 0.2 μm, Q = 5.6-8.5, Qmr = 5.9-7.1, Qmm =
6.5 ± 0.6, n = 20 spores per 3 specimens],
elongate-lacrimoid to clavate, smooth, hyaline,
inamyloid. Basidia 24-26 × 6-8 μm, 4-spored.
Basidioles 25-30 (-34) × 6-9 μm, fusoid to
clavate. Cheilocystidia numerous, composed of
Siccus-type broom cells; main body (8-) 11-20
(-22) × (5-) 7-11 μm, cylindrical to clavate or
broadly clavate, hyaline to light yellow,
inamyloid, thick-walled; apical setulae (3-) 4-6
(-8) × 0.5-1 (-2) μm, crowded, cylindrical to
conical, subacute to obtuse, hyaline, yellow to
light brownish yellow, inamyloid, thick-walled.
Pleurocystidia absent. Pileipellis mottled,
composed of a hymeniform layer of Siccustype broom cells with scattered seta-like cells;
main body (8-) 10-20 (-26) × 5-7 (-10) μm,
subcylindrical to clavate or subclavate, hyaline,
inamyloid, thick-walled; apical setulae (3-) 6-9
(-12) × 0.5-1 μm, cylindrical to conical, light
yellow to light brown or brownish yellow,
inamyloid, thick-walled. Seta-like cells rare,
similar to Siccus-type broom cells but with
only 2-4 very long setulae; main body (10-) 1530 (-42) × 6-7 μm, cylindrical to subcylindrical,
light yellow, inamyloid, thick-walled; apical
setulae (15-) 20-33 (-35) × 1.5-3 (-4) μm,
brownish yellow, inamyloid, thick-walled.
Pileus trama interwoven; hyphae 4-10 μm
diam, cylindrical, hyaline, strongly dextrinoid,
thin-walled, non-gelatinous. Lamellar trama
regular; hyphae 4-7 μm diam, cylindrical,
hyaline, dextrinoid to strongly dextrinoid, thinwalled, non-gelatinous. Stipe tissue monomitic;
cortical hyphae 4-7 μm diam, parallel,
cylindrical, smooth, light yellow (apex) to
yellowish brown (base), inamyloid to weakly
dextrinoid, thick-walled (0.5 µm), nongelatinous; medullary hyphae parallel, 6-11 μm
diam, cylindrical, smooth, hyaline, dextrinoid
to strongly dextrinoid, thin-walled, weakly
gelatinous. Caulocystidia absent. Clamp
connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves. Malaysia (Selangor), Singapore.
Material examined: Malaysia, Selangor, Sungai
Chongkak Forest Reserve, N 03°12.705’, E 101°50.472’,
elev. 122 m, 16 May 2005, Yee-Shin Tan, TYS 464
(KLU-M#33, SFSU); same location, 16 May 2005, YeeShin Tan, TYS 467 (KLU-M#34, SFSU). Singapore,
Garden's Jungle, 8 Aug. 1940, Corner s.n. (Holotype: E
#205880).
93
Fig. 49. Marasmius adhaesus (TYS 464 = KLU–M#36). a. Basidiomes. b. Basidiospores. c. Basidia and basidioles. d.
Cheilocystidia. e. Pileipellis. Bars: a = 20 mm, b-e = 10 µm.
Notes:
Marasmius
adhaesus
is
characterized by a truncately conical to convex,
light brown to brownish olive, sulcate-plicate
pileus; subdistant (10-20) lamellae with brown
edges; a glabrous stipe lacking caulocystidia;
basidiospores in the range of 25-30 × 3-5 μm
with mean 27.9 × 4.4 μm; an absence of
pleurocystidia; a few seta-like cells in the
pileipellis with 2-4 setulae 15-30 µm long; and
growth on the dicotyledonous leaves.
94
Marasmius adhaesus is similar to the sympatric
species M. selangorensis but the latter differs
in forming pilei with reddish brown to golden
brown tones (not olive), smaller basidiospors in
the range 18-24 × 3.5-5 µm with mean 20.9 ×
4.1 µm, and pileipellis broom cells with
numerous setulae 7-15 µm long (seta-like cells
with setulae 15-30 µm long are absent). For a
comparison of M. adhaesus to M. aciebrunneus,
see the notes on the latter species.
Fungal Diversity
Fig. 50. Marasmius adhaesus (Holotype: E#205880). a. Basidiospores. b. Cheilocystidia. c. Pileipellis. Bar = 10 µm.
