Fungal Diversity
A monograph of Marasmius (Basidiomycota) from Northern Thailand based on
morphological and molecular (ITS sequences) data
Wannathes, N.1, Desjardin, D.E.2*, Hyde, K.D. 3,4, Perry, B.A.2 and Lumyong, S.5
1
Biology Program, Faculty of Science and Technology, Pibulsongkram Rajabhat University, 156 Moo 5 Plaichoompol,
Muang, Phitsanulok 65000, Thailand
2
Department of Biology, San Francisco State University, 1600 Holloway Ave., San Francisco, California 94132, USA
3
Kunming University of Science and Technology, Kunming, Yunnan, P.R. China
4
School of Science, Mae Fah Luang University, Tasud, Chiang Rai, Thailand
5
Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand
Wannathes, N., Desjardin, D.E., Hyde K.D., Perry, B.A. and Lumyong, S. (2009). A monograph of Marasmius
(Basidiomycota) from Northern Thailand based on morphological and molecular (ITS sequences) data. Fungal
Diversity 37: 209-306
Sixty-four taxa of Marasmius are reported representing fifty-seven species from northern Thailand. Comprehensive
descriptions, illustrations, photographs, a dichotomous key to aid in their identification, and comparisons with phenetically
similar taxa are provided. Seventeen species are described as new: Marasmius araneocephalus, M. auratus, M. bondoi, M.
brunneoolivascens, M. cafeyen, M. coarctatus, M. cupreostipes, M. delicatulus, M. ganyao, M. graminipes, M. imitarius,
M. inthanonensis, M. jasminodorus, M. makok, M. pseudopellucidus, M. suthepensis and M. tantulus. Molecular
phylogenetic reconstructions are presented based on ITS regions in Bayesian, likelihood and parsimony analyses.
Phylogenetic data were strongly correlated with morphological data and were useful to aid in delimiting species and
distinguishing among closely related species. ITS sequences were of limited use, however, in recognizing currently
circumscribed infrageneric taxa at the series rank and higher.
Key words: Agaricales, fungi, phylogeny, taxonomy
Article Information
Received 12 January 2008
Accepted 3 March 2009
Published online 1 August 2009
*Corresponding author: Dennis E Desjardin; e-mail: ded@sfsu.edu
Introduction
This paper is part of a series dealing with
the macrofungi of northern Thailand (Le et al.,
2007a,b; Wannathes et al., 2009). The genus
Marasmius Fr. (Agaricales, Marasmiaceae) is
worldwide in distribution and is composed of
approximately
600
species.
Recently
Marasmius of Southeast Asia has received
renewed interest through the research of
Desjardin et al. (2000) in Indonesia, Tan et al.
(2007) in Malaysia, and Desjardin et al. (2004)
and Wannathes et al. (2004, 2007) in Thailand.
To date, no monographic treatment of
Marasmius has been published from Thailand.
During the years 2003 to 2007, we conducted
extensive fieldwork in three provinces (Chiang
Mai, Chiang Rai and Phrae) of northern
Thailand. As a result of these expeditions, we
present here a monograph of Marasmius from
the region based on morphological and
molecular (ITS sequences) data.
In this treatment we accept the generic
circumscription of Marasmius as delimited by
Wilson and Desjardin (2005). They restricted
Marasmius to a monophyletic lineage based on
nLSU rDNA sequences data including only
sections Globulares, Hygrometrici, Leveilleani,
Marasmius, Neosessiles, Scotyphysini and Sicci
as defined by Singer (1976, 1986). Other
sections recognized by Singer (1976, 1986) are
currently excluded from Marasmius. Wilson
and Desjardin (2005) transferred members of
Marasmius section Alliacei to the genus
Mycetinis. Section Androsacei was elevated to
generic rank as Setulipes by Antonín (1987);
section Epiphylli belongs in the family
Physalacriaceae (Wilson and Desjardin 2005);
209
and section Fusicystides is treated as a
synonym of Setulipes (Desjardin unpubl.).
groups, and 3 species of Crinipellis for
outgroup rooting purposes.
Materials and methods
DNA extraction, Polymerase Chain Reaction
and sequencing:
Genomic DNA was extracted from two
sources: mostly from dried herbarium specimens,
with fifteen extractions from living cultures
(Table 1). Extraction of dried herbarium tissue
(1-10 mg) was done using the E.Z.N.A. fungal
DNA Miniprep Kit or E.Z.N.A. Forensic DNA
Extraction Kit (Omega Bio-tek Inc., Norcross,
GA, USA). Pure culture extractions were
performed with a modified SDS extraction
protocol of Roger and Bendich (1994) and
Johnson (1998).
ITS regions 1 and 2, and the 5.8S rDNA,
were amplified using primers ITS1-F, ITS5,
ITS 4, ITS4-B (Gardes and Bruns, 1993; White
et al., 1990) and ITS4-m (TTG AGC TTT TCC
CTC TTC AC) which was designed in this study.
When the amplification failed the intermediate
primers ITS2 and ITS3 (White et al., 1990)
were also used. The DNA fragment was
amplified on GeneAmp PCR System 9600
(Applied
Biosystems®,
Perkin-Elmer
Corporation, Nortwalk, CT, USA) or PCT100™ (MJ research, Inc., Watertown, MA, USA)
automated thermal cycle. PCR products were
purified using ExoSAPIT Kit (USB
Corporation, Cleveland, OH, USA) or GFX™
PCR DNA or Gel band Purification Kit
(Amersham Biosciences, USA) and run on an
ABI 3100 Genetic Analyzer System (Applied
Biosystems®, Foster City, CA, USA). Sequence
editing was performed with Sequencher 4.2
software (GeneCodes Co., Ann Arbor, MI,
USA). Edited sequences have been deposited
in GenBank (Table 1).
Morphological protocols
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
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. Specimens are deposited in the
mycological herbarium at Chiang Mai
University (CMU), and the H. D. Thiers
Herbarium at San Francisco State University
(SFSU).
Phylogenetic protocols
Taxon sampling:
One hundred and twenty five Internal
Transcribed Spacer (ITS) nrDNA sequences
representing 62 species were used in the
analyses, of which 116 sequences of
Marasmius were generated as part of this study
(Table 1), 4 sequences of Marasmius and 2
sequences of Crinipellis were obtained from
Y.S. Tan (Univ. of Malaya), 2 sequences of
Marasmius were obtained from GenBank, and 1
sequence of Crinipellis were obtained from J.
Kerekes (San Francisco State Univ.). The
sequences represent 59 species of Marasmius
in nine traditionally accepted infrageneric
210
Phylogenetic analyses:
Initial
sequence
alignment
was
performed with Clustal X (Thompson et al.,
1997) using the default settings, followed by
manual alignment with MacClade 4.04
(Maddison and Maddison, 2001). Parsimony
analyses were conducted using PAUP* 4.0b10
(Swofford, 2002). Searches employed a
heuristic search method with all characters
weighted equally, gaps treated as missing data,
random stepwise sequence addition, tree
bisection reconnection (TBR), and collapse of
Fungal Diversity
Table 1. List of Marasmius specimens sequenced (ITS1-5.8S-ITS2) for this study, itemized by
infrageneric group.
GenBank
accession No.
EU935544
NW422
Globulares
M. albimyceliosus
EU935481
NW331
Globulares
M. calvus
EU643514
NW152
Globulares
M. grandiviridis
EU643515
NW349**
Globulares
M. grandiviridis
EU643510
NW231
Globulares
M. laticlavatus
EU643512
NW293
Globulares
M. laticlavatus
EU643511
NW412
Globulares
M. laticlavatus
EU643516
DED7726
Globulares
M. mokfaensis
EU643517
NW020**
Globulares
M. mokfaensis
EU935508
NW321
Globulares
M. pellucidus
EU935509
NW342
Globulares
M. pellucidus
EU935510
NW352
Globulares
M. pellucidus
EU935539
NW158
Globulares
M. purpureostriatus
EU935538
NW318
Globulares
M. purpureostriatus
EU643513
NW286
Globulares
M. pseudopurpureostriatus
M. leveilleanus
Leveilleani
NW248
EU935566
M. leveilleanus
Leveilleani
NW268
EU935567
M. apatelius
Marasmius/ Marasmius
NW427
EU935561
M. apatelius
Marasmius/ Marasmius
NW437**
EU935562
M. somalomoensis
Marasmius/ Marasmius
NW232
EU935559
M. tantulus
Marasmius/ Marasmius
NW239
EU935560
NW128
EU935558
Marasmius/ Sicciformes
M. brevicollus
NW058**
EU935546
M. cafeyen
Marasmius/ Sicciformes
NW130***
EU935547
M. cafeyen
Marasmius/ Sicciformes
EU935548
Marasmius/ Sicciformes
NW200*
M. cafeyen
EU935555
Marasmius/ Sicciformes
NW348
M. crinis-equi
EU935564
Marasmius/ Sicciformes
M. aff. crinis-equi
NW182
EU935565
Marasmius/ Sicciformes
M. aff. crinis-equi
NW205
EU935552
Marasmius/ Sicciformes
M. guyanensis
NW254*
EU935553
Marasmius/ Sicciformes
NW280*
M. guyanensis
EU935554
Marasmius/ Sicciformes
TYS314*
M. guyanensis
EU935570
Marasmius/ Sicciformes
NW162
M. nigrobrunneus
EU935572
Marasmius/ Sicciformes
NW223
M. nigrobrunneus
EU935571
Marasmius/ Sicciformes
NW416*
M. nigrobrunneus
EU935576
Marasmius/ Sicciformes
NW025
M. nigrobrunneus “f. cinnamomeus”
EU935573
NW119
M. nigrobrunneus “f. cinnamomeus”
Marasmius/ Sicciformes
EU935578
NW120
M. nigrobrunneus “f. cinnamomeus”
Marasmius/ Sicciformes
EU935574
NW260*
M. nigrobrunneus “f. cinnamomeus”
Marasmius/ Sicciformes
EU935577
NW327*
M. nigrobrunneus “f. cinnamomeus”
Marasmius/ Sicciformes
EU935575
TYS281
M. nigrobrunneus “f. cinnamomeus”
Marasmius/ Sicciformes
EU935556
NW215*
M. purpureobrunneolus
Marasmius/ Sicciformes
EU935557
NW370
M. purpureobrunneolus
Marasmius/ Sicciformes
EU935563
NW155
M. purspureisetosu
Marasmius/ Sicciformes
EU935550
NW257*
M. ruforotula
Marasmius/ Sicciformes
EU935551
NW312
M. rufurotula
Marasmius/ Sicciformes
EU935549
NW256
M. straminiceps
Marasmius/ Sicciformes
EU935579
NW140
M. subruforotula
Marasmius/ Sicciformes
M. tenuissimus
Neosessiles
NW192*
EU935568
M. tenuissimus
Neosessiles
NW199
EU935569
M. araucariae var. siccipes
Sicci/ Atrorubentes
NW364
EU935511
M. auratus
Sicci/ Atrorubentes
NW076
EU935501
M. auratus
Sicci/ Atrorubentes
NW175
EU935502
*genomic DNA extracted from pure cultures, ** only ITS1-5.8S sequences were generated, *** only 5.8S-ITS2
sequences were generated
Species
Sections/ Series
Collection No.
211
Table 1 (Continued). List of Marasmius specimens sequenced (ITS1-5.8S-ITS2) for this study,
itemized by infrageneric group.
GenBank
accession No.
EU935515
NW414
Sicci/ Atrorubentes
M. inthanonensis
EU935486
NW276
Sicci/ Atrorubentes
M. iras
EU935487
NW375
Sicci/ Atrorubentes
M. iras
EU935512
NW067**
Sicci/ Atrorubentes
M. jasminodorus
EU935513
NW294
Sicci/ Atrorubentes
M. jasminodorus
EU935514
NW353
Sicci/ Atrorubentes
M. jasminodorus
EU935506
NW138
Sicci/ Atrorubentes
M. luteolus
EU935507
NW304
Sicci/ Atrorubentes
M. luteolus
NW299
Sicci/ Atrorubentes
M. ochroleucus
EU935503
NW186
Sicci/ Atrorubentes
M. pseudopellucidus
EU935504
NW305
Sicci/ Atrorubentes
M. pseudopellucidus
EU935505
JFK69
Sicci/ Atrorubentes
M. xestocephalus
EU935488
NW344
Sicci/ Atrorubentes
M. xestocephalus
EU935489
EU935521
NW329
Sicci/ Leonini
M. bambusiniformis
NW368
Sicci/ Leonini
M. bambusiniformis
EU935522
NW410
Sicci/ Leonini
M. bambusiniformis
EU935523
NW269
Sicci/ Leonini
M. corneri
EU935482
TYS274*
Sicci/ Leonini
M. corneri
EU935483
WN366
Sicci/ Leonini
M. cremeus
EU935494
TYS320
Sicci/ Leonini
M. cremeus
EU935495
NW150
Sicci/ Leonini
M. cupreostipes
EU935485
NW297
Sicci/ Leonini
M. imitarius
EU935496
NW423
Sicci/ Leonini
M. imitarius
EU935497
NW425
Sicci/ Leonini
M. imitarius
EU935498
NW201
Sicci/ Leonini
M. makok
EU935524
NW011
Sicci/ Haematocephali
M. bondoi
EU935472
NW237
Sicci/ Haematocephali
M. bondoi
EU935473
EU935474
NW320
Sicci/ Haematocephali
M. bondoi
EU935475
NW384
Sicci/ Haematocephali
M. bondoi
EU935476
NW386
Sicci/ Haematocephali
M. bondoi
EU935477
NW390
Sicci/ Haematocephali
M. bondoi
EU935478
NW399
Sicci/ Haematocephali
M. bondoi
EU935516
NW112
Sicci/ Haematocephali
M. brunneoolivascens
EU935519
NW277**
Sicci/ Haematocephali
M. brunneoolivascens
EU935517
NW373
Sicci/ Haematocephali
M. brunneoolivascens
EU935518
NW397***
Sicci/ Haematocephali
M. brunneoolivascens
NW005
Sicci/ Haematocephali
M. ganyao
EU935499
NW078
Sicci/ Haematocephali
M. graminipes
EU935479
M. haematocephalus “f. atrobrunneus”
Sicci/ Haematocephali
NW117
EU935525
M. haematocephalus “f. violaceus”
Sicci/ Haematocephali
NW193*
EU935531
M. haematocephalus “f. violaceus”
Sicci/ Haematocephali
NW339
EU935532
M. haematocephalus “f. violaceus”
Sicci/ Haematocephali
NW413
EU935533
M. haematocephalus f. haematocephalus
Sicci/ Haematocephali
NW296
EU935526
M. haematocephalus f. haematocephalus
Sicci/ Haematocephali
NW409
EU935527
M. haematocephalus f. haematocephalus
Sicci/ Haematocephali
NW428
EU935528
M. haematocephalus f. haematocephalus
Sicci/ Haematocephali
NW434
EU935529
M. haematocephalus f. haematocephalus
Sicci/ Haematocephali
TYS277
EU935530
M. haematocephalus “f. luteocephalus”
Sicci/ Haematocephali
NW310
EU935534
M. haematocephalus “f. robustus”
Sicci/ Haematocephali
NW330
EU935537
M. haematocephalus “f. robustus”
Sicci/ Haematocephali
NW433
EU935536
M. haematocephalus “f. variabilis”
Sicci/ Haematocephali
NW430
EU935535
M. hypophaeus
Sicci/ Haematocephali
NW285
EU935484
M. aff. pallescens
Sicci/ Haematocephali
NW424
EU935500
M. suthepensis
Sicci/ Haematocephali
TYS280*
EU935520
*genomic DNA extracted from pure cultures, ** only ITS1-5.8S sequences were generated, *** only 5.8S-ITS2
sequences were generated
Species
212
Sections/ Series
Collection No.
Fungal Diversity
Table 1 (Continued). List of Marasmius specimens sequenced (ITS1-5.8S-ITS2) for this study,
itemized by infrageneric group.
GenBank
accession No.
M. araneocephalus
Sicci/ Spinulosi
NW358
EU935540
M. coarctatus
Sicci/ Spinulosi
NW315
EU935541
M. coarctatus
Sicci/ Spinulosi
NW385
EU935542
M. coklatus
Sicci/ Spinulosi
TYS301
EU935543
M. nummularius
Sicci/ Spinulosi
NW266
EU935492
M. nummularius
Sicci/ Spinulosi
NW396
EU935493
M. trichotus
Sicci/ Spinulosi
NW262*
EU935490
M. trichotus
Sicci/ Spinulosi
NW263*
EU935491
*genomic DNA extracted from pure cultures, ** only ITS1-5.8S sequences were generated, *** only 5.8S-ITS2
sequences were generated
Species
Sections/ Series
zero length branches. Analyses were performed
in two steps: first, 1000 random sequence
addition replicates were performed, saving no
more than 5 trees per replicate, with MaxTrees
set to auto increase. Second, with MaxTrees
reset to 15 K, the most parsimonious trees
resulting from the first step were used as
starting trees and the analysis was allowed to
swap to completion.