Type study of Marasmius adhaesus:
Corner originally labeled the packet M.
adhaerens, but apparently changed it to M.
adhaesus at time of publication. The holotype
specimen (Singapore, Garden's Jungle, 8 Aug.
1940, Corner s.n., E #205880) consists of
approx. 8 fragmented basidiomes in fair
condition, loose or attached to dicot leaves. As
dried: Pileus 10-22 mm diam, plano-convex,
plicate, glabrous, evenly brown. Lamellae
adnexed, not collariate, distant with no
lamellulae, broad, pale brown. Stipe 25-40 × 1
mm, central, cylindrical, non-insititious, arising
from cream-coloured, strigose basal mycelium,
glabrous, brown; no rhizomorphs.
Basidiospores 25-30.5 × 4.3-5.5 µm [x =
27.6 × 4.9 µm], elongate-clavate, smooth,
hyaline, inamyloid, thin-walled. Hymenial
cells poorly reviving. Basidia not seen.
Basidioles 40 × 8-10 µm, clavate to subfusoid.
Cheilocystidia abundant, edge mostly sterile, of
Siccus-type broom cells; main body 6.5-20 × 510 µm, subcylindrical to clavate, sometimes
lobed, hyaline, inamyloid, thin-walled; apical
setulae 4-14(-19) × 1-2 µm, irregularly
cylindrical to subconical, sometimes furcate,
acute, brown, thick-walled. Pleurocystidia
absent. Pileipellis not mottled, a hymeniform
layer of Siccus-type broom cells; main body 816 × 5-8 µm, subcylindrical to clavate, hyaline
and thin-walled at base, apically pale brown
and thick-walled, inamyloid; apical setulae 36.5(-8) × 0.5-1.5 µm, narrowly conical, rarely
furcate, acute, brown, inamyloid, thick-walled;
with a few scattered seta-like cells. Pileus
trama strongly dextrinoid. Lamellar trama
dextrinoid. Stipe tissue monomitic; cortical
hyphae 3-6.5 µm diam, parallel, cylindrical,
pale yellowish brown, strongly dextrinoid;
medullary hyphae 3-8 µm diam, hyaline,
dextrinoid, thin-walled. Caulocystidia absent.
Clamp connections present.
Marasmius sect. Sicci Singer, subsect. Siccini
Singer, ser. Haematocephali Singer, Fl.
Neotrop. Monogr. 17: 201. 1976.
Type species: Marasmius haematocephalus (Mont.) Fr.
42. Marasmius distantifolius Y.S. Tan &
Desjardin, sp. nov.
(Fig. 51, Plate 3H)
MycoBank: MB 512632
Etymology: distans- (Latin) = distant; -folius
(Latin) = leaf, referring to the distant lamellae.
Pileus 2-8 mm latus, conico-umbonatus vel late
convexus, papillatus, striatus vel sulcatus, subvelutinus,
brunneo-rufus, brunneo-violaceus vel griseo-rufus.
Lamellae adnatae, distantes (7-8), pallide incarnateae,
rubri-marginatae. Stipes 11-12 × 0.1-0.2 mm, cylindricus,
glabrus, haud insititius, apicaliter pallide rufus, basim
atrobrunneus, ad basim tomento albido affixus.
Basidiosporae 18.5-23 × 3.5-5 μm, fusiodeae, hyalinae,
inamyloideae. Basidiola 23-30 × 6-7 μm, clavatae vel
fusiodeae. Cheilocystidia typi Sicci, 10-25 × 5-10 μm,
clavata, hyalina; setulae ad apicem 3-6 × 0.5 μm, densae,
cylindricae vel conicae, subacutae vel obtusae,
95
Fig. 51. Marasmius distantifolius (Holotype: TYS 478 = KLU-M#72). a. Basidiomes. b. Basidiospores. c. Basidioles. d.
Cheilocystidia. e. Pleurocystidia. f. Pileipellis. Bars: a = 20 mm, b-f = 10 µm.
stramineae, inamyloideae, tenui-tunicatae. Pleurocystidia
rara, 37-42 × 9-12 μm, fusiodea vel clavata, mucronata,
flavida,
inamyloidea,
tenui-tunicata.
Pileipellis
hymeniformis, typi Sicci, cellulae 7-17 ×6-10 μm,
cylindricae vel clavatae, hyalinae, dextrinoideae, crassetunicatae; setulae ad apicem 3.5-6 × 0.5-1 μm, densae,
cylindricae vel conicae, acutae vel subacutae, fulvae,
dextrinoideae. Caulocystidia nulla. Fibulae presentes.
Gregarius, ad folia putrida plantarum dicotyledonearum.
Holotypus: Malaysia, Selangor, Pangsun, Mount Nuang
Forest Reserve, 17 May 2005, Yee-Shin Tan, TYS 478
(Holotypus: KLU-M#72).
Pileus 2-8 mm broad, obtusely conical
with umbonate disc when young, expanding in
age to convex or broadly convex-depressed
with an acute papilla; margin striate to sulcate;
surface dull, dry, subvelutinous; brownish red
(10D7-8) to brownish violet (11D7-8) or
greyish red (10C5), with ruby to pinkish
brown tones, not violet. Lamellae adnate,
distant (7-8) with no lamellulae, pinkish white
with darker pinkish red edges. Stipe 11-22 ×
0.1-0.2 mm, cylindrical, equal, tough, terete,
dry, dull, glabrous, non-insititious, pale red
96
(12A3) at the apex, dark brown at the base,
with white basal mycelium.
Basidiospores 18.5-23 (-25) × 3.5-5 (-6)
µm [xm = 21.4 ± 1.5 × 4 ± 0.6 µm, Q = 4-7.2,
Qm = 5.5 ± 0.7, n = 40], elongate-lacrymoid to
fusoid, hyaline, smooth, inamyloid. Basidia
not observed. Basidioles 23-30 × 6-7 µm,
clavate to fusoid. Cheilocystidia numerous,
composed of Siccus-type broom cells; main
body (5-) 10-25 × 5-10 (-11) µm, clavate to
broadly clavate, hyaline, inamyloid, thickwalled; apical setulae 3-6 × 0.5 µm, crowded,
cylindrical to conical, subacute to obtuse,
seldom branched or forked, light yellow,
inamyloid,
thin-walled.
Pleurocystidia
scattered, 37-42 × 9-12 µm, fusoid to clavate
with a constricted mucro, refractive, yellow,
inamyloid, thin-walled. Pileipellis mottled,
composed of a hymeniform layer of Siccustype broom cells; main body 7-17 (-20) × 6-10
(-13) µm, cylindrical to clavate or subclavate,
seldom lobed, hyaline, weakly dextrinoid,
Fungal Diversity
thick-walled; apical setulae 3.5-6 (-7) × 0.5-1
µm, crowded, cylindrical to conical, acute to
subacute, rarely forked, light brownish yellow
to tawny, weakly dextrinoid, thick-walled.
Pileus trama subparallel; hyphae 4-5 µm diam,
cylindrical, hyaline, dextrinoid, thin-walled,
non-gelatinous. Lamellar trama regular;
hyphae 3-5 µm diam, cylindrical, hyaline,
dextrinoid to strongly dextrinoid, thin-walled,
non-gelatinous. Stipe tissue monomitic;
cortical hyphae 4.5-6 µm diam, parallel,
cylindrical, smooth, dark brownish yellow,
very weakly dextrinoid, thick-walled (1-2 µm
thick), non-gelatinous; medullary hyphae 5-6
µm diam, parallel, cylindrical, smooth, hyaline
to light yellow, dextrinoid, thin-walled (0.5-1
µm), non-gelatinous. Caulocystidia absent.
Clamp connections present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves in montane forest. Malaysia (Selangor).
Material examined: Malaysia, Selangor, Pangsun,
Mount Nuang Forest Reserve, N 03° 12.593’, E 101 °
52.575’, 319 m, 17 May 2005, Yee-Shin Tan, TYS 478
(Holotype: KLU-M#72, Isotype: SFSU).
Notes: Marasmius distantifolius is
characterized by the following features: a
small (2-8 mm diam), sulcate, papillate,
convex-umbonate to broadly convex-depressed
pileus coloured deep brownish red to brownish
violet or greyish red (not violet); distant (7-9),
pinkish white lamellae with darker pinkish red
edges; a glabrous stipe 11-22 × 0.5 mm
coloured pale red at the apex and dark brown at
the base with white basal mycelium;
basidiospores in the range 18.5-25 × 3.5-5 µm
with mean 21.4 × 4 µm; small, broadly clavate
(not strangulate) pleurocystidia in the range 3742 × 9-12 µm; Siccus-type cheilocystidia and
pileipellis broom cells with narrowly conical,
acute setulae 3-6 µm long; and growth on
leaves. It is similar to Old World populations
of M. haematocephalus (Mont.) Fr. but the
latter differs in forming more violet pigmented
pilei, more lamellae (10-18), and slightly
shorter basidiospores with mean length in the
range 18-21 µm (see below; Desjardin &
Horak, 1997; Desjardin et al., 2000). The two
species are sister taxa in the ITS phylogeny
with strong support (100% BS, 1.0 PP),
although they are on long branches relative to
each other and to other species in the tree.