The appropriate model of sequence
evolution for phylogenetic analyses was
determined using the program MrModeltest
v2.3 (Nylander, 2004). Maximum likelihood
(ML) analyses were conducted in PAUP*, and
employed the GTR+I+G model of sequence
evolution. ML searches followed an iterative
search strategy. First, a neighbor joining (NJ)
tree was constructed using a Jukes-Cantor (JC)
distance model. Next, with the search criterion
reset to ML, the model parameters were
estimated from the NJ tree, fixed, and an ML
search was conducted. After this initial search was
completed, the model parameters were reestimated from the resulting ML tree(s), fixed,
and another ML search was intiated with
starting trees obtained via NJ. This step was
completed a third and final time, for a total of
three independent ML searches. Upon
completion of the final search, trees resulting
from all three iterations were compared to
insure that the searches had converged on
equally likely topologies. Maximum likelihood
bootstrapping was performed in GARLI v0.951
(Zwickl, 2006) with 300 replicates under the
GTR+I+G model of sequence evolution, with
all parameter values estimated by the program
and using the default run termination settings.
Collection No.
Bayesian phylogenetic analyses were
carried out using Metropolis-coupled Markov
chain Monte Carlo (MCMCMC) methods with
MrBayes v3.1.2 (Huelsenbeck & Ronquist,
2001; Ronquist and Huelsenbeck, 2003), under
a GTR+I+G model as determined above. The
analyses consisted of two parallel searches, run
for 5,000,000 generations, with six chains and
random starting trees. Default settings in
MrBayes were used for the incremental heating
scheme of the chains (3 heated and 1 cold
chain),
unconstrained
branch
length
(unconstrained: exponential (10.0)), and
uninformative topology (uniform) priors. The
chains were sampled every 1000 generations.
Trees sampled prior to searches reaching a split
deviation frequency of 0.03 were discarded as
the burn-in, and the resulting trees were used to
calculate Bayesian posterior probabilities.
To further analyze the relationships
among three Marasmius species complexes,
sub-alignments were constructed by pruning
taxa from the larger ITS dataset to represent the
species complexes and sister taxa. Boostrap
analyses were performned in PAUP* on
resulting sub-alignments, consisting of 2000
replicates under a branch and bound search
algorithm with furthest sequence addition and
MulTrees on.
Results
Molecular Phylogeny
Phylogenetic analyses were performed on
an ITS dataset of 125 sequences of which 112
sequences were Marasmius species and 3
sequences were Crinipellis species, with the
213
latter used as an outgroup for rooting purposes.
Two new Thai Marasmius species were not
included in the analyses because of the
presence of multiple different ITS copies in
each species. Cloning was not conducted to
address this problem. Sequencing products
ranged from 270 nucleotides (Marasmius
mokfaensis NW020; partial sequence) to 883
nucleotides (Marasmius tantulus NW239). All
sequences were aligned and the ends trimmed
to create a dataset of 701 nucleotides that
included 354 parsimony informative characters.
Each of the ML analysis iterations
recovered a single tree, the likelihood values of
which did not differ significantly. We have
selected the topology resulting from the third
iteration to present here (Fig. 1; -lnL =
10983.475). Parsimony analyses produced 15
K equally parsimonious trees 819 steps, C.I. =
0.427, R.I. = 0.701),), which did not differ
significantly in topology from those recovered in
the ML analyses. Bayesian analyses reached a
standard deviation of split frequencies of 0.03
after 1,592,000 generations, and the intial 4000
trees recovered were excluded as the burn-in.
Maximum likelihood bootstrap values (BS) and
Bayesian posterior probalilites (PP) support
many of the terminal nodes in the phylogeny,
but fail to recover the deeper nodes with strong
support.
All members of sections Globulares plus
Sicci belong to a monophyletic clade with
relatively strong support (1.0 PP and 76% BS;
clade A), although the sections themselves are
not monophyletic. Eight species currently
placed in sect. Globulares fall into three
distinct subclades (AA, AD, AF), although five
of them belong to a single subclade (AD) but
with low support (.67 PP, 52% BS). Members
of section Marasmius, comprised of species
with lamellae attached to a collarium, do not
form a monophyletic group (clades B-F and
other species at the base of the tree). Section
Marasmius subsection Marasmius (collariate
species with Rotalis-type broom cells) is
monophyletic for the four included species
with 1.0 PP and 83% BS support (clade D).
214
Section Marasmius subsection Sicciformes
(collariate species with Siccus-type broom cells)
is not monophyletic, with species clustering
into more than three separate clades: clade B
with 0.91 PP and 60% BS; clade C with 0.92
PP and 85 % BS; clade E with no statistical
support; plus four unsupported taxa at the base
of the tree. A single member of section
Neosessiles (M. tenuissimus) and the monotypic
section Leveilleani (M. leveilleanus) form a
clade (F) with relatively strong support (0.97
PP and 82% BS) although its placement in the
tree is not supported statistically. Within sect.
Sicci, the four infrasectional taxa circumscribed
by Singer (1976) based on micromorphological
features
(viz.,
series
Atrorubentes,
Haematocephali, Leonini, Spinulosi) are not
monophyletic (Fig. 1). Members of ser.
Atrorubentes fall into two clades (AA, AC);
members of ser. Haematocephali belong to
three clades (AA, AB, AE) and two isolated
lineages; members of ser. Leonini are scattered
throughout Clade A; while members of ser.
Spinulosi fall into two clades (AC, AF) plus an
isolated lineage sister to AA (M. coarctatus).
Clade AF, with 1.0 PP support, contains
members of sect. Globulares, sect. Sicci series
Leonini and Spinulosi.
In the ML analysis, three species
complexes were uncovered, based on M.
bondoi (sect. Sicci ser. Haematocephali; clade
AB) with 1.0 PP and 74% BS support, M.
haematocephalus
(sect.
Sicci
ser.
Haematocephali; clade AE) with 1.0 PP and
99% BS support, and M. nigrobrunneus (sect.
Marasmius subsect. Sicciformes) in an
unresolved position near the base of the tree
(Fig. 1). To help clarify the taxonomic
boundaries of these species complexes, three
separate bootstrap branch and bound analyses
were performed based on datasets that included
the ingroup taxa with their sister taxa (as
resolved in the ML tree) used as outgroup.
These results will be discussed in the
commentaries that follow the species
descriptions.
Fungal Diversity
0.96/ 87 M. pellucidus NW321 -G
1.0/ 100
M. pellucidus NW342 -G
M. pellucidus NW352 -G
M. araucariae var. siccipes NW364 -SA
M. jasminodorus NW414 -SA
M. jasminodorus NW067 (type) -SA
0.98/
M. jasminodorus NW294 0.97/ 78
M. inthanonensis NW353 (type) -SA
1.0/ 77 1.0/ 100 M. auratus NW175 (type) -SA
M. auratus NW076 0.66/ 54
M. ochroleucus NW299 -SA
1.0/ 97
M. pseudopellucidus NW305 (type) -SA
1.0/ 87
M. pseudopellucidus NW186 -SA
0.98/
0.99/ 92
M. luteolus NW138 -SA
63
M. luteolus NW304 -SA
1.0/
0.95/ 74
M. brunneoolivasscens NW112 (type) -SH
M. brunneoolivascens NW373 -SH
M. brunneoolivascens NW277 -SH
M. brunneoolivascens NW397 -SH
M. suthepensis TYS280 (type) -SH
1.0/ 100 M. coarctatus NW315 -SS
M. coarctatus NW385 (type) -SS
0.93/ 85
M. bambusiniformis NW329 -SL
1.0/ 100
M. bambusiniformis NW368 -SL
M. bambusiniformis NW410 -SL
M. makok NW201 (type) -SL
0.64/ 60
0.52 / 53 M. bondoi NW390 -SH
M. graminipes NW078 (type) -SH
M. bondoi NW011 -SH
M. bondoi NW386 (type) -SH
1.0/ 74
M. bondoi NW237 -SH
0 78/ 84 M. bondoi NW399 -SH
0.84/ 59
M. bondoi NW384 -SH
1.0/ 77
M. bondoi NW320-SH
M. plicatulus NW439 -SL
M. calvus NW331 M. ganyao NW005 (type) -SH
1.0/ 100 M. corneri NW269 -SL
M. corneri TYS274 M. hypophaeus NW285 -SH
M. cupreostipes NW150 (type) -SL
1.0/ 99 M. imitarius NW423 (type) -SL
M. imitarius NW425 -SL
1.0/ 87 M. imitarius NW297 -SL
1.0/ 100 M. iras NW276 1.0/ 0.93 M. iras sNW375 -SA
M. nummularius NW266 0.77
M. nummularius NW396 M.
xestocephalus JFK69 -SA
1.0/ 100
1.0/ 97
M. xestocephalus NW344 -SA
M. trichotus NW263 -SS
M. trichotus NW262 -SS
M. aff. pallescens NW424 -SH
1.0/ 99 M. cremeus WN366 -SL
M. cremeus TYS320 -SL
1.0/ 100 M. grandiviridis NW152 (type) -G
0.81/ 56
M. grandiviridis NW349 -G
0.54
M. purpureostriatus NW158 -G
M. purpureostriatus NW318 -G
1.0/ 100
M. pseudopurpureostriatus NW286 (type) -G
M. laticlavatus NW231 (type) -G
0.67/ 52
M. laticlavatus NW412 -G
M. laticlavatus NW293 -G
1.0/ 100 1.0/ 98 M. mokfaensis DED7726 (type) -G
M. mokfaensis NW020 -G
0.9
0.76 M. haematocephalus f. haematocephalus NW296 -SH
M. haematocephalus f. haematocephalus NW428 -SH
1.0/ 100
M. haematocephalus f. haematocephalus NW409 -SH
1.0/ 93 M. haematocephalus f. haematocephalus TYS277 -SH
M. haematocephalus f. haematocephalus NW434 -SH
0.79/ 63
M. haematocephalus f. atrobrunneus NW117 -SH
0.86/ 65
M. haematocephalus f. violaceus NW193 -SH
1.0/
M. haematocephalus f. violaceus NW413 -SH
1.0/
1.0/ 76
M. haematocephalus f. violaceus NW339 -SH
M. haematocephalus f. variabilis NW430 -SH
0.52
M. haematocephalus f. luteocephalus NW310 -SH
M. haematocephalus f. robustus NW433 -SH
0.65/ 64
M. haematocephalus f. robustus NW330 -SH
M. siccus DED 5255 -SH
0.52/ 71
0.95/ 60
M. coklatus TYS301 -SS
0.63
M. hypochroides NW405 -SL
0.76
M. oreades DQ490641 -G
1.0
M. albimyceliosus NW422 -G
0.97/ 55
M. araneocephalus NW358 (type) -SS
M. cohaerans var. lachnophyllus DED 4071 -SS
0.77/ 78
AA
AB
A
AC
AD
AE
AF
10
Fig. 1. Maximum likelihood tree (-lnL = 10983.745) generated using a GTR+I+G model of nucleotide evolution.
Numbers to the left of / are Bayesian posterior probabilities, and those to the right are ML bootstrap percentages. G – sect.
Globulares; L – sect. Leveilleani; MM – sect. Marasmius subsect. Marasmius; MS – sect. Marasmius subsect.
Sicciformes; N – sect. Neosessiles; SA – sect. Sicci ser. Atrorubentes; SH – sect. Sicci ser. Haematocephali; SL – sect. Sicci ser.
Leonini; SS – sect. Sicci ser. Spinulosi.
215
1.0
M. cafeyen NW130 (type) -MS
1.0/ 100 M. cafeyen NW200 -MS
0.94/ 62
M. cafeyen NW058 -MS
1.0/ 88 M. straminiceps NW256 -MS
M. ruforotula NW257 -MS
M.rufurotula NW312 -MS
0.91/ 60
1.0/ 93 M. guyanensis NW254 -MS
0.99/ 94
M. guyanensis TYS314 -MS
M. guyanensis NW280 -MS
1.0/ 93
M. crinisequi NW348 -MS
1.0/ 100 M. purpureobrunneolus NW215 -MS
1.0
0.92/ 85
M. purpureobrunneolus NW370 -MS
M. brevicollus NW128 -MS
M. rotula DQ182506 -MM
0.91/ 64
0.52
M. apatelius NW427 -MM
1.0
M. apatelius NW437 -MM
1.0/ 83
M. tantulus NW239 (type) -MM
0.99
M. somalomoensis NW232 -MM
100 M. aff. crinisequi NW182 -MS
M. aff. crinisequi NW205 -MS
M. purpureosetosus NW155 -MS
0.97/ 82
1.0/ 94 M. tenuissimus NW199 -N
1.0/ 97 M. tenuissimus NW192 -N
0.63
M. tenuissimus DED 7659 -N
M. leveilleanus NW248 -L
1.0/ 100
M. leveilleanus NW268 -L
M. nigrobrunneus f. cinnamoneus NW119 -MS
M. nigrobrunneus f. cinnamoneus NW260 -MS
M. nigrobrunneus f. cinnamoneus TYS281 (type) -MS
M. nigrobrunneus f. cinnamoneus NW025 -MS
0.84/ 67
M. nigrobrunneus f. cinnamoneus NW327 -MS
M. nigrobrunneus NW162 -MS
M. nigrobrunneus NW416 -MS
M. nigrobrunneus NW223 -MS
M. nigrobrunneus f cinnamoneus NW120 -MS
M. curreyi DED 5142 -MS
M. subruforotula NW140 -MS
1.0/ 89
C. malesiana TYS 346
C. brunneipurpurea JFK 84
1.0
C. dipterocarpi TYS111
B
C
D
E
F
10 changes
Fig. 1 (Continued). Maximum likelihood tree (-lnL = 10983.745) generated using a GTR+I+G model of nucleotide
evolution. Numbers to the left of / are Bayesian posterior probabilities, and those to the right are ML bootstrap percentages. G
– sect. Globulares; L – sect. Leveilleani; MM – sect. Marasmius subsect. Marasmius; MS – sect. Marasmius subsect. Sicciformes;
N – sect. Neosessiles; SA – sect. Sicci ser. Atrorubentes; SH – sect. Sicci ser. Haematocephali; SL – sect. Sicci ser. Leonini; SS –
sect. Sicci ser. Spinulosi.
216
Fungal Diversity
Taxonomy
Key to species of Marasmius from northern
Thailand
1.
1.
2.
2.
3.
3.
4.
4.
5.
5.
6.
6.
7.
7.
Collarium present; stipe insititious..........................
................................................ (sect. Marasmius) 2
Collarium absent; stipe insititious or non-insititious
............................................................................. 21
Pileipellis composed of Rotalis-type broom cells
...........................................(subsect. Marasmius) 3
Pileipellis composed of Siccus-type broom cels.......
..........................................(subsect. Sicciformes) 6
Basidiospores (17-) 20-22 × 4 µm; basidiomes very
small, with pilei up to 1.5 mm diam and stipes up to
1 mm long.......................................... 1. M. tantulus
Basidiospores ≤ 15 µm long; basidiomes larger,
with pilei usually > 1.5 mm diam and stipes > 5 mm
long......................................................................... 4
Lamellae distant (6-10), non-marginate; pileus
yellowish white to brownish cream with a light
brown central spot .........................2. M. delicatulus
Lamellae subdistant to close (9-18), brownmarginate; pileus brown with a hint of olive or
brownish cream to yellowish brown....................... 5
Pileus 2-4 mm diam, brown with a hint of olive,
with a reddish brown papilla in the center surround
by a pale zone; stipe 8-47 mm long ..........................
...............................................3. M. somalomoensis
Pileus 2-10 mm diam, light brownish cream to
yellowish brown without olive tones, with or
without a small brown papilla, lacking a pale central
zone; stipe 8-15 mm long .................4. M. apatelius
Pileus minutely hispidulous from the presence of
pileosetae ................................................................ 7
Pileus glabrous, lacking pileosetae ......................... 8
Pileosetae uncommon, 36-82 µm long; pileipellis
broom cells and cheilocystidia with numerous (5-20)
apical setulae; lamellae distant (7-8); pileus 0.2-2
mm diam; stipe arising directly from rhizomorphs...
................................................ 5. M. berambutanus
Pileosetae common, 45-350 µm long; pileipellis
broom cells and cheilocystidia with few (2-6)
apical setulae; lamellae subdistant (13-17); pileus
1-4 mm diam; stipe not arising directly from
rhizomorphs..........................6. M. purpureisetosus
8.
8.
Basidiospores > 13 µm long.................................. 9
Basidiospores ≤ 12 µm long................................ 10
9.
Basidiospores 16-23 × 3.5-5.5 µm, xmm = 19.7 ×
4.6 µm; lamellae distant (6-9) ......7. M. brevicollus
Basidiospores 13-16 × 3-4 µm, xmm = 14.2 × 3.2
µm; lamellae distant to subdistant (9-13) ................
......................................8. M. purpureobrunneolus
9.
10. Stipe arising directly from rhizomorphs.............. 11
10. Stipe arising directly from the substrate;
rhizomorphs present or absent............................. 16
11. Basidiomes and rhizomorphs associated with wood;
lamellae remote to distant (5-10) ............................. 12
11. Basidiomes and rhizomorphs associated with
leaves, typically bamboo and other grasses or rarely
dicot leaves; lamellae subdistant (8-17) .............. 14
12. Pileus very pale, cream-colored . 9. M. pallenticeps
12. Pileus more deeply pigmented, reddish brown to
orangish brown, orange to greyish orange .......... 13
13. Pileus reddish brown to orangish brown; lamellae
distant (8-10); basidiospores 10-12 × 4-4.5 µm, xmm
= 10.6 × 4.6 µm, Qm = 2............. 10. M. crinis-equi
13. Pileus orange to greyish orange; lamellae remote
(5-8); basidiospores 8-10 × 4-6 µm, xmm = 9 × 4.9
µm, Qm = 1.9 ....................... 11. M. aff. crinis-equi
14. Pileus 2-6 mm diam, reddish brown; stipe 4-10 mm
long ....................................... 12. M. subruforotula
14. Pileus 2-28 mm diam, black, dark grey, greyish
brown, brown or brownish orange; stipe 8-215 mm
long ..................................................................... 15
15. Pileus black, dark grey or greyish brown; stipe 50215 mm long ...........................................................
.............13A. M. nigrobrunneus f. nigrobrunneus
15 Pileus brown to brownish orange; stipe 8-60 mm
long ... 13B. M. nigrobrunneus “f. cinnamomeus”
16. Pileus reddish brown; lamellae subdistant (11-13);
stipe 4-10 mm long ............... 12. M. subruforotula
16. Pileus brown, brownish orange, brownish yellow,
rusty brown, cream-brown, light yellow; lamellae
distant (8-12); stipe 8-35(-60) mm long .............. 17
17. Pileus light yellow to yellowish white; rhizomorphs
absent ...................................... 14. M. straminiceps
17. Pileus more deeply pigmented, brown, brownish
orange, brownish yellow, rusty brown or creambrown; rhizomorphs present or absent ................ 18
18. Stipe arising directly from rhizomorphs or from
bamboo or other grass leaves ..................................
.......... 13B. M. nigrobrunneus “f. cinnamomeus”
18. Stipe arising directly from dicot leaves............... 19
19. Basidiospores narrow, 10-12 × 3-4 µm, xmm = 11.1
× 3.5 µm, Qmm = 3.2.................. 15. M. guyanensis
19. Basidiospores broader, 9-12 × 4-7 µm, xmmr = 1010.2 × 4.3-5.8, Qmmr = 1.8-2.4............................. 20
20. Pileus disc brown with a cream-brown margin;
basidiospores 9-11 × 4-5 µm, xmm = 10 × 4.3 µm,
Qmm = 2. .......................................... 16. M. cafeyen
20. Pileus disc brownish yellow or rusty brown with a
greyish orange to light yellow margin;
basidiospores 10-12 × 4.5-7 µm, xmm = 10.2 × 5.8
µm, Qmm = 1 ................................17. M. ruforotula
217
21. Pileipellis composed of a hymeniform layer of
clavate to pyriform, non-setulose cells....................
............................................ (sect. Globulares)... 22
21. Pileipellis composed of Siccus-type broom cells 29
29. Stipe none or rudimentary, lateral to strongly
eccentric, less than 2 mm long ................................
................(sect. Neosessiles)... 26. M. tenuissimus
29. Stipe central, well-developed .............................. 30
22. Pleurocystidia present, fusoid............ 18. M. calvus
22. Pleurocystidia absent........................................... 23
30. Stipe insititious; lamellae free, well-separated from
the stipe apex; coarse black rhizomorphs may be
present ..... (sect. Leveilleani)... 27. M. leveilleanus
30. Stipe non-insititious; lamellae attached, typically
adnexed to adnate; rhizomorphs absent ..................
........................................................ (sect. Sicci) 31
23. Basidiospores < 10 µm long................................ 24
23. Basidiospores > 18 µm long................................ 25
24. Cheilocystidia indistinct, basidiomorphous; pileus
context thin but opaque, subhygrophanous;
lamellae relatively broad (2-3 mm), strongly
intervenose; pileus with brown to creamy brown
disc and striae; basidiomes usually solitary.............
..............................................19. M. albimyceliosus
24. Cheilocystidia distinct, irregularly cylindrical to
fusoid, ventricose or clavate; pileus context
extremely thin, pellucid, hygrophanous; lamellae
narrow (< 2 mm), often forked and intervenose;
pileus with cream to orange white disc, pure white
elsewhere; basidiomes usually in dense cespitose
clusters ........................................20. M. pellucidus
25. Pileus with bluish grey, purple or violaceous
pigments .............................................................. 26
25. Pileus lacking bluish grey, purple or violaceous
pigments .............................................................. 28
26. Basidiomes robust, with pilei 30-90 mm diam,
bluish grey to purplish grey overall when young
becoming grayish brown with paler ridges in age;
basidiospores 27-33 × 5-6 µm (xmm = 30 × 5.3 µm)
................................................... 21. M. mokfaensis
26. Basidiomes smaller, with pilei 13-38 mm diam,
striped, with dark violet to grayish violet disc and
plicae and yellow to grayish yellow ridges;
basidiospores 20-30 × 4-7 µm (xmr = 21.3-28.2 ×
4.6-6.2 µm).......................................................... 27
27. Pileus rather small, 13-20 mm diam, with dark
violet to violet disc and plicae, and narrow pale
grayish yellow to grayish white ridges; stipe up to
1.5 mm thick; basidiospores 21-30 × 4-7 µm (xmm =
23.8 × 5.3 µm).................. 22. M. purpureostriatus
27. Pileus larger, 14-38 mm diam, with violet to purple
disc and narrow plicae, and broad yellow to cream
ridges; stipe 1.5-3 mm thick; basidiospores 20-24.5
× 5-6 µm (x = 22.7 × 5.6 µm) .................................
...............................23. M. pseudopurpureostriatus
28. Pileus 37-88 mm diam, yellowish green with darker
olive green plicae; stipe 3-7 mm thick;
cheilocystidia clavate to irregular, sometimes lobed,
5-12 µm diam; basidiospores 26-30 × 4-5 µm (x =
26.7 × 4.4 µm).........................24. M. grandiviridis
28. Pileus 15-19 (-33) mm diam, yellowish grey to
greyish cream with a brown disc; stipe 1-2 mm
thick; cheilocystidia broadly clavate, 11-16 µm
diam; basidiospores 26-35 × 5-7 µm (xmm = 30.4 ×
5.6 µm) ......................................25. M. laticlavatus
218
31. Setae present on pileus, lamellae and/or stipe.........
.................................................. (ser. Spinulosi) 32
31. Setae absent......................................................... 37
32. Pleurocystidia and pleurosetae absent................. 33
32. Pleurocystidia or pleurosetae present.................. 35
33. Basidiospores 6-7 × 2.5-3.5 µm; lamellae very
crowded, > 22 with 4-5 series of lamellulae; pileus
brown with a hint of olive ...........28. M. coarctatus
33. Basidiospores 12-15 × 3-5 µm; lamellae close to
subdistant, 12-22 with 2-3 series of lamellulae;
pileus orange, brownish orange or reddish brown...
............................................................................ 34
34. Pileus orange to brownish orange; lamellae close
(16-22); pileosetae abundant, golden, up to 300 µm
or more long; caulosetae simple, acicular ...............
...................................................... 29. M. trichotus
34. Pileus reddish brown with brown margin; lamellae
subdistant (12-18); pileosetae absent or rare,
reddish brown, up to 115 µm long; caulosetae often
with apical setulae................. 30. M. nummularius
35. Basidiospores 10-13 × 5.5-7 µm .... 31. M. coklatus
35. Basidiospores 6-8 × 2.5-4 µm ............................. 36
36. Pileus smooth to striatulate, not venose-reticulate,
brown with a hint of olive; lamellae very crowded
(> 22) with 4-5 series of lamellulae; pileipellis
formed from Siccus-type broom cells, simple
pileosetae up to 60 µm long and transitional cells
with 2-4 apical setulae up to 37 µm long ................
....................................................28. M. coarctatus
36. Pileus venose-reticulate, dark purplish brown;
lamellae subdistant (10-12) with 3 series of
lamellulae; pileipellis formed from Siccus-type
broom plus arachnoid cells with 2-4 setulae 30-70
µm long ..............................32. M. araneocephalus
37. Stipe pruinose overall, pruinosity formed from
irregularly cylindrical, obtuse to subacute, nonsetulose cells, Siccus type cells typically absent on
stipe surface (if present then they are in
combination with non-setulose cells) ......................
............................................ (ser. Atrorubentes) 38
37. Stipe glabrous or pruinose, if pruinose then
pruinosity formed only from Siccus-type broom
cells ..................................................................... 47
Fungal Diversity
38. Cheilocystidia simple, cylindrical to clavate,
Siccus-type broom cells absent from lamellar edge
............................................................................. 39
38. Cheilocystidia of Siccus-type broom cells .......... 40
39. Pileus disc golden yellow, margin orange to
yellowish orange; stipe base dark reddish brown;
basidiospore xmm = 12.2 × 4.1 µm, Qmm = 3.0;
basidiomes solitary on dicot leaves . 33. M. auratus
39. Pileus disc pale yellowish white, margin white,
fading to white overall in age; stipe base pale
orange to grayish orange; basidiospore xmm = 11.0
× 4.1 µm, Qmm = 2.7; basidiomes cespitose on
bamboo debris ................. 34. M. pseudopellucidus
40. Pileus light yellow to cream overall; lamellae close
to crowded (20-24).................. 35. M. ochroleucus
40. Pileus more deeply pigmented, olive brown, brown,
reddish brown, brownish orange or yellowish
brown; lamellae subdistant to close (12-20)........ 41
41. Caulocystidia of two types: Siccus-type broom cells
plus simple cylindrical cells ................................ 42
41. Caulocystidia of one type: simple or lobed,
cylindrical cells, Siccus-type broom cells absent
from stipe surface ................................................ 44
42. Pleurocystidia present as scattered Siccus-type
broom cells; pileus bright brownish orange to
golden orange with paler creamy orange margin;
lamellae distant (8-13), narrow (1-2 mm diam) ......
.........................................................36. M. luteolus
42. Pleurocystidia absent; pileus dark brown to dark
reddish brown with light brown to brownish orange
margin; lamellae subdistant (12-20), broad (2-5
mm diam) ............................................................ 43
43. Odor strongly of jasmine tea; lamellae pale
yellowish white; pileus disc rugulose, dark reddish
brown .................................... 37. M. jasminodorus
43. Odor absent; lamellae brownish orange to grayish
brown or brown; pileus disc smooth, dark brown to
brown or brownish orange.......................................
............................38B. M. araucariae var. siccipes
44. Caulocystidia apically lobed or forked; pileus with
distinct yellow tones when young (disc yellowish
brown, margin grayish yellow to yellowish orange
...............................................39. M. xestocephalus
44. Caulocystidia all simple, cylindrical cells; pileus
lacking yellow tones when young, with dark brown,
reddish brown or olive tones ............................... 45
45. Basidiospores 12-14 × 3-4 µm with xmm = 12.8 ×
3.7 µm, Qmm = 3.5; pileus dark brown overall when
young..................................................... 40. M. iras
45. Basidiospores 8-12 × 3-4 µm with xmr = 10.4-10.5
× 3.9-4.0 µm, Qmr = 2.6-2.7; pileus olive or dark
reddish brown overall when young ..................... 46
46. Pileus disc smooth, dark olive to olive when young;
stipe long, up to 100 mm long.................................
............................................. 41. M. inthanonensis
46. Pileus disc rugulose, dark reddish brown when
young; stipe shorter, up to 45 mm long...................
..................... 38A. M. araucariae var. araucariae
47. Pleurocystidia absent....................(ser. Leonini) 48
47. Pleurocystidia present .(ser. Haematocephali)... 54
48. Pileus white to cream colored ........ 42. M. cremeus
48. Pileus more deeply pigmented, olive, brown,
reddish brown, brownish orange, greyish orange or
orange.................................................................. 49
49. Basidiospores 9-12 × 5-6 µm, Qm ≤ 2.0 ..................
...............................................43. M. hypochroides
49. Basidiospores in the range 12-26 × 3-5 µm, Qm ≥
4.0 ....................................................................... 50
50. Pileus conical, with a dark greyish brown disc and
olive margin ......................................44. M. makok
50. Pileus convex, brown, reddish brown, brownish
orange, greyish orange or orange, lacking olive
tones .................................................................... 51
51. Stipe distinctly copper colored overall, 70-250 mm
long ......................................... 45. M. cupreostipes
51. Stipe base brown, dark brown or reddish brown
(never copper colored), 13-90 mm long.............. 52
52. Pileus 10-43 mm diam; stipe 1-3 mm thick ............
.........................................................46. M. corneri
52. Pileus 2-15 mm diam; stipe < 0.5 mm thick........ 53
53. Pileus brown to reddish brown; lamellae distant
(10-12); basidiospore xmm = 18.6 × 4.4 µm;
basidiomes mostly on woody sticks ........................
......................................................47. M. imitarius
53. Pileus brownish orange to greyish orange; lamellae
subdistant (10-18); basidiospore xmm = 16.2 × 3.7
µm; basidiomes on dicot leaves ..............................
.........................................48. M. bambusiniformis
54. Basidiospores 8-12 × 4-5(-6) µm, xmm = 9.6 × 4.8
µm, Qmm = 2.1 .............................. 49. M. confertus
54. Basidiospores 11-32 × 3.5-5.5 µm, xmmr = 12.628.2 × 3.7-4.5 µm, Qmmr = 3.0-6.6....................... 55
55. Basidiospores 25-32 × 4-4.5 µm, xmm = 28.2 × 4.3
µm, Qmm = 6.6; stipe 100-195 × 0.5 mm; lamellae
distant (6-8); basidiomes on bamboo leaves ...........
.........................................................50. M. ganyao
55. Basidiospores 11-25 × 3.5-5.5 µm, xmmr = 12.6-22
× 3.7-4.5 µm, Qmmr = 3.0-5.9; stipe < 100 mm long;
lamellae distant to close (8-24); basidiomes
typically on dicot leaves or wood, rarely on
bamboo or other grass debris .............................. 56
219
56. Lamellae close (20-24); stipe surface with
caulocystidia formed from grass-like clusters of
setulae 6-40 x 1-2 µm lacking a basal cell or arising
from a rudimentary basal cell.... 51. M. graminipes
56. Lamellae distant to subdistant (8-18); stipe surface
glabrous, lacking caulocystidia ........................... 57
57. Basidiospores 11-15 × 3.5-5.5 µm, xmmr = 12.613.2 × 4-4.5 µm, Qmmr = 3.0-3.7.......................... 58
57. Basidiospores 13-25 × 3.5-5 µm, xmmr = 16.1-22 ×
3.7-4.2 µm, Qmmr = 4.2-5.9.................................. 59
58. Pileus dark brown overall or with brown margin in
age; lamellae olive brown to grayish olive or
yellowish olive; Siccus-type broom cells often
scattered on lamellar sides.......................................
.......................................52. M. brunneoolivascens
58. Pileus disc bright brown, margin brownish orange
to brownish yellow; lamellae yellowish white;
Siccus type broom cells absent on lamellar sides ....
...................................................53. M. suthepensis
59. Pileus deep reddish brown, maroon, dark violet,
purple, purplish brown or yellowish orange with
white margin; basidiospores 19-25 × 3.5-5 µm, xmm
= 20.7 × 4.0 mm, Qmm = 5.2 forms of .....................
.........................................54. M. haematocephalus
59. Pileus dark brown to brown, ferruginous, brownish
orange, brownish yellow or grayish brown (lacking
deep red, violet, purple or light yellow tones);
basidiospores 13-19 × 3.5-5 µm, xmmr = 16.1-18.1 ×
4.0-4.2 µm, Qmmr = 4.2-4.3.................................. 60
60. Basidiomes on monocot leaves; pileus typically
small, 1-5(-8) mm diam....................................... 61
60. Basidiomes on dicot leaves and wood; pileus
typically larger, 4-30(-48) mm diam ................... 62
61. Pileus dark brown overall; lamellae distant (8-11);
stipe 5-15 mm long..................................................
........54B. M. haematocephalus “f. atrobrunneus”
61. Pileus disc brown to ferruginous, margin
ferruginous to grayish orange; lamellae subdistant
(11-13); stipe 22-35 mm long..................................
..................................................55. M. hypophaeus
62. Pileus 4-15 mm diam, pallid, light brown to grayish
brown with paler margin; cheilocystidia setulae 3-6
µm long ................................ 56. M. aff. pallescens
62. Pileus more robust, 8-30(-48) mm diam, more
deeply pigmented, brown to brownish orange,
yellowish brown with brownish cream margin;
cheilocystidia setulae 3-15(-21) µm long ...............
.......................................................... 57. M. bondoi
Enumeration of taxa
Section Marasmius, subsect. Marasmius
Type: Marasmius rotula (Scop.: Fr.) Fr.
= sect. Pararotulae Singer, Sydowia 18: 339.
220
1965. [Type: Marasmius pararotula Singer].
= subsect. Pararotulae (Singer) Singer, Fl.
Neotrop. Monogr. 17: 92. 1976.
1. Marasmius tantulus Wannathes, Desjardin
& Lumyong, sp. nov.
(Fig. 2)
MycoBank: MB512412.
Etymology: ‘tantulus’ = very small; referring to
the very tiny basidiomes.
Pileus < 1.5 mm diametro, convexus, breviter
umbilicatus, sine papilla, striatus, glaber, hebetatus,
cinereo-creameus. Contextus creameus, tenuis. Lamellae
adnatae usque collarium infirmus evolutum, remotae (34), angustae, creameae, haud marginatae, haud
intervenosae. Stipes < 1 × < 0.3 mm, excentricus, sursum
angustatus cum basi subbulbosa, subvelutinus, insititius,
brunneus, sine rhizomorpha. Odor saporque non propria.
Basidiosporae (17-)20-22 × 4 µm, fusoideae usque
anguste clavatae, laeves, hyalinae, inamyloideae,
tenuitunicatae. Basidia non observata. Basidiolae
fusoideae usque clavatae. Cheilocystidia vulgaria, typi
Rotalis; 18-23 × 10-18 µm, subglobosa usque late
clavata, hyalina, inamyloidea, tenuitunicata; setulis
divergentibus 1-2 × 1 µm, gongylodibus usque conicis,
obtusis, confertis ad dimidium superiorem, hyalinis,
tenui- usque crassetunicatis. Pleurocystidia nulla.
Pileipellis hymeniformis, haud maculosus, typi Rotalis;
27-35 × 16-22 µm, late clavatus, subglobosus, pyriformis
vel adumbratim inaequabilis, hyalinus, inamyloideus,
tenuitunicatus; setulis divergentibus 1-2 × 1 µm,
gongylodibus usque conicis, obtusis, saepe confertis ad
dimidium superiorem, flavis, crassetunicatis. Trama pilei
intertexta, inamyloidea. Trama lamellae intertexta usque
regularis, hyphis 3-6 µm diametro, cylindratis, laevibus,
hyalinis, inamyloideis, tenuitunicatis, haud gelatinosis.
Stipitipellis parallelus, hyphis 5-10 µm diametro,
cylindratis cum eminentiis digitiformibus dispersis, laete
flavis
usque
brunneo-flavis,
laevibus,
leniter
dextrinoideis usque dextrinoideis, tenuitunicatis, haud
gelatinosis. Trama stipitis subparallela, hyphis 4-6 µm
diametro, cylindratis, hyalinis, laevibus, inamyloideis,
tenuitunicatis, haud gelatinosis. Caulocystidia nulla.
Fibulae praesentes in omnibus texturis.
Typus: Chiang Mai Province, Doi Inthanon
National Park Junction of Highway 1,009 and road to
Mae Chaem, N 18º 31.6’ E 98º 29.6’ alt. 1,703 m.,
dispersus usque gregarius in ligno, 28 July 2004, N.
Wannathes 239 (CMU: holotypus).
Pileus < 1.5 mm diam., convex,
shallowly umbilicate, without papilla, striate,
glabrous, dull, greyish cream. Context cream,
thin. Lamellae adnate to a poorly developed
collarium, remote (3-4), narrow, cream, nonmarginate, non-intervenose. Stipe < 1 × < 0.3
mm, eccentric, tapering upward with subbulbous
at base, subvelutinous, insititious, brown;
rhizomorphs absent. Odor and taste not
distinctive.
Fungal Diversity
brownish yellow, smooth, weakly dextrinoid to
dextrinoid, thin-walled, non-gelatinous. Stipe
trama hyphae 4-6 µm diam., subparallel,
cylindrical, hyaline, smooth, inamyloid, thinwalled, non-gelatinous. Caulocystidia absent.
Clamp connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on wood, Northern
Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Inthanon National Park Junction of
Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E
98º 29.6’ alt. 1,703 m., 28 July 2004, N. Wannathes 239
(CMU: holotype).
Fig. 2. Marasmius tantulus (N. Wannathes 239). 1.
Basidiomes 2. Basidiospores 3. Basidioles 4.
Cheilocystidia 5. Pileipellis 6. Stipitipellis, Scale bar 1 =
2 mm, 2-6 = 10 µm
Basidiospores (17-)20-22 × 4 µm [xm =
20.1 ± 1.3 × 4.0 ± 0 µm, Q = 4.3-5.5, Qm = 5.0,
n = 9 spores, s = 1 specimen], fusoid to
narrowly clavate, smooth, hyaline, inamyloid,
thin-walled. Basidia not observed. Basidioles
fusoid to clavate. Cheilocystidia common, of
Rotalis-type broom cells; main body 18-23 ×
10-18 µm, subglobose to broadly clavate,
hyaline, inamyloid, thin-walled; divergent
setulae 1-2 × 1 µm, knob-like to conical, obtuse,
dense over upper half of cell, hyaline, thin- to
thick-walled. Pleurocystidia absent. Pileipellis
hymeniform, not mottled, composed of Rotalistype broom cells; main body 27-35 × 16-22 µm,
broadly clavate, subglobose, pyriform, or
irregular in outline, hyaline, inamyloid, thinwalled; divergent setulae 1-2 × 1 µm, knob-like
to conical, obtuse, often dense over upper half
of cell, yellow, thick-walled. Pileus trama
interwoven, inamyloid. Lamellar trama hyphae
3-6 µm diam., interwoven to regular,
cylindrical, smooth, hyaline, inamyloid, thinwalled, non-gelatinous. Stipitipellis hyphae 510 µm diam., parallel, cylindrical with
scattered finger-like projections, light yellow to
Discussion: Marasmius tantulus is
characterized by very tiny basidiomes with
greyish cream pilei less than 1.5 mm diam,
collariate and remote lamellae (3-4), an
eccentric stipe less than 1 mm long, and very
long basidiospores with mean 20.1 × 4.0 µm.
Few species of sect. Marasmius subsect.
Marasmius have been described with
basidiospores longer than 11 µm, and none
were previously known with basidiospores 2022 µm long. The most similar species are M.
arimanus Dennis from Trinidad, and M.
rosulatus Desjardin & R.H. Petersen from New
Zealand. Marasmius arimanus differs in
forming mummy brown pilei 2-3 mm diam.
and basidiospores 11-14 × 3.5-4 µm (fide
Singer, 1976). Marasmius rosulatus differs in
forming pure white basidiomes with
basidiospores 7.8-9.6 × 4.0-4.8 µm and grows
on Cyperales leaves (Desjardin and Petersen,
1989).
2. Marasmius
delicatulus
Desjardin & Lumyong, sp. nov.
Wannathes,
(Figs 3, 17-1)
MycoBank: MB512413
Etymology: ‘delicatulus’ = delicate; referring to
the tiny, delicate basidiomes.
Pileus 1-3 mm diametro, convexus usque late
convexus, subumbilicatus, sine papilla, striatus glaber,
hebetatus, flavo-albus usque brunneo-creameus cum
macula laete brunnea ad centrum. Contextus flavo-albus,
tenuis. Lamellae adnatae usque collarium parvum,
distantes (6-10), angustae, flavo-albae usque creameae,
haud marginatae, haud intervenosae. Stipes 12 × 0.1-0.2
mm, centralis, cylindratus, filo metallico similis, nodis
dispersis, glaber, insititius, brunneus usque rubrobrunneus, plerumque rhizomorphis brunneis. Odor
saporque non propria. Basidiosporae 8-10 × 3.5-5 µm,
ellipsoideae,
laeves,
hyalinae,
inamyloideae,
tenuitunicatae. Basidia non observata. Basidiolae
221
fusoideae usque clavatae. Cheilocystidia vulgaria, typi
Rotalis; 13-16 × 9-13 µm, clavata usque late clavata,
subglobosa, hyalina, inamyloidea, tenuitunicata; setulis
divergentibus 1-2 × 1 µm, gongylodibus usque conicis,
obtusis, confertis ad dimidium superiorem, hyalinis
usque pallide flavis, crassetunicatis. Pleurocystidia nulla.
Pileipellis hymeniformis, haud maculosus, typi Rotalis;
18-23 × 14-18 µm, late clavatus usque subglobosus,
pyriformis, hyalinus, inamyloideus, tenuitunicatus;
setulis divergentibus 1-2 × 1 µm, gongylodibus usque
conicis, obtusis, confertis ad dimidium superiorem,
brunneo-flavis usque laete brunneis, crassetunicatis.
Trama pilei intertexta, inamyloidea. Trama lamellae
intertexta, hyphis 4-8 µm diametro, cylindratis usque
inflatis, laevibus, hyalinis, inamyloideis, tenuitunicatis,
haud gelatinosis. Stipitipellis parallelus, hyphis 3-5 µm
diametro, cylindratis, brunneis, laevibus, leniter
dextrinoideis usque inamyloideis, crassetunicatis (usque
ad 1 µm), haud gelatinosis. Trama stipitis parallela,
hyphis 2-4 µm diametro, cylindratis, hyalinis, laevibus,
inamyloideis,
tenuitunicatis,
haud
gelatinosis.
Caulocystidia nulla. Fibulae praesentes in omnibus
texturis.
Typus: Phrae Province, Maung District, Pa Dang,
Nong Kam Village, Nong Kam reservoir, dispersus
usque gregarius in folis plantae dicotyledoneae, 18
August2005, N. Wannathes 426 (CMU: holotypus;
SFSU: isotypus).
Pileus 1-3 mm diam., convex to broadly
convex, subumbilicate, without papilla, striate,
glabrous, dull, yellowish white to brownish
cream with a light brown central spot. Context
yellowish white, thin. Lamellae adnate to a
small collarium, distant (6-10), narrow,
yellowish white to cream, non-marginate, nonintervenose. Stipe 5-12 × 0.1-0.2 mm, central,
Basidiospores 8-10 × 3.5-5 µm [xmr = 8.6
× 3.9-4.2 µm, xmm = 8.6 ± 0 × 2.7 ± 0.3 µm,
Qmr = 2.1-2.2, Qmm = 2.2 ± 0.1, n = 25 spores, s
= 2 specimens], ellipsoid, smooth, hyaline,
inamyloid, thin-walled. Basidia not observed.
Basidioles fusoid to clavate. Cheilocystidia
common, of Rotalis-type broom cells; main
body 13-16 × 9-13 µm, clavate to broadly
clavate, subglobose, hyaline, inamyloid, thinwalled; divergent setulae 1-2 × 1 µm, knob-like
to conical, obtuse, dense over upper half of cell,
hyaline to pale-yellow, thick-walled. Pleurocystidia absent. Pileipellis hymeniform, not
mottled, composed of Rotalis-type broom cells;
main body 18-23 × 14-18 µm, broadly clavate
to subglobose, pyriform, hyaline, inamyloid,
thin-walled; divergent setulae 1-2 × 1 µm,
knob-like to conical, obtuse, dense over upper
half of cell, brownish yellow to light brown,
thick-walled. Pileus trama interwoven, inamyloid. Lamellar trama hyphae 4-8 µm diam.,
interwoven, cylindrical to inflated, smooth,
hyaline, inamyloid, thin-walled, non-gelatinous.
Stipitipellis hyphae 3-5 µm diam., parallel,
cylindrical, brown, smooth, weakly dextrinoid
to inamyloid, thick-walled (up to 1 µm), nongelatinous. Stipe trama hyphae 2-4 µm diam.,
parallel, cylindrical, hyaline, smooth, inamyloid,
thin-walled, non- gelatinous. Caulocystidia
absent. Clamp connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Chiang Mai University, back area of Ang
Kaew reservoir, 18 August 2004, N. Wannathes 255
(CMU, SFSU); Phrae Province, Maung District, Pa Dang,
Nong Kam Village, Nong Kam reservoir, 18
August2005, N. Wannathes 426 (CMU: holotype; SFSU:
isotype).
Fig. 3. Marasmius delicatulus (N. Wannathes 426). 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pileipellis Scale bar 1 = 4.5 mm, 2-4 = 10 µm
cylindrical, wiry, with scattered nodes,
glabrous, insititious, brown to reddish brown;
brown rhizomorphs usually present. Odor and
taste not distinctive.
222
Discussion: Distinctive features of M.
delicatulus include tiny basidiomes with
convex, yellowish white to brownish cream
pilei 1-3 mm diam. that lack a papilla,
collariate and distant lamellae (6-10), a small
central, insititious stipe not arising from
rhizomorphs, and moderately-sized basidiospores (xmm = 8.6 × 4.1 µm; Qmm = 2.2). The
new Thai species is morphologically similar to
several other species. Marasmius diminutivus
Y.S. Tan, Desjardin & Vikineswary, from
Malaysia, differs in forming smaller (0.5-1 mm
Fungal Diversity
diam.), conical pilei with a dark brown papilla,
and slightly longer and narrower basidiospores
with mean Q = 2.4 (Tan et al., 2009).
Marasmius leucorotalis Singer, as reported
from Java by Desjardin et al. (2000), differs in
forming larger, pure white pilei up to 7 mm
diam. with a distinct black conical papilla, a
larger stipe up to 32 mm long, and has slightly
smaller basidiospores with mean 8.4 x 3.7 µm.
Marasmius arimanus Dennis, from Trinidad,
differs in forming mummy brown pilei and
basidiospores 11-14 × 3.5-4 µm, while
Marasmius manuripiensis Singer, from Bolivia,
differs in forming white pilei 2-5 mm diam and
a more robust stipe up to 0.5 mm thick (Singer,
1976).
Antonín,
3. Marasmius
somalomoensis
Mycotaxon 88: 66. 2003.
(Figs 4, 17-2)
Pileus 2-4 mm diam., convex, umbilicate,
with a tiny papilla when young, papilla absent
in age, sulcate, glabrous, dull, with a reddish
brown spot in centre, surrounded by a pale
zone, reddish brown with a hint of olive
elsewhere. Context cream, thin. Lamellae
adnate to a small collarium, subdistant (12-18),
broad (0.5-1 mm), cream with brown edge,
non-intervenose. Stipe 8-47 × 0.3 mm, central,
cylindrical, wiry, with scattered nodes,
glabrous, insititious, apex cream, base dark
brown; rhizomorphs absent. Odor and taste not
distinctive.
Basidiospores 8-10 × 4-5 µm [xm = 8.8
± 0.6 × 4.3 ± 0.3 µm, Q = 1.8 - 2.2, Qm = 2.1, n
= 25 spores, s = 1 specimen], ellipsoid, smooth,
hyaline, inamyloid, thin-walled. Basidia not
observed. Basidioles fusoid to clavate.
Cheilocystidia common, of Rotalis-type broom
cells; main body 11-19 × 7-11 µm, clavate to
broadly clavate, hyaline, inamyloid, thinwalled; divergent setulae 2-3 × 1 µm, knob-like
to conical, obtuse, dense over upper half of cell,
brownish yellow to yellow, thick-walled.
Pleurocystidia absent. Pileipellis hymeniform,
not mottled, composed of Rotalis-type broom
cells; main body 18-23 × 14-18 µm, broadly
clavate, subglobose, turbinate to pyriform,
hyaline, inamyloid, thin-walled; divergent
setulae 1-3 × 1 µm, knob-like to conical, obtuse,
dense over upper half of cell, brown to light
Fig. 4. Marasmius somalomoensis (N. Wannathes 232).
1. Basidiomes 2. Basidioles 3. Cheilocystidia 4.
Basidiospores 5. Pileipellis, Scale bar 1 = 10 mm, 2-5 =
10 µm
brown, thick-walled. Pileus trama interwoven,
inamyloid. Lamellar trama hyphae 2-4(-8) µm
diam., interwoven, cylindrical to inflated,
smooth, hyaline, inamyloid, thin-walled, nongelatinous. Stipitipellis hyphae 3-5 µm diam.,
subparallel, cylindrical, dark brown, smooth,
weakly dextrinoid to inamyloid, thick-walled
(up to 1 µm), non-gelatinous. Stipe trama
hyphae 3-6 µm diam., subparallel, cylindrical,
hyaline, smooth, inamyloid, thin- to thickwalled (up to 1 µm), non-gelatinous.
Caulocystidia absent. Clamp connections
present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Sangasabhasri
Land N 18º 48.4’ E 98º 54.6’ alt. 1,146 m., 27 July 2004,
N. Wannathes 232 (CMU, SFSU).
Discussion: Maramius somalomoensis,
originally described from material collected in
Cameroon, is characterized by having a light
brown to brownish orange pileus with darker
papilla surrounded by a pale zone, moderately
distant lamellae, basidiospores in the range 710 × 3.7-5.5 µm, and Rotalis-type pileipellis
cells (Antonín, 2003). The Thai material differs
from the protologue only in forming pilei with
more reddish brown tones, more lamellae (1218 vs 11-13) with brown edges, and slightly
longer stipes (up to 47 mm vs 10-17 mm).
4. Marasmius apatelius Singer, Bull, Jard.
Bot.Burx. 43: 332. 1964.
(Figs 5, 17-3)
223
Pileus 2-10 mm diam., hemispherical to
convex with or without a small papilla in
umbilicus, striate, glabrous, dull, light
brownish cream to yellowish brown with a
brown central spot. Context yellowish white to
greyish cream, thin. Lamellae adnate to a small
collarium, subdistant (9-16), narrow, cream
with or without brown edges, non-intervenose.
Stipe 7-33 × 0.1-0.4 mm, central, cylindrical,
wiry, glabrous, insititious, brown to reddish
brown; rhizomorphs absent. Odor and taste not
distinctive.
Basidiospores (6-)8-9(-10.5) × 3.5-4(5.5)
µm [xmr = 7.2-8.9 × 3.9-4.6 µm, xmm = 8.0 ±
0.6 × 4.0 ± 0.4 µm, Qmm = 2.0 ± 0.2, Qmr = 1.72.3, n = 25 spores, s = 7 specimens], ellipsoid,
curved in profile, smooth, hyaline, inamyloid,
thin-walled. Basidia 16-24 × 7-9 µm, clavate 4spored. Basidioles clavate to fusoid.
Cheilocystidia common, of Rotalis-type broom
cells; main body (13)17-32(-34) × (7-)12-20
µm, clavate to broadly clavate, pyriform,
hyaline, inamyloid, thin-walled; divergent
setulae 0.5-4 × 0.8-1.5 µm, knob-like to conical,
obtuse, pale-yellow to light brown, thickwalled. Pleurocystidia absent or of scattered
Rotalis-type broom cells like the cheilocystidia.
Pileipellis hymeniform, not mottled, composed
of Rotalis-type broom cells; main body 16-35 ×
10-20 µm, clavate to broadly clavate, pyriform
or turbinate, hyaline, inamyloid, thin- to thickwalled; divergent setulae 1-3 × 0.5-1 µm,
knob-like to conical, obtuse, dense over upper
half of cell, yellow to light brown, thick-walled.
Pileus trama interwoven, inamyloid. Lamellar
trama hyphae 3-6(-13) µm diam., interwoven
to regular, cylindrical to inflated, smooth,
hyaline, inamyloid, thin-walled, non-gelatinous.
Stipitipellis hyphae 3-6 µm diam., parallel to
subparallel, cylindrical, light brown to brown,
smooth, weakly dextrinoid to inamyloid, thinto thick-walled (up to 1 µm), non-gelatinous.
Stipe trama hyphae (2-)3-7(-10) µm diam.,
parallel, cylindrical, hyaline, smooth, inamyloid, thin-walled, non-gelatinous. Caulocystidia
absent. Clamp connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
224
Fig. 5. Marasmius apatelius (N. Wannathes 251). 1.
Basidiomes 2. Basidioles and basidia 3. Basidiospores 4.
Cheilocystidia 5. Pileipellis, Scale bar 1 = 10 mm, 2-5 =
10 µm
leaves, bamboo debris or wood, Africa and
northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mokfa
Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
m. 3 July 2003, N. Wannathes 017 (CMU, SFSU); same
location, 21 August 2004, N. Wannathes 265 (CMU);
Chiang Mai Province, Mushroom Research Centre, 27
km marker on Hwy 1095, 27 July 2003, N. Wannathes
035 (CMU); same location, 27 July 2003, N. Wannathes
043 (CMU); Chiang Mai province, Chiang Mai
University, 11 August 2004, N. Wannathes 251 (CMU,
SFSU); Phrae Province, Maung District, Pa Dang, Nong
Kam Village, near Nong Kam reservoir, 18 August 2005,
N. Wannathes 427 (CMU, SFSU); Chiang Mai Province,
Chiang Mai University, close to Ang Kaew reservoir, 22
August 2005, N. Wannathes 437 (CMU, SFSU)
Discussion: The specimens from
Thailand matched quite closely the descriptions
of populations from Africa (Singer 1965;
Pegler 1977), except for forming a few more
lamellae per basidiome (13-14 vs 9-12).
Distinctive features of this species include an
entirely brown, umbilicate pileus lacking a
dark central spot, collariate, subdistant lamellae
with or without a pale brown edge, an absence
of well-developed rhizomorphs, and small
basidiospores with mean 7.2 × 3.9 µm. This is
the first report of this species outside Africa.
Marasmius tubulatus Petch, is similar, but
Fungal Diversity
differs in forming darker brown pilei, fewer
lamellae (10-13) that are also broader, and
slightly longer basidiospores with mean 9.3 × 4
µm (Tan et al., 2007).
A tetrapolar mating system was reported
for Thai specimens of M. apatelius by
Wannathes et al. (2007).
Section Marasmius, subsect. Sicciformes
Antonín, Acta Mus. Moraviae, Sci. Nat. 76:
145. 1991.
Type: Marasmius curreyi Berk. &
Broome.
= subsect. Penicillati Singer sensu Singer, Fl.
Neotrop. Monogr. 17: 121. 1976.
[Type: Marasmius graminum (Lib.) Berk.
sensu Singer].
5. Marasmius berambutanus Desjardin, Retn.
& E. Horak, Sydowia 52: 116. 2000. (Fig. 6)
Pileus 0.2-2 mm diam., hemispherical to
convex, umbilicate, with or without a reddish
brown to black papilla, striate, velutinous to
hispidulous, dull, cream-brown. Context cream,
thin. Lamellae adnate to a small collarium,
distant (8), broad, cream, non-marginate, nonintervenose. Stipe 3-12 × 0.1-0.2 mm, central,
cylindrical, wiry, with scattered nodes,
glabrous, insititious, apex white, base black,
arising directly from black rhizomorphs. Odor
and taste not distinctive.
Basidiospores 7-10 × (3.5-)4-5.5 µm [xm
= 8.4 ± 0.6 × 4.7 ± 0.6 µm, Q = 1.4 - 2.3, Qm =
2.1, n = 25 spores, s = 1 specimen], ellipsoid,
smooth, hyaline, inamyloid, thin-walled.
Basidia not observed. Basidioles cylindrical to
clavate. Cheilocystidia abundant, of Siccustype broom cells; main body 13-20 × 7-11 µm,
cylindrical to clavate or broadly clavate,
hyaline, inamyloid, thin-walled; apical setulae
3-19 × 1.5-2.5 µm, cylindrical, often wavy and
forked, obtuse to subacute, hyaline, thin-walled.
Pleurocystidia absent. Pileipellis hymeniform,
weakly mottled, composed of 3 types of cells: a)
Siccus-type broom cells with main body 9-20 ×
8-12 µm, clavate to broadly clavate or pyriform,
hyaline, inamyloid, thin-walled; apical setulae
3-15 × 1-2.5 µm, conical to cylindrical, obtuse
to subacute, yellow, thin-walled; b) transitional
cells, similar to Siccus-type broom cells with
main body 5-11 × 6-9 µm, cylindrical to
clavate, with 3-5 large apical setulae 9-28 × 2-3
Fig. 6. Marasmius berambutanus (N. Wannathes 044). 1.
Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5a. Siccus-type pileipellis 5b. Cells
transitional between broom cells and setae 5c. Pileosetae,
Scale bar 1 = 2 mm, 2-5 = 10 µm
µm, cylindrical, conical to subacute, thickwalled; c) pileosetae 36-82 × 6-8 µm, common,
lanceolate to fusoid, hyaline to yellow,
inamyloid, thick-walled. Pileus trama interwoven, inamyloid. Lamellar trama hyphae 3-6 µm
diam., interwoven, cylindrical to inflated,
smooth, hyaline, inamyloid, thin-walled, nongelatinous. Stipitipellis hyphae (2-)3-5 µm
diam., subparallel, cylindrical, dark brown,
smooth, dextrinoid, thick-walled (up to 1 µm),
non-gelatinous. Stipe trama hyphae 2-4 µm
diam., subparallel, cylindrical, hyaline, smooth,
inamyloid, thin-walled, non-gelatinous.
Caulocystidia absent. Clamp connections
present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on undetermined
dicotyledonous leaves or on wood, Java,
Malaysia and northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Mushroom Research Centre, 27 km marker on
Hwy 1095, 27 June 2003, N. Wannathes 004 (CMU).
Discussion: Marasmius berambutanus is
characterized by a very small, cream-brown
225
pileus with reddish brown to black papilla and
a suface that is covered with small setae,
collariate lamellae, and stipes that arise directly
from rhizomorphs. The Thai material differs
subtly from that described by Desjardin et al.
(2000) from Indonesia in forming pilei with
more brownish tones, and in forming
basidiospores that are slightly shorter (mean
8.4 × 4.7 µm vs 9.4 × 4 µm), and fits nicely
into the range of variation reported by Tan et al.
(2007) from populations in Malaysia.
A tetrapolar mating system was reported
by Tan et al. (2007) based on Malaysian
material.
6. Marasmius purpureisetosus Corner, Nova
Hedwigia 111: 90. 1996.
(Figs 7, 17-5)
Pileus 1-4 mm diam., hemispherical to
convex, striate, hispid with brown to reddish
brown setae, dull, margin orangish cream, disc
brown and dark brown spot at centre. Context
cream, thin. Lamellae adnate to a small
collarium, subdistant (13-17), broad (0.5-1
mm), cream, non-marginate, non-intervenose.
Stipe 15-55 × 0.2-0.3 mm, central, cylindrical,
wiry, with scattered nodes, glabrous, insititious,
apex creamish brown, base light brown to
brown; rhizomorphs present. Odor and taste
not distinctive.
Basidiospores 8-10 × 3-5 µm [xm = 8.6 ±
0.6 × 3.9 ± 0.4 µm, Q = 1.6 - 2.9, Qm = 2.3, n =
25 spores, s = 1 specimen], ellipsoid, smooth,
hyaline, inamyloid, thin-walled. Basidia not
observed. Basidioles fusoid to clavate.
Cheilocystidia abundant, composed of 2 types
of cells: a) Siccus-type broom cells with main
body 13-16 × 7-10 µm, clavate to broadly
clavate, hyaline to yellow, inamyloid, thickwalled; apical setulae 5-10 × 2-3 µm,
cylindrical to conical, obtuse, ranging from 2-4
per cell, yellow, thick-walled; b) non-setulose
cells, 23-38 × 6-8 µm, fusoid, hyaline,
inamyloid, thin- to thick-walled. Pleurocystidia
absent. Pileipellis hymeniform, not mottled,
composed of 3 types of cells: a) Siccus-type
broom cells with main body 14-20 × 8-15 µm,
clavate to broadly clavate or pyriform, hyaline
to pale yellow, inamyloid, thick-walled; apical
setulae 4-15 × 2-3 µm, conical to cylindrical,
obtuse, brown to dark brown, thick-walled; b)
transitional cells, similar to Siccus-type broom
226
Fig. 7. Marasmius purpureisetosus (N. Wannathes 155).
1. Basidiomes 2. Basidiospores 3a. Cheilocystidia 3b.
Cheilosetae 4a. Siccus-type pileipellis 4b. Cells transitional
between broom cells and setae 4c. Peleosetae, Scale bar 1 =
6.5 mm, 2-4 = 10 µm
cells with main body 17-36 × 9-12 µm, clavate
to broadly clavate, with only 2-3 large apical
setulae 15-27 × 3-5 µm, cylindrical, conical to
subacute, thick-walled; c) pileosetae 80-350 ×
8-12 µm, common, lanceolate to fusoid, brown,
inamyloid, thick-walled (up to 3 µm). Pileus
trama interwoven, inamyloid. Lamellar trama
hyphae 3-8(-10) µm diam., interwoven, cylindrical to inflated, smooth, hyaline, inamyloid,
thin-walled, non-gelatinous. Stipitipellis hyphae 3-5 µm diam., subparallel, cylindrical,
dark brown, smooth, inamyloid to weakly
dextrinoid, thick-walled (up to 1 µm), nongelatinous. Stipe trama hyphae 3-6 µm diam.,
subparallel, cylindrical, hyaline, smooth,
weakly dextrinoid, thin-walled, non-gelatinous.
Caulocystidia absent. Clamp connections
present in all tissues.
Habit, habitat and known distribution.
Scattered to gregarious on undetermined
dicotyledonous leaves, Java, Malaysia,
northern Thailand and Singapore.
Fungal Diversity
Material examined: Thailand. Chiang Mai
Province, Doi Inthanon National Park, At 25 km marker
on Highway 1,009, N 18º 32.5’ E 98º 33.5’ alt. 1,076 m.,
10 June 2004, N. Wannathes 155 (CMU, SFSU).
Discussion: Marasmius purpureisetosus is
characterized by a small pileus colored
orangish cream with a brown disc and dark
brown central spot, a hispid surface with brown
to reddish brown pileosetae, subdistant and
collariate lamellae, and a glabrous stipe with
scattered nodes. Marasmius purpureisetosus
and M. berambutanus are the only known
members of section Marasmius with pileosetae.
The two species differ by the features presented
in the Key. The Thai specimens are nearly
indistinguishable from those reported from
Indonesia (Desjardin et al., 2000) and Malaysia
(Corner, 1996; Tan, 2008), although the
pileosetae lacked purple tones.
7. Marasmius brevicollus Corner, Beih. Nova
Hedwigia 111: 37. 1996.
(Fig. 8)
Pileus 0.5-6 mm diam., hemispherical,
umbilicate, striate to sulcate, with or without a
papilla, glabrous to minutely velutinous, dull,
brownish red to brownish purple. Context
cream, thin. Lamellae adnate to a small
collarium, distant to subdistant (6-9), broad,
cream with brown edge, non-intervenose. Stipe
10-25 × 0.1-0.2 mm, central, cylindrical, wiry,
with or without nodes, glabrous, insititious,
brownish yellow, rhizomorph absent. Odor and
taste not distinctive.
Basidiospores (16-)18-22(-23) × (3.5-)45.5 µm [xmr = 19.0-21.1 × 4.0-5.1 µm, xmm =
19.7 ± 0.6 × 4.6 ± 0.5 µm, Qmr = 3.8-4.9, Qmm
= 4.6 ± 0.5, n = 25 spores, s = 5 specimens],
clavate to fusoid, smooth, hyaline, inamyloid,
thin-walled. Basidia 30-40 × 10-12 µm,
cylindrical to clavate, 4-spored. Basidioles
fusoid to clavate. Cheilocystidia abundant, of
Siccus-type broom cells; main body 12-26 × 615(-19) µm, cylindrical to clavate, broadly
clavate or pyriform, often branched, hyaline,
inamyloid, thin-walled; apical setulae 1-6 × 11.5 µm, conical to cylindrical, obtuse to
subacute, yellowish brown to dark brown,
thick-walled. Pleurocystidia absent. Pileipellis
hymeniform, mottled, composed of Siccus-type
broom cells; main body 7-27(-30) × 9-15 µm,
clavate to broadly clavate, pyriform or irregular
in outline, usually branched, hyaline, inamyloid,
Fig. 8. Marasmius brevicollus (N. Wannathes 128). 1.
Basidiomes 2. Basidia 3. Basidiospores 4. Cheilocystidia
5. Pileipellis, Scale bar 1 = 6.5 mm, 2-5 = 10 µm
thin- to thick-walled; apical setulae 1-3(-6) × 11.5 µm, conical to cylindrical, subacute to
obtuse, yellowish brown to dark brown, thickwalled. Pileus trama interwoven, inamyloid.
Lamellar trama hyphae 3-5(-7) µm diam.,
interwoven, cylindrical to inflated, smooth,
hyaline, inamyloid, thin-walled, non-gelatinous.
Stipitipellis hyphae 2-5 µm diam., subparallel,
cylindrical, yellowish brown, smooth, dextrinoid, thin-walled, non-gelatinous. Stipe trama
hyphae 2-5 µm diam., parallel, cylindrical,
hyaline, smooth, inamyloid, thin-walled, nongelatinous. Caulocystidia absent. Clamp
connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on undetermined
monocotyledonous leaves and stems, Malaysia,
northern Thailand and Singapore.
Material examined: Thailand. Chiang Mai
Province, Mae Taeng District, Pa Deng Village,
Pathammikaram temple on Hwy 1095, 7 August 2003, N.
Wannathes 094(CMU) and 095 (CMU, SFSU); same
location, 23 August 2003, N. Wannathes 123 and 126
(CMU); same location, 25 August 2003, N. Wannathes
128 (CMU, SFSU).
Discussion: Diagnostic features of
Marasmius brevicollus include striate to
227
sulcate, brownish red to brownish purple pilei,
distant and collariate lamellae with brown
edges, a brownish yellow stipe, and clavate
basidiospores with mean 19.7 × 4.6 µm. The
Thai material matches quite closely the
Malaysian material reported by Tan et al.
(2007) except that the Thai populations form
slightly larger basidiospores (with mean 19.7 ×
4.6 µm vs 18.8 × 3.7 µm) and fewer lamellae
(6-9 vs 9-11). Both Thai and Malaysian
specimens differ from the protologue based on
Singaporean material (Corner, 1996) in
forming a shorter stipe (10-25 mm vs 20-60
mm) and paler stipe color (brownish yellow vs
dark brown). The Singaporean material has a
broad range of the lamellae number (6-12)
which encompasses the range of lamellae
number for Thai and Malaysian populations.
Corner (1996) suggested that M. brevicollus
was allied with M. nigrobrunneus (Pat.) Sacc.,
but our molecular data (Fig. 1) indicate that
M. brevicollus is allied with M. purpureobrunneolus and together they are placed in a
clade distantly related to M. nigrobrunneus.
A tetrapolar mating system was reported
for M. brevicollus by Tan et al. (2007) based on
Malaysian material.
8. Marasmius purpureobrunneolus Henn.,
Monsunia 1: 151. 1900.
(Fig. 9)
= Marasmius acierufus Corner, Beih. Nova
Hedwigia 111: 25. 1996
Pileus 2-5 mm diam., hemispherical to
truncately conical, umbilicate, with or without
a papilla, striate, pruinose to pubescent, dull,
dark reddish brown (9F8) to violet brown
(11F4) with a reddish brown spot in centre.
Context cream, thin. Lamellae adnate to a small
collarium, subdistant (9-13), narrow, yellowish
white with reddish brown edge, nonintervenose. Stipe 10-32 × 0.1-0.2 mm, central,
cylindrical, wiry, with or without nodes,
glabrous, insititious, brown (6E5) to dark
brown (7F7) overall; rhizomorphs absent. Odor
and taste not distinctive.
Basidiospores (12-)13-16 × 3-4 µm [xmr
= 14.0-14.4 × 3.2-3.3 µm, xmm = 14.2 ± 0.2 ×
3.2 ± 0.1 µm, Qmr = 4.4-4.5, Qmm = 4.4 ± 0.1, n
= 25 spores, s = 2 specimens], narrowly
ellipsoid to fusoid, smooth, hyaline, inamyloid,
thin-walled. Basidia not observed. Basidioles
fusoid to clavate. Cheilocystidia abundant, of
228
Fig. 9. Marasmius purpureobrunneolus (N. Wannathes
370). 1. Basidiomes 2. Cheilocystidia 3. Basidiospores 4.
Pileipellis, Scale bar 1 = 13.5 mm, 2-4 = 10 µm
Siccus-type broom cells; main body 12-17 × 813 µm, clavate to broadly clavate or irregular
in outline, hyaline, inamyloid, thin- to thickwalled; apical setulae 1-3 × 1 µm, cylindrical
to conical, obtuse to subacute, pale yellow,
thick-walled. Pleurocystidia absent. Pileipellis
hymeniform, weakly mottled, composed of
Siccus-type broom cells; main body 15-25 × 715 µm, clavate to broadly clavate, pyriform or
irregular in outline, hyaline, inamyloid, thin- to
thick-walled; apical setulae 1-3 × 1 µm, conical
to cylindrical, subacute to obtuse, yellowish
brown, thick-walled. Pileus trama interwoven,
inamyloid. Lamellar trama hyphae 4-8(-13)
µm diam., regular to interwoven, cylindrical to
inflated, smooth, hyaline, inamyloid, thinwalled, non-gelatinous. Stipitipellis hyphae 3-5
µm diam., subparallel, cylindrical, yellowish
brown to brown, smooth, dextrinoid, thickwalled (up to 1 µm), non-gelatinous. Stipe
trama hyphae (2-)3-6 µm diam., subparallel,
cylindrical, hyaline, smooth, inamyloid, thinwalled, non-gelatinous. Caulocystidia absent.
Clamp connections present in all tissues.
Habit, habitat and known distributions.
Scattered to gregarious on undetermined
dicotyledonous leaves or on wood, Java,
Malaysia and northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Mae Taeng District, “Water Conservation taste
not distinctive. Area” Highway 1,095 at 22 km marker,
N. 19º 7.5’ E 98º 45.7’ alt. 724 m. 5 July 2004, N.
Wannathes 215 (CMU, SFSU); Chiang Mai Province,
Doi Inthanon National Park, At 25 km marker on
Highway 1,009, N 18º 32.5’ E 98º 33.5’ alt. 1,076 m., 27
June 2005, N. Wannathes 370 (CMU, SFSU).
Discussion: Marasmius purpureobrunneolus is characterized by striate, dark reddish
brown to violet brown pilei, subdistant (9-13)
lamellae with reddish brown edges, fusoid
Fungal Diversity
basidiospores with mean 14.2 × 3.2 µm, and
growth on dicotyledonous debris. We follow
the species concept of M. purpureobrunneolus
redefined by Desjardin et al. (2000) based on
recently collected specimens from the type
locality in the Bogor Botanical Garden, Java.
The Thai specimens match nicely the data
reported for material from Java (Desjardin et
al., 2000) and Malaysia (Tan, 2008). In the
field M. purpureobrunneolus and M. brevicollus look very similar in size and color of the
pilei. The two species differ in lamellae
spacing (9-13 lamellae vs 6-9, respectively)
and in basidiospore size (xmm = 14.2 × 3.2 µm
vs 19.7 × 4.6 µm, respectively).
9. Marasmius pallenticeps Singer, Fl. Neotrop.
Monogr. 17: 127. 1976
(Fig. 10)
Pileus 0.5-1.5 mm diam., convex,
subumbilicate with a papilla, striate, pruinose
to glabrous, dull, cream to pale yellowish white
(2A2) with a dark brown papilla. Context
cream, thin. Lamellae adnate to a small
collarium, distant (7-9), narrow, cream, nonmarginate, non-intervenose. Stipe 3-6 × 0.1 mm,
central, cylindrical, wiry, glabrous, insititious,
dark brown to black overall, arising directly
from black rhizomorphs. Odor and taste not
distinctive.
Basidiospores (7-)8-10(-11) × 4.5-5.5 µm
[xm = 9.0 ± 0.8 × 4.9 ± 0.3 µm, Q = 1.4 - 2.2,
Qm = 1.7, n = 25 spores, s = 1 specimen],
ellipsoid, smooth, hyaline, inamyloid, thinwalled. Basidia not observed. Basidioles fusoid
to clavate. Cheilocystidia abundant, of Siccustype broom cells; main body 11-18 × 5-10 µm,
cylindrical to clavate or broadly clavate,
hyaline, inamyloid, thin-walled; apical setulae
2-4 × 1(-1.5) µm, cylindrical to conical, obtuse
to subacute, pale yellow to hyaline, thickwalled. Pleurocystidia absent. Pileipellis
hymeniform, weakly mottled, composed of
Siccus-type broom cells; main body 9-13 × 411 µm, cylindrical to clavate or broadly clavate,
hyaline, inamyloid, thin-walled; apical setulae
1-4 × 1 µm, cylindrical to conical, subacute,
pale yellow, thick-walled. Pileus trama
interwoven, inamyloid. Lamellar trama hyphae
Fig. 10. Marasmius pallenticeps (N. Wannathes 346). 1.
Basidiomes 2. Cheilocystidia 3. Basidiospores 4.
Pileipellis, Scale bar 1 = 3.5 mm, 2-4 = 10 µm
3-5 µm diam., regular to interwoven,
cylindrical, smooth, hyaline, inamyloid, thinwalled, non-gelatinous. Stipitipellis hyphae 3-5
µm diam., subparallel, cylindrical, brown to
dark brown, smooth, inamyloid to weakly
dextrinoid, thick-walled (up to 1 µm), nongelatinous. Stipe trama hyphae 1.5-3 µm diam.,
subparallel, cylindrical, hyaline, smooth,
inamyloid,
thin-walled,
non-gelatinous.
Caulocystidia absent. Clamp connections
present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on wood (dicotyledonous tree), Argentina, New Zealand, Indonesia
and northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Mae Taeng District, Mae Sae Village, Pang Sa
Ded Water Conservation Area, N 19º 14.6', E 98º 38.5'
alt. 962 m., 26 June 2005, N. Wannathes 346 (CMU,
SFSU).
Discussion: Distinctive features of
Marasmius pallenticeps include a tiny pale
cream-coloured pileus with dark brown papilla,
distant (7-9), collariate lamellae, and a stipe
arising directly from rhizomorphs. Our
specimen is indistinguishable from the
specimen reported from Java by Desjardin et al.
(2000).
10. Marasmius crinis-equi F. Muell. ex
Kalchbr., Grevillea 8: 153. 1880.
(Fig. 11)
≡ 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 var. equicrinis (F. Muell.
ex Berk.) Dennis, Trans. Brit. Mycol. Soc. 34: 416. 1951.
229
= 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.
Pileus 0.5-4 mm diam., convex,
umbilicate, papillate, striate, glabrous, dull,
reddish brown to orange brown with dark
brown papilla or spot in the center. Context
cream, thin. Lamellae adnate to a small
collarium, distant (8-10), narrow, cream to
orangish
yellow,
non-marginate,
nonintervenose. Stipe 4-6 × 0.1-0.2 mm, central,
cylindrical, wiry, glabrous, insititious, dark
brown to black overall, arising directly from
copious, branched, black rhizomorphs. Odor
and taste not distinctive.
Basidiospores (8-)10-12 × 4-4.5 µm [xmr
= 10.5-10.6 × 4.1-5.1 µm, xmm = 10.6 ± 0.1 ×
4.6 ± 0.7 µm, Qmr = 2.1-2.6, Qmm = 2.3 ± 0.4, n
= 25 spores per 2 specimens], ellipsoid, smooth,
hyaline, inamyloid, thin-walled. Basidia 22-23
× 6-7 µm, clavate, 4-spored. Basidioles fusoid
to clavate. Cheilocystidia abundant, of Siccustype broom cells; main body 11-22 × 7-12 µm,
clavate to broadly clavate, subglobose or
irregular in outline, hyaline, inamyloid, thinwalled; apical setulae 2-5 × 1-2 µm, conical to
cylindrical, obtuse to subacute, pale yellow,
thick-walled. Pleurocystidia absent. Pileipellis
hymeniform, weakly mottled, composed of
Siccus-type broom cells; main body9-22 ×8-21
µm, clavate to broadly clavate, pyriform,
subglobose or irregular in outline, often
branched, hyaline, inamyloid, thin-walled;
apical setulae 1-5 × 1-1.5 µm, conical to
cylindrical, subacute to obtuse, yellow to
yellowish brown, thick-walled. Pileus trama
interwoven, inamyloid. Lamellar trama hyphae
3-8(-12) µm diam., regular to interwoven,
cylindrical to inflated, smooth, hyaline, inamyloid, thin-walled, non- gelatinous. Stipitipellis
hyphae (2-)3-6 µm diam., subparallel, cylindrical, brown to dark brown, smooth,
dextrinoid, thick-walled (up to 3 µm), nongelatinous. Stipe trama hyphae 2-5 µm diam.,
subparallel, cylindrical, hyaline, smooth,
inamyloid, thin-walled, non-gelatinous. Caulocystidia absent. Clamp connections present in
all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on wood, cosmo230
Fig. 11. Marasmius crinis-equi (N. Wannathes 182). 1.
Basidiomes 2. Cheilocystidia 3. Basidiospores 4.
Pileipellis, Scale bar 1 = 6.5 mm, 2-4 = 10 µm
politan in tropical habitats.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mokfa
Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
m., 25 July 2003, N. Wannathes 029 (CMU, SFSU);
Chiang Mai Province, Mae Taeng District, Mae Sae
Village, Pang Sa Ded Water Conservation Area, N 19º
14.6', E 98º 38.5' alt. 962 m., 26 June 2005, N.
Wannathes 348 (CMU, SFSU).
Discussion: Marasmius crinis-equi is one
of the more widely distributed species in the
genus, reported thoughout tropical areas of the
world. It is the causal agent of 'Horse-hair
Blight' that is known to be pathogenic on tea,
rubber, cocoa and nutmeg crops (Pegler, 1983).
This species grows on leaves and petioles of
living or dead trees, and on monocotyledonous
stems and clums (Singer, 1976) and its
rhizomorphs are often found covering aerial
branches of living trees. Marasmius crinis-equi
is easy to recognize by a small, brown to
brownish orange pileus with a dark brown
central papilla, distant collariate lamellae,
stipes that arise directly from black rhizomorphs, and Siccus-type pileipellis elements
and cheilocystidia.
11. Marasmius aff. crinis-equi F. Muell. ex
Kalchbr., Grevillea 8: 153. 1880
(Fig. 12)
Pileus 1-2 mm diam., hemispherical to
convex, shallowly umbilicate, papillate, striate
to sulcate, glabrous, dull, light orange (6A4) to
greyish orange with a brown papilla. Context
greyish cream, thin. Lamellae adnate to a small
collarium, distant (5-8), broad, cream, nonmarginate or with orangish cream edges, nonintervenose. Stipe 3-8 × 0.1-0.2 mm, central,
Fungal Diversity
cylindrical, wiry, glabrous, insititious, black
overall, arising directly from copious, branched,
black rhizomorphs. Odor and taste not
distinctive.
Basidiospores 8-10 × 4-6 µm [xmr = 8.99.2 × 4.6-5.1 µm, xmm = 9.0 ± 0.2 × 4.9 ± 0.4
µm, Qmr = 1.8-2.0, Qmm = 1.9 ± 0.1, n = 25
spores, s = 2 specimens], ellipsoid, smooth,
hyaline, inamyloid, thin-walled. Basidia not
observed. Basidioles fusoid to clavate. Cheilocystidia abundant, of Siccus-type broom cells;
main body 10-22 × 8-13 µm, clavate to broadly
clavate or pyriform, hyaline, inamyloid, thinwalled; apical setulae 1-4 × 1 µm, conical to
cylindrical, obtuse to subacute, yellow, thickwalled. Pleurocystidia absent. Pileipellis
hymeniform, weakly mottled, composed of
Siccus-type broom cells; main body 11-19 × 716 µm, clavate to broadly clavate, pyriform or
irregular in outline, hyaline, inamyloid, thin- to
thick-walled; apical setulae 2-7 × 1 µm,
cylindrical to conical, subacute to obtuse,
yellowish brown to brown, thick-walled. Pileus
trama interwoven, inamyloid. Lamellar trama
hyphae 3-8 µm diam., interwoven, cylindrical,
smooth, hyaline, inamyloid, thin-walled, nongelatinous. Stipitipellis hyphae 4-8 µm diam.,
subparallel, cylindrical, black to dark brown,
smooth, weakly dextrinoid to inamyloid, thickwalled (up to 1 µm), non-gelatinous. Stipe
trama hyphae 2-3(-5) µm diam., subparallel,
cylindrical, hyaline, smooth, inamyloid, thinwalled, non-gelatinous. Caulocystidia absent.
Clamp connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on wood, northern
Thailand.
Fig. 12. Marasmius aff. crinis-equi (N. Wannathes 348).
1. Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5. Pileipellis, Scale bar 1 = 4.5 mm, 2-5 =
10 µm
of this taxon is collected, we tentatively
identify it as M. aff. crinis-equi.
12. Marasmius subruforotula Singer, Bull.
Jard. Bot. Etat Brux. 34: 339. 1964 (Fig. 13)
Pileus 2-6 mm diam., hemispherical to
convex, umbilicate, with or without a papilla,
striate to sulcate, minutely velutinous, dull,
reddish brown to brown with dark brown
papilla or spot at the centre. Context cream,
thin. Lamellae adnate to a small collarium,
subdistant (11-13), narrow, cream-grey, nonmarginate, non-intervenose. Stipe 4-10 × 0.10.2 mm, central, cylindrical, wiry, glabrous,
shiny, insititious, black overall, arising from
leafy substrate or seldom directly from black
rhizomorphs. Odor and taste not distinctive.
Material examined: Thailand. Chiang Mai
Province, Doi Inthanon National Park, Junction of
Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E
98º 29.6’ alt. 1,703 m., 25 June 2004, N. Wannathes 182
(CMU, SFSU); same location, 3 July 2004, N.
Wannathes 205(CMU, SFSU)).
Discussion: The two specimens reported
here are morphologically similar to Marasmius
crinis-equi in all taxonomically significant
details except for having broader basidiospores
with Qmm = 1.9 (vs. 2.3) and paler pilei. The
molecular data (Fig. 1), however, indicate that
these specimens are not closely allied to M.
crinis-equi sensu stricto. Until further material
Fig. 13. Marasmius subruforotula (N. Wannathes 140).
1. Basidiomes 2. Cheilocystidia 3. Basidiospores 4.
Pileipellis, Scale bar 1 = 10 mm, 2-4 = 10 µm
Basidiospores 8-10 × 4-5 µm [xm = 9.5 ±
1.0 × 4.5 ± 0.5 µm, Q = 1.8 - 2.5, Qm = 2.1, n =
25 spores, s = 1 specimen], ellipsoid, smooth,
231
hyaline, inamyloid, thin-walled. Basidia not
observed. Basidioles fusoid to clavate. Cheilocystidia abundant, of Siccus-type broom cells;
main body 14-16 × 6-12 µm, cylindrical to
clavate or broadly clavate, hyaline, inamyloid,
thin-walled; apical setulae 2-5 × 1 µm,
cylindrical, often forked, subacute, pale yellow,
thick-walled. Pleurocystidia absent. Pileipellis
hymeniform, weakly mottled, composed of
Siccus-type broom cells; main body 8-20 × 1015 µm, clavate to broadly clavate or pyriform,
hyaline, inamyloid, thin-walled; apical setulae
3-5 × 1 µm, cylindrical to conical, subacute,
yellowish brown to brown, thick-walled. Pileus
trama interwoven, inamyloid. Lamellar trama
hyphae 5-12(-18) µm diam., interwoven,
cylindrical to inflated, smooth, hyaline,
inamyloid, thin-walled, non-gelatinous. Stipitipellis hyphae 3-6 µm diam., subparallel,
cylindrical, dark brown to black, smooth,
inamyloid, thick-walled (up to 1 µm), nongelatinous. Stipe trama hyphae 3-6 µm diam.,
subparallel, cylindrical, hyaline to yellow,
smooth, inamyloid, thin-walled, non-gelatinous.
Caulocystidia absent. Clamp connections
present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on undetermined
dicotyledonous leaves or on wood, Africa and
Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Sangasabhasri
Land on the way to Huai Kok Ma Village, N 18º 48.4’ E
98º 54.6’ alt. 1,146 m., 29 August 2003, N. Wannathes
140 (CMU, SFSU).
Discussion: Marasmius subruforotula is
characterzied by a small (2-6 mm), reddish
brown pileus, subdistant (11-13) collariate
lamellae, a short stipe 4-10 mm long, and
ellipsoid basidiospores with mean 9.5 × 4.5 µm.
The Thai specimens differ from M.
nigrobrunneus “f. cinnamomeus” in forming
reddish brown pilei and usually arising directly
from leaves, although some basidiome arise
from rhizomorphs. Our molecular data (Fig. 1)
suggest that this species is distinct from M.
nigrobrunneus and its forms, and is closer to M.
curreyi. Until further material is available for
comparison with the African, M. subruforotula,
we are only tentatively accepting the Thai
species as M. subruforotula.
232
13A. Marasmius nigrobrunneus (Pat.) Sacc. f.
nigrobrunneus Syll. Fung. 11:37. 1895.
(Figs 14, 17-4)
Pileus 3-28 mm diam, hemispherical to
convex when young, becoming plano-convex
and shallowly umbilicate in age, with or
without a small black papilla in the umbilicus,
striate to plicate; surface dull, dry, glabrous to
pruinose; disc dark grey to black, margin
greyish brown (6F3) to brownish grey (9F2).
Context thin, greyish white to cream. Lamellae
adnate to a collarium, subdistant (9-17) with no
lamellula, non-intervenose, broad (2-4 mm),
yellowish white when young, becoming paleyellow in age, with brown to dark brown edges.
Stipe 52-214 × 0.3-0.5 mm, central, cylindrical,
wiry, shiny, solid, rarely with a few nodes,
glabrous, insititious, apex olive brown grading
to dark brown at the base when young, dark
brown overall in age, arising directly from
black rhizomorphs. Odor and taste not
distinctive.
Basidiospores (6-)8-10(-13) × (3-)4-5(-7)
μm [xmr = 7.4-10.6 × 4.1-6.3 μm, xmm = 9.0 ±
0.9 × 4.7 ± 0.6 μm, Q = 1.4-2.9, Qmr = 1.7-2.2,
Qmm = 1.9 ± 0.1, n = 25 spores, s = 11
specimens], ellipsoid, smooth, hyaline,
inamyloid, thin-walled. Basidia 25-27 × 5.0-7.5
μm, clavate to cylindrical, 4-spored. Basidioles
fusoid to cylindrical. Cheilocystidia common,
of Siccus-type broom cells; main body (11-)1420(-30) × (4-)7-14(-19) μm, cylindrical to
clavate, pyriform or sometimes irregular in
outline, hyaline, thin- to thick-walled; apical
setulae (2-)3-9(-15) × 1-2(-2.5) μm, crowded,
cylindrical or irregular in outline, obtuse to
subacute, dark grey to black, inamyloid, thickwalled. Pleurocystidia absent. Pileipellis
hymeniform, mottled to weakly mottled,
composed of 3 types of cells: a) Siccus-type
broom cells with main body (11-)15-27 ×
(7-)10-15(-18) μm, clavate to broadly clavate,
pyriform or sometimes irregular in outline,
yellowish brown, inamyloid, thin- to thickwalled; apical setulae 2-9 × 1-2(-2.5) μm,
cylindrical to conical or irregular in outline,
obtuse to subacute, dark grey to black,
inamyloid,
thinto
thick-walled;
b)
Globulares-type cells, 14-28 × 11-12 μm,
subglobose to clavate, sometimes irregular in
Fungal Diversity
Fig. 14. Marasmius nigrobrunneus (N. Wannathes 294).
1. Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5a. Siccus-type pileipellis 5b. Cells
transitional between broom cells and non-setulose cells
5c. Globulares-type pileipellis, Scale bar 1 = 20 mm, 2-5
= 10 µm
outline, dark grey to black, inamyloid, thinwalled; c) cells transitional between broom
cells and non-setulose cells with main body 2022 × 15-17 μm, broadly clavate to pyriform
with a few knob- like apical setulae 6-13 × 4-8
μm, dark grey to black, thin- to thick-walled.
Pileus trama interwoven; lamellar trama
regular; hyphae 3-7 μm diam, cylindrical to
inflated, smooth, hyaline, inamyloid, thinwalled, non-gelatinous. Stipitipellis hyphae 4-8
μm diam, parallel, cylindrical, smooth, dark
brown, dextrinoid, thick-walled (up to 2 μm),
non-gelatinous. Stipe trama hyphae 3-8 μm
diam, parallel, cylindrical, smooth, hyaline,
inamyloid, thick-walled (up to 2 μm).
Caulocystidia absent. Clamp connections
present in all tissues.
Habit, habitat, and known distribution:
Scattered to gregarious on bamboo leaves and
stems, or rarely on undetermined dicotyledonous leaves. Africa, Caribbean region, South
America, India, Sri Lanka, Papua New Guinea,
Vietnam, Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Sangra
Sabhasri Land to Huai Kok Ma Village, N18°48.402’,
E98°54.617’, 1146 meters, 30 Jun. 2003, N. Wannathes
010 (CMU, SFSU); Chiang Mai Province, Doi SuthepPui National Park, Mok Fa waterfall, N19°6.581’,
E98°46.353’, 1014 meters, 13 Aug. 2003, N. Wannathes
101 (CMU, SFSU); same location, 29 Jun 2004, Y. S.
Tan 275 (CMU, SFSU); same location, 7 July 2004, N.
Wannathes 223 (CMU, SFSU); same location, 25 Jun
2005, N. Wannathes 339 (CMU, SFSU); Chiang Mai
Province, Mae Taeng District, Phadeng Village,
Pathummikaram temple, N19º06' 28.8", E98º44' 47.3",
1050 meters, 25 Aug 2003, N. Wannathes 129 (CMU,
SFSU); Chiang Mai Province, Mae Taeng District, Tung
Joaw Village, N19º8.07’, E98º38.90’, 1423 meters, 20
Jun 2004, N. Wannathes 162 (CMU, SFSU); Chiang Mai
Province, Mae Taeng District, Mae Sae Village, Pang Sa
Ded Water Conservation Area, N19º14.60', E98º38. 46',
962 meters, 26 Jun 2005, N. Wannathes 347 (CMU,
SFSU); Phrae Province, Maung District, Cherng Thong
waterfall, 16 Aug 2005, N. Wannathes 416 (CMU,
SFSU); same location, 19 Aug 2005, N. Wannathes 435
(CMU, SFSU); Chiang Rai Province, Maung District,
Khun Con waterfall, N19º5' 38.6", E99º39' 7.4", 577
meters, 12 Jun 2005, N. Wannathes 302 and 311 (CMU,
SFSU).
Discussion: The specimens of Marasmius
nigrobrunneus from northern Thailand match
quite closely those reported by Singer (1958,
1976) and Pegler (1983, 1986) from Sri Lanka,
India, Sierra Leone, Trinidad, Bolivia and
Argentina, including forming lamellae with
greyish brown edges and stipes that often arise
from black rhizomorphs. They differ only
slightly from specimens reported from Vietnam
(type; Patouillard, 1891) and Papua New
Guinea (Desjardin and Horak, 1997) that lack a
colored lamellar edge, and were not reported to
arise from black rhizomorphs. The species is
commonly found on bamboo debris and the
combination of dark grey to greyish brown
pileus, distant collariate lamellae with or
without colored edge, long, black, wiry stipes
often associated with rhizomorphs, and
versiform pileipellis broom cells are distinctive
features for the species. A recent type study
was published by Yang (2000).
A tetrapolar mating system was reported
by Wannathes et al. (2007) for M.
nigrobrunneus based on specimens from
northern Thailand.
13B. Marasmius nigrobrunneus “f. cinnamomeus” Wannathes, Desjardin & Lumyong
nom. prov.
(Fig. 15)
Etymology: ‘cinnamomeus’ = cinnamon; referring
to the color of the pileus.
Pileus 2-7(-15) mm diam., obtusely
conical, umbilicate, with or without a papilla,
striate to sulcate, glabrous to pruinous, dull,
brown to orangish brown (6C5-C8) with a
brown papilla or spot in the centre. Context
233
cream, thin. Lamellae adnate to a small
collarium, distant to subdistant (8-11(-15)),
broad, cream, non-marginate or with orange to
brown edges, non-intervenose. Stipe 8-60 ×
0.1-0.3 mm, central, cylindrical, wiry, glabrous,
insititious, brown to dark brown overall,
usually arising directly from rhizomorphs.
Odor and taste not distinctive.
Basidiospores 8-11 × 4-5(-6) µm [xmr =
8.9-10.1 × 4.3-5.4 µm, xmm = 9.5 ± 0.5 × 4.7 ±
0.4 µm, Qmr = 1.9-2.1, Qmm =2.0 ± 0.1, n = 25
spores, s = 7 specimens], ellipsoid, smooth,
hyaline, inamyloid, thin-walled. Basidia 31-37
× 6-10 µm, cylindrical to clavate, 4-spored.
Basidioles cylindrical to clavate. Cheilocystidia
abundant, of Siccus-type broom cells; main
body 10-21(-32) × 6-12 µm, cylindrical to
clavate or broadly clavate, hyaline, inamyloid,
thin-walled; apical setulae 2-4(-7) × 1-1.5 µm,
cylindrical to conical, obtuse to subacute, pale
yellow, thick-walled. Pleurocystidia absent.
Pileipellis hymeniform, weakly mottled,
composed of 2 types of cells: a) Siccus-type
broom cells with main body 8-18(-24) × 7-16
µm, clavate to broadly clavate, pyriform or
irregular in outline, hyaline, inamyloid, thin- to
thick-walled; apical setulae 2-6 × 1-1.5 µm,
cylindrical to conical, subacute to obtuse,
yellow to yellowish brown, thick-walled; b)
non-setulose cells, scattered, 12-17 × 8-15 µm,
clavate to broadly clavate, often branched,
hyaline, inamyloid, thick-walled. Pileus trama
interwoven, inamyloid. Lamellar trama hyphae
Fig. 15. Marasmius nigrobrunneus “f. cinnamomeus” (N.
Wannathes 119). 1. Basidiomes 2. Cheilocystidia 3.
Basidiospores 4. Pileipellis, Scale bar 1 = 20 mm, 2-4 =
10 µm
234
2-7(-12) µm diam., interwoven, cylindrical to
inflated, smooth, hyaline, inamyloid, thinwalled, non-gelatinous. Stipitipellis hyphae 34(-8) µm diam., subparallel, cylindrical, brown,
smooth, dextrinoid, thin- to thick-walled, nongelatinous. Stipe trama hyphae 3-6(-10) µm
diam., parallel, cylindrical, hyaline, smooth,
inamyloid, thin-walled, non-gelatinous. Caulocystidia absent. Clamp connections present in all
tissues.
Habit, habitat and known distribution:
Scattered to gregarious on undetermined
monocotyledonous leaves and stems, Northern
Thailand.
Material examined: Thailand. Chiang Mai
Province, Mae Taeng District, Pong Dued Hot Spring,
on Hwy1095, 25 July 2003, N. Wannathes 025 (CMU,
SFSU); Chiang Mai Province, Doi Suthep-Pui National
Park, Mokfa Waterfall, on Hwy 1095, N19º 6.5’.E 98º
46.3’ alt. 1014 m., 25 July 2003, N. Wannathes 030
(CMU); same location, 29 June 2004, Y. S. Tan 281
(CMU, SFSU); same location, 25 June 2005, N.
Wannathes 327 (CMU, SFSU); Chiang Mai Province,
Mae Taeng District, Pa Deng Village, street to
Pathammikaram temple on Hwy 1095, 22 August, 2003,
N. Wannathes 119 and 120 (CMU, SFSU); Chiang Mai
Province, Mae Taeng District, New Waterfall, on 36 km.
marker of Hwy1095, 20 August 2004, N. Wannathes 260
(CMU, SFSU).
Discussion: Marasmius nigrobrunneus “f.
cinnamomeus” is distinct because of the
following features: a brown to brownish orange
(cinnamon) pileus 2-7(-15) mm diam., a stipe
in the range 8-60 × 0.1-0.3 mm that arises
directly from rhizomorphs or from the substrate,
and is usually associated with non-bamboo
grasses. Marasmius nigrobrunneus f. nigrobrunneus differs in forming basidiomes with a
nearly black or dark greyish brown pileus,
much larger stipse (52-214 × 0.5 mm,) and
basidiomes that typically arise directly from a
bamboo substrate (only occasionally arising
from rhizomorphs). Marasmius curreyi Berk.
& Broome described from Europe and North
America, is another grass-loving species but it
differs in forming pilei with more orange tones,
and lacks of rhizomorphs.
A molecular dataset composed only of
ITS sequences of M. nigrobrunneus with M.
curreyi as outgroup, indicates that f.
nigrobrunneus forms a clade distinct from “f.
cinnamomeus”,
providing
support
for
recognition of the latter as a distinct form (Fig. 16)
Fungal Diversity
M. nigrobrunneus NW162
88
M. nigrobrunneus NW416
M. nigrobrunneus NW223
74
M. nigrobrunneus “f. cinnamomeus” NW119
M. nigrobrunneus “f. cinnamomeus” NW260
M. nigrobrunneus “f. cinnamomeus” TYS281
M. nigrobrunneus “f. cinnamomeus” NW025
M. nigrobrunneus “f. cinnamomeus” NW327
M. nigrobrunneus “f. cinnamomeus” NW120
M. curreyi DED 5142
5 changes
Fig. 16. Branch and bound tree of the Marasmius nigrobrunneus species complex. Numbers above branches represent
support from 2000 bootstrap replicates
14. Marasmius straminiceps Wannathes,
Desjardin, & Lumyong, Mycol. Res. 111: 991.
2007.
(Fig. 18)
Pileus 2-4 mm diam, hemispherical to
convex, subumbilicate to depressed, with or
without a papilla, striate, dull, dry, glabrous,
light yellow (4A4) to yellowish white (3A2)
with a reddish brown central spot. Context thin,
yellowish white (3A2). Lamellae adnate to a
collarium, distant (9-10) with no lamellula,
non-intervenose, narrow, cream, non-marginate.
Stipe 11-35 × 0.1-0.3 mm, central, cylindrical,
tough, terete, glabrous, insititious, yellowish
white at apex, brown to dark brown at base;
rhizomorphs absent. Odor and taste not
distinctive.
Basidiospores (7-)8-10(-12) × 4-5(-6.5)
μm [xmr = 8.4-9.4 × 4.4-5.1 μm, xmm = 8.9 ± 0.7
× 4.8 ± 0.5 μm, Q = 1.4-2.2, Qmr = 1.8-1.9, Qmm
= 1.8 ± 0.1, n = 25 spores, s = 2 specimens],
ellipsoid, smooth, hyaline, inamyloid, thinwalled. Basidia 25-30 × 6-8 μm, clavate, 4spored. Basidioles clavate to fusoid. Cheilocystidia common, of Siccus type broom cells;
main body (7-)10-18 × (4-)7-12 μm, clavate to
broadly clavate, pyriform, cylindrical or
irregular in outline, rarely lobed, hyaline, thinwalled; apical setulae (2-)4-6(-7) × 0.8-1.5 μm,
cylindrical to conical or irregular in outline,
obtuse to subacute, hyaline to pale-yellow,
inamyloid, thin- to thick walled. Pleurocystidia
absent. Pileipellis hymeniform, not mottled,
composed of Siccus-type broom cells; main
body 11-20 × 6-10 μm, cylindrical to clavate,
broadly clavate or sometimes irregular in
outline, rarely lobed, hyaline, thick-walled
apically, thin-walled elsewhere; apical setulae
(1-)3-6(-7) × 1-2 μm, cylindrical to conical,
usually wavy in outline, obtuse to subacute,
pale yellow, inamyloid, thin- to thick-walled.
Pileus trama and lamellar trama interwoven;
hyphae (2-)3-6 μm diam, cylindrical, smooth,
hyaline, inamyloid, thin-walled, non-gelatinous.
Stipitipellis hyphae 3-7 μm diam, parallel,
cylindrical, smooth, brown, weakly dextrinoid
to dextrinoid, thin-walled, non-gelatinous;
Stipe trama hyphae (2-)4-8 μm diam, parallel,
cylindrical, smooth, hyaline, weakly dextrinoid
to dextrinoid, thin- to thick-walled ( up to 1
μm). Caulocystidia absent. Clamp connections
present in all tissues.
Habit, habitat, and known distribution.
Scattered to gregarious on undetermined
dicotyledonous leaves or wood. Northern
Thailand.
Material examined. Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mok Fa
waterfall on Hwy 1095, N19º 6.581’, E98º 46.353’, 1014
meters, 13 Aug. 2003, N. Wannathes106 (CMU, SFSU);
Chiang Mai Province, Doi Inthanon National Park,
junction of Hwy1009 and road to Mae Chem, N18º
31.58’, E98º 29.64’, 1703 meters, 28 Jul. 2004, N.
Wannathes 244 (CMU: holotype; SFSU: isotype);
Chiang Mai Province, Mae Taeng District, New
Waterfall, on 36 km. marker of Hwy1095, N. Wannathes
256 (CMU, SFSU).
Discussion: Distinctive features
new species include the following:
yellow (stramineus), umbilicate pileus
reddish brown spot; collariate,
of this
a pale
with a
distant
235
Fig. 17. Basidiomata of Marasmius section Marasmius. 1. M. delicatulus (N. Wannathes 426) 2. M. somalomoensis (N.
Wannathes 232) 3. M. apatelius (N. Wannathes 427) 4. M. nigrobrunneus (N. Wannathes 435) 5. M. purpureisetosus (N.
Wannathes 155), Scale bars = 20 mm
(9-10), non-marginate lamellae; a lack of
obvious rhizomorphs; basidiospores with mean
of 8.9 × 4.8 µm and Qmm = 1.8; Siccus-type
broom cells on pileipellis and lamellar edges;
and growth on dicotyledonous leaves. These
features indicate placement in sect. Marasmius
subsect. Sicciformis. Marasmius straminiceps
is phenetically similar to M. conicopapillatus
Henn., but the latter species differs in forming
a white to cream-buff pileus with a distinctive,
rather large conical papilla, forms black
236
rhizomorphs, has thick-walled cheilocystidia,
and narrower basidiospores with mean 10.3 ×
4.3 µm and Qmm = 2.4 (Desjardin et al. 2000).
The new species is macromorphologically
similar to M. platyspermus Singer, a species
described from Argentina, but the latter forms
Rotalis-type broom cells and belongs in subsect.
Marasmius (Singer 1976).
A tetrapolar mating system was reported
for M. straminiceps by Wannathes et al. (2007).
Fungal Diversity
Fig. 18. Marasmius straminiceps (N. Wannathes 244). 1.
Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5. Pileipellis, Scale bar 1 = 10 mm, 2-5 =
10 µm
Fig. 19. Marasmius guyanensis (N. Wannathes 254). 1.
Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5. Pileipellis, Scale bar 1 = 5 mm, 2-5 =
10 µm
15. Marasmius guyanensis Mont., Ann. Sci.
Nat., Bot., sér. 4, 1:114. 1854.
(Fig. 19)
Pileus 0.5-4 mm diam, hemispherical to
convex, umbilicate, with or without small dark
reddish brown papilla in umbilicus, striate, dull,
dry, glabrous; disc brown (7E6-7), margin
brownish yellow to orange (6B7-8). Context
thin, yellowish orange (4A6). Lamellae adnate
to a collarium, distant (6-11) with no lamellula,
non-intervenose, narrow, yellowish orange
(4A6), non-marginate. Stipe 15-26 × 0.2-0.3
mm, central to eccentric, cylindrical, terete,
tough, glabrous, insititious, dark reddish brown
overall; rhizomorphs usually present. Odor and
taste not distinctive.
Basidiospores 10-12 (-15) × 3-4(-5) μm
[xmm = 11.6 ± 0.7 × 3.8 ± 0.3 μm, xmr = 11.012.4 × 3.4-4.2 μm, Qmm = 3.1 ± 0.1, Qmr = 3.03.3, n = 25 spores, s = 5 specimens], ellipsoid,
siccus-type broom cells; main body 10-21 × 611(-20) μm, cylindrical to pyriform, turbinate,
subglobose or sometimes irregular in outline,
rarely lobed, hyaline, inamyloid, thin-walled;
apical setulae 1-5 × 1 μm, cylindrical to conical
or irregular in outline, obtuse to subacute,
brownish yellow, inamyloid, thin- to thickwalled. Pileus trama interwoven, inamyloid.
Lamellar trama hyphae 3-7(-9) μm diam,
interwoven, cylindrical, smooth, hyaline,
inamyloid, thin-walled, non-gelatinous. Stipiti-
pellis hyphae 3-6 μm diam, parallel, cylindrical,
smooth, light brown to brown, inamyloid,
thick-walled (up to 2 μm), non-gelatinous. Stip
trama hyphae 2-5 μm diam, parallel,
cylindrical, smooth, hyaline to pale-yellow,
weakly dextrinoid to inamyloid, thin-walled.
Caulocystidia absent. Clamp connections
present in all tissues.
Habit, habitat, and known distribution:
Scattered to gregarious on undetermined
dicotyledonous leaves. Caribbean region,
Indonesia, South America, Singapore, Thailand.
Material examined: Thailand. Chiang Mai
Province, Mae Taeng District, “Water Conservation
Area” Highway 1,095 at 22 km marker, N. 19º 7.5’ E 98º
45.7’ alt. 724 m., 2 July 2003, N. Wannathes 015 (CMU,
SFSU); same location, 5 July 2004, Y. S. Tan 314 (CMU,
SFSU); Chiang Mai Province, Mushroom Research
Centre, 27 km marker on Hwy 1095, 28 July 2003, N.
Wannathes 046 (CMU); Chiang Mai Province, Chiang
Mai University, 11 Aug. 2004, N. Wannathes 254 (CMU,
SFSU); Chiang Mai Province, Mae Taeng District, Doi
Huai Kom Bang in Papae Village, 23 August 2004, N.
Wannathes 280 (CMU, SFSU).
Discussion: The material from Thailand
differs slightly from that described by
Desjardin et al. (2000) from Indonesia in
forming pilei with more brown tones and less
orange tones, and in forming basidiospores that
are slightly shorter (mean of 11.1 × 3.5 µm vs.
12.1 × 3.6 µm). In other details, the two
populations are similar.
237
A tetrapolar mating system was reported
for M. guyanensis by Wannathes et al. (2007)
based on Thai specimens.
16. Marasmius cafeyen Wannathes, Desjardin
& Lumyong, sp. nov.
(Fig. 20)
MycoBank: MB512414
Etymology: ‘cafeyen’ (Thai) = coffee with milk;
referring to the color of the pileus.
Pileus 2-8 mm diametro, hemisphaericus usque
convexus, umbilicatus, cum vel sine papilla atrobrunnea,
striatus usque sulcatus, minute velutinus, hebetatus,,
margine creameo-brunneo usque creameo-flavo, disco
brunneo cum papilla vel macula atrobrunnea ad centrum..
Contextus creameus, tenuis. Lamellae adnatae usque
collarium parvum, distantes (8-12), latae, flavo-albae,
haud marginatae, haud intervenosae. Stipes 6-31 × < 0.2
mm, centralis, cylindratus, filo metallico similis, glaber,
insititius, apice flavo-albo, basi atrobrunnea interdum
cum rhizomorphis. Odor saporque non propria.
Basidiosporae 9-11 × 4-5 µm, ellipsoidae, laeves,
hyalinae, inamyloidae, tenuitunicatae. Basidia non
observata. Basidiolae fusoideae usque clavatae.
Cheilocystidia abundantia, typi Sicci; 9-17 × 8-13 µm,
clavata usque late clavata vel pyriformia, hyalina,
inamyloidea, tenuitunicata; setulis apicalibus 2-6 × 1 µm,
cylindratis usque conicis, saepe furcatis, obtusis usque
subacutis, pallide flavis, crassetunicatis. Pleurocystidia
nulla. Pileipellis hymeniformis, leniter maculosus, typi
Sicci; 12-18(-25) × 7-14 µm, clavatus usque late clavatus,
pyriformis vel adumbratim inaequabilis, raro ramosus,
hyalinus usque pallide flavus, inamyloideus, tenui- usque
crassetunicatus; setulis apicalibus 2-3 × 1 µm, conicis
usque cylindratis, subacutis usque obtusis, flavobrunneis usque brunneis, crassetunicatis. Trama pilei
intertexta, inamyloidea. Trama lamellae regularis usque
intertexta, hyphis 4-7(-10) µm diametro, cylindratis
usque inflatis, laevibus, hyalinis, inamyloideis,
tenuitunicatis, haud gelatinosis. Stipitipellis subparallelus,
hyphis 3-5 µm diametro, cylindratis, brunneis usque
atrobrunneis, laevibus, dextrinoideis, crassetunicatis
(usque ad 1 µm), haud gelatinosis. Trama stipitis
subparallela, hyphis 2-5 µm diametro, cylindratis,
hyalinis, laevibus, inamyloideis, tenuitunicatis, haud
gelatinosis. Caulocystidia nulla. Fibulae praesentes in
omnibus texturis.
Typus: Chiang Mai Province, Mae Taeng District,
Pathummikaram temple in Pa Deng Village, dispersus
usque gregarius in folis plantae dicotyledoneae, 25
August 2003, N. Wannathes 130 (CMU: holotypus;
SFSU: isotypus).
Pileus 2-8 mm diam., hemispherical to
convex, umbilicate, with or without dark brown
papilla, striate to sulcate, minutely velutinous,
dull, margin creamish brown to creamish
yellow, disc brown with a dark brown papilla
or spot at the centre. Context cream, thin.
Lamellae adnate to a small collarium, distant
(8-12), broad, yellowish white, non-marginate,
238
non-intervenose. Stipe 6-31 × < 0.2 mm,
central, cylindrical, wiry, glabrous, insititious,
apex yellowish white, base dark brown;
rhizomorphs sometimes present. Odor and
taste not distinctive.
Basidiospores 9-11 × 4-5 µm [xmr = 9.410 × 4.2-4.4 μm, xmm = 9.8 ± 0.3 × 4.3 ± 0.1
μm, Qmr = 2.2-2.4, Qmm = 2.3 ± 0.1, n = 25
spores, s = 3 specimens], ellipsoid, smooth,
hyaline, inamyloid, thin-walled. Basidia not
observed. Basidioles fusoid to clavate.
Fig. 20. Marasmius cafeyen (N. Wannathes 130). 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pileipellis, Scale bar 1 = 13 mm, 2-4 = 10 µm
Cheilocystidia abundant, of Siccus-type broom
cells; main body 9-17 × 8-13 µm, clavate to
broadly clavate or pyriform, hyaline, inamyloid,
thin-walled; apical setulae 2-6 × 1 µm,
cylindrical to conical, often forked, obtuse to
subacute,
pale
yellow,
thick-walled.
Pleurocystidia absent. Pileipellis hymeniform,
weakly mottled, composed of Siccus-type
broom cells; main body 12-18(-25) × 7-14 µm,
clavate to broadly clavate, pyriform or irregular
in outline, seldom branched, hyaline to pale
yellow, inamyloid, thin- to thick-walled; apical
setulae 2-3 × 1 µm, conical to cylindrical,
subacute to obtuse, yellowish brown to brown,
thick-walled. Pileus trama interwoven,
inamyloid. Lamellar trama hyphae 4-7(-10)
µm diam., regular to interwoven, cylindrical to
inflated, smooth, hyaline, inamyloid, thinwalled, non-gelatinous. Stipitipellis hyphae 3-5
µm diam., subparallel, cylindrical, brown to
dark brown, smooth, dextrinoid, thick-walled
(up to 1 µm), non-gelatinous. Stipe trama
hyphae 2-5 µm diam., subparallel, cylindrical,
hyaline, smooth, inamyloid, thin-walled, nongelatinous. Caulocystidia absent. Clamp
connections present in all tissues.
Fungal Diversity
Habit, habitat and known distribution.
Scattered to gregarious on undetermined
dicotyledonous leaves, northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Sangasabhasri
Land on the way to Huai Kok Ma Village, N 18º 48.4’ E
98º 54.6’ alt. 1,146 m., 29 July 2003, N. Wannathes 058
(CMU, SFSU); Chiang Mai Province, Mae Taeng
District, Pathummikaram temple in Pa Deng Village, 25
August 2003, N. Wannathes 130 (CMU: holotype;
SFSU: isotype); Chiang Mai Province, Doi Inthanon
National Park, Junction of Highway 1,009 and road to
Mae Chaem, N 18º 31.6’ E 98º 29.6’ alt. 1,703 m., 3 July
2004, N. Wannathes 200 (CMU, SFSU).
Discussion: Marasmius cafeyen is
characterized by the following features: small,
striate to sulcate, umbilicate pilei colored
creamish brown to creamish yellow (coffee
with milk coloured) with a darker disc, distant
(8-12) collariate lamellae with, non-marginate
edges, ellipsoid basidiospores with mean 9.8 ×
4.3 μm and mean Q = 2.3, and dicotyledonous
leaf substrate. The Thai species is most
phenetically similar to the following species.
Marasmius gracilichorda Corner, described
from Singapore, differs in forming a brownish
orange to reddish brown or dark brown pileus,
brown marginate lamellae, and pileipellis
broom cells with warted and often forked
setulae up to 12 µm long (Corner, 1996).
Marasmius pallipes Singer, described from
South America, differs in forming more
cinnamon buff to ochraceus buff pileus, a pale
stipe (stramineus to light chestnut), and slightly
broader basidiospores measuring 4.5-6.2 µm
wide with mean diameter near 5 µm (Singer,
1976). Marasmius ruforotula Singer, described
from North America and reported here from
Thailand, differs in forming pilei with reddish
brown to rusty brown or brownish yellow disc
and greyish orange to light yellow margin,
orange-marginate lamellae, and broader
basidiospores with mean width 5.8 µm.
17. Marasmius ruforotula Singer, Sydowia 2:
34. 1948.
(Fig. 21)
Pileus 3-7 mm diam, paraboloid to
convex, umbilicate, with a small reddish brown
papilla in the umbilicus, striate, dull, dry,
glabrous to minutely subvelutinous; disc
brownish yellow to pale rusty brown, margin
grayish orange (6B4-5) to light yellow (4A4).
Context thin, cream (3A2). Lamellae adnate to
a collarium, distant (9-10) with no lamellula,
non-intervenose, broad (0.8-1 mm), cream
(3A2) with orange edges. Stipe 27-34 × 0.1-0.2
mm, central to eccentric, cylindrical, terete,
wiry, tough, glabrous, insititious, reddish
brown (7E6) overall; rhizomorphs absent. Odor
and taste not distinctive.
Fig. 21. Marasmius ruforotula (N. Wannathes 257). 1.
Basidiomes 2. Basidioles and basidium 3. Basidiospores
4. Cheilocystidia 5. Pileipellis, Scale bar 1 = 20 mm, 2-5
= 10 µm
Basidiospores (8-)10-12 × (4.5-)5-7 μm
[xmr = 9.3-11.1 × 5.1-6.5 μm, xmm = 10.2 ± 0.9
× 5.8 ± 0.7 μm, Q = 1.5-2.0, Qmr = 1.6-2.0, Qmm
= 1.8 ± 0.2, n = 25 spores, s = 1 specimen],
ellipsoid, smooth, hyaline, inamyloid, thinwalled. Basidia and basidioles clavate to
cylindrical. Cheilocystidia common, of Siccustype broom cells; main body 12-15 × 6-8 μm,
cylindrical to pyriform, clavate or broadly
clavate, rarely lobed, hyaline, thin-walled;
apical setulae 3-5 × 0.8-1.0 μm, crowded,
cylindrical to conical or wavy, obtuse to
subacute, yellow, inamyloid, thin-walled.
Pleurocystidia absent. Pileipellis hymeniform,
weakly mottled, composed of Siccus-type
broom cells; main body 14-20 × 8-14 μm,
pyriform to clavate or broadly clavate,
sometimes irregular in outline, seldom lobed,
pale yellow, inamyloid, thin- to thick-walled;
apical setulae (2-)4-8 × 1-1.5(-2) μm,
cylindrical to conical or irregular in outline,
239
obtuse to subacute, yellow to yellowish brown
or tawny, inamyloid, thick-walled. Pileus
trama and lamellar trama interwoven; hyphae
4-8 μm diam, cylindrical, smooth, hyaline,
inamyloid,
thin-walled,
non-gelatinous.
Stipitipellis hyphae 3-4 μm diam, parallel,
cylindrical, smooth, light brown, inamyloid,
thick-walled (up to 1 μm), non-gelatinous.
Stipe trama hyphae 2-3 μm diam, parallel,
cylindrical, smooth, hyaline, inamyloid, thinwalled.
Caulocystidia
absent.
Clamp
connections present in all tissues.
Habit, habitat, and known distribution:
Scattered to gregarious on undetermined
dicotyledonous leaves. Florida, Guadeloupe,
Mexico, Ecuador, Papua New Guinea, Java,
Thailand.
Material examined: Thailand. Chiang Mai
Province, New waterfall near 36 km marker on Hwy
1095, 20 Aug. 2004, N. Wannathes 257 (CMU, SFSU).
Discussion: The material from northern
Thailand fits within the concept of M.
ruforotula as it occurs in the old world tropics
as reported by Desjardin and Horak (1997) and
Desjardin et al. (2000) although there are
subtle differences. The Thai material has larger
basidiospores with mean 10.2 × 5.8 µm versus
8.9 × 4.3 mm from Indonesia. Spore size and
other basidiome features match quite closely
those reported for M. gordipes Sacc. & Paol., a
species described from Melaka State, Malaysia.
The latter species, however, was characterized
as having a reddish brown pileus and an
exceptionally long stipe, measuring 60-130 mm,
and with Siccus-type broom cells that have
fewer and more blunt setulae (Pegler 1986).
A tetrapolar mating system was reported
for M. ruforotula by Wannathes et al. (2007)
based on Thai specimens.
Section Globulares Kühner, Botaniste 25:
100. 1933 (ut Globularineae).
Type: Marasmius globularis Fr. (= M. wynneae
Berk. & Broome).
18. Marasmius calvus Berk. & Broome, J.
Linn. Soc. Bot. 14: 34. 1873
(Figs 22, 30-1)
Pileus 42-87 mm diam., convex to planoconvex, sulcate to plicate, glabrous, dull,
greyish orange (5B5) to light brown (6D5)
overall. Context brownish orange (5C5), thin.
Lamellae free to narrowly adnexed, subdistant
240
Fig. 22. Marasmius calvus (N. Wannathes 331). 1.
Basidiomes 2. Basidia 3. Basidiospores 4. Cheilocystidia
5. Pleurocystidia 6. Pileipellis, Scale bar 1 = 10 mm, 2-6
= 10 µm
to close (14-19) with 2-3 series of lamellulae,
broad (6-12 mm), brownish orange (5C5), nonmarginate, non-intervenose. Stipe 82-125 × 2-5
mm, central, cylindrical with subbulbous base,
hollow, glabrous to silky, non-insititious, light
brown to buff