43. Marasmius haematocephalus (Mont.) Fr.,
Epicr. Syst. Mycol.: 382. 1838.
(Fig. 52, Plate 3E)
= Agaricus haematocephalus Mont., Ann. Sci.
Nat. Bot., sér. 2, 8: 369. 1837.
= Androsaceus haematocephalus (Mont.) Pat., J.
Bot. (Morot) 3: 336. 1889.
Taxonomic synonyms:
= Marasmius rhodocephalus Fr., Nova Acta
Regiae Soc. Sci. Uppsal. ser. 3, 1: 31. 1851.
= Androsaceus rhodocephalus (Fr.) Pat. &
Gaillard, Bull. Soc. Mycol. France 4: 20. 1888.
= Marasmius semipellucidus Berk. & Broome, J.
Linn. Soc., Bot. 14: 36. 1873.
= Marasmius sanguineus Cooke & Massee,
Grevillea 17: 59. 1889.
= Marasmius atropurpureus Murrill, N. Amer. Fl.
9: 262. 1915.
= Marasmius vinosus Beeli, Bull. Soc. Roy. Bot.
Belgique 60: 158. 1928.
Type: Brazil, Rio de Janeiro, A. de Saint
Hilaire. Not extant.
Pileus 2-7 mm in diam, obtusely conical
to convex with a small central papilla; margin
striate; surface dry, dull, minutely velutinous;
dark purple (14F7-8) to dark violet (15F7-8).
Lamellae adnate, subdistant (12-13) with no
lamellulae, pinkish white (11A2) to pale red
(11A3), non-marginate. Stipe 8-23 × 0.5-1 mm,
central, cylindrical, twisted-fibrous, dry, dull,
glabrous, non-insititious, dark brown overall,
arising from white basal mycelium.
Basidiospores (15-) 17-20 (-23) × 3.5-4.5
μm [x = 18.2 ± 1.6 × 3.8 ± 0.5 μm, Q = 4-5.3,
Qm = 4.8 ± 0.6, n = 40 spores], elongatelacrymoid to fusoid or clavate, smooth, hyaline,
inamyloid. Basidia not observed. Basidioles
17-28 × 8-12.5 μm, fusoid to clavate.
Cheilocystidia numerous, composed of Siccustype broom cells; main body 8-21 × (5-) 7-10
μm, subcylindrical to clavate or broadly clavate,
hyaline, inamyloid, thin- to thick-walled; apical
setulae 2-6 × 0.5-2 μm, crowded, cylindrical to
conical, subacute to obtuse, hyaline to light
yellow, thin- to thick-walled. Pleurocystidia
scattered, 30-42 × 7-10 µm, fusoid to clavate,
seldom with a constricted mucro, nonrefractive or refractive, hyaline, inamyloid,
thin-walled, arising from the lamellar trama
and projecting up to 13 µm beyond basidioles.
Pileipellis mottled, composed of a hymeniform
layer of Siccus-type broom cells; main body 920 × 5-13 μm, subcylindrical to clavate or
broadly clavate, hyaline to yellowish brown,
97
Fig. 52. Marasmius haematocephalus (TYS 523 = KLU-M#8). a. Basidiomes. b. Basidiospores. c. Basidioles. d.
Cheilocystidia. e. Pleurocystidia. f. Pileipellis. Bars: a = 10, b-f = 10 µm.
inamyloid, thin- to thick-walled; apical setulae
3-8 × 0.5-1 μm, cylindrical to conical, acute or
subacute, tawny to reddish brown, dextrinoid,
thick-walled. Pileus trama interwoven ; hyphae
3-4 μm diam, cylindrical, hyaline, dextrinoid,
thin-walled, non-gelatinous. Lamellar trama
regular; hyphae 3-5 μm diam, cylindrical,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Stipe tissue monomitic; cortical hyphae 3-5 μm
diam, parallel, cylindrical, smooth, dark brown,
dextrinoid, thick-walled; medullary hyphae 4-6
μm diam, parallel, cylindrical, smooth, hyaline,
dextrinoid to strongly dextrinoid, thin-walled.
Caulocystidia absent. Clamp connections
present.
Habit, habitat and known distribution:
gregarious on undetermined dicotyledonous
leaves. Putatively pantropical; Africa, Asia,
98
North, Central and South America, Malaysia
(Selangor), New Zealand, Papua New Guinea.
Material examined: Malaysia, Selangor, Ulu
Gombak, University of Malaya’s Field Study Center, 5
Sept. 2005, Yee-Shin Tan, TYS 523 (KLU-M #8, SFSU).
Notes: This is the purple form of M.
haematocephalus that is widespread in the Old
World tropics. The Malaysian material matches
that described from Sri Lanka (Pegler, 1986),
Papua New Guinea (Desjardin & Horak, 1997)
and Indonesia (Desjardin et al., 2000),
although the Malaysian population forms less
constricted pleurocystidia. Basidiospore size in
this species is apparently variable, with
populations in Sri Lanka having a spore mean
of 14 × 3.7 μm, in Malaysia and Papua New
Guinea the spore mean is 18.2 × 3.8 μm,
whereas in Indonesia (Java) they measure 20.5
× 4.0 μm.
Fungal Diversity
The mating system of New World
(Puerto Rico) specimens of M. haematocephalus was reported as bipolar (specimen TENN
50917), secondary homothallic (TENN 50916;
12 of 12 ssi were dikaryotic) and
amphithallic/bipolar (TENN 50918; 7 of 12 ssi
were dikaryotic) by Gordon et al. (1994).
Although no single-spore isolates (ssi) of the
Malaysian specimen germinated to form
dikaryotic hyphae (and hence the specimen is
not amphithallic), the report of a bipolar mating
system for a Malaysian population of M.
haematocephalus (Tan et al. 2007) is
consistent with that of New World populations.
Conclusions
A total of 43 species of Marasmius sensu
stricto are reported from Malaysia, including
two species (M. corneri and M. abundans var.
campanulatus) that have not yet been collected
in Malaysia but are expected to occur. Out of
the 41 species that have been collected, 19
species are documented as new records in
Malaysia, while 9 species are described herein
as new to science (viz., M. angustilamellatus,
M. diminutivus, M. distantifolius, M. iras, M.
kanchingensis, M. kuthubutheenii, M. musicolor, M. ochropoides, and M. olivascens).
Three additional species from Malaysia were
previously described by us as new to science,
viz., M. acerosus, M. nummularioides and M.
selangorensis (Tan et al., 2007). The 43
morphological species reported herein belong
to six sections of Marasmius as defined by
suites of morphological characters (sections
Sicci, Globulares, Marasmius, Hygrometrici,
Neosessiles and Leveilleani). Of the 121
species of Marasmius sensu lato reported from
Malesia by Corner (1996), type studies
revealed that only 20 of these species belong to
Marasmius sensu stricto. The other 101 species
belong to numerous other genera.
The ITS sequences supported species
concepts based on shared morphological
features where nearly all morphospecies were
clustered in well-resolved clades. The ITS
derived phylogeny was unable to resolve
currently accepted infrageneric taxa as
monophyletic groups, with the exception of
sect. Marasmius subsect. Marasmius which
was monophyletic with 96% BS and 1.0 PP
support. The inclusion of additional taxa and
additional genes are necessary to clarify the
infrageneric delimitation of Marasmius sensu
stricto. The ITS sequence data supported the
use of selected micromorphological features,
such as basidiospore size, cystidium-type, and
pileipellis anatomy, in delimiting species and
clusters of closely related species.
Acknowledgements
The authors would like to thank the University of
Malaya for the Vote F grants (F0159/2003D,
F0194/2005A and P0171/2006C), and the National
Science Foundation (USA) PEET grant (DEB-0118776)
to Desjardin, and IRPA grant (01-02-03-1002) from
Ministry of Science, Technology and Innovation
(MOSTI) Malaysia. We would also like to thank the
Johor State Government and University of Malaya for
the Vote F grant (F0738/2002A) for Johor National
Park-Endau Rompin Expedition, and Langkawi
Development Authority (LADA) and Malaysian Nature
Society (MNS) for the Langkawi Expedition.
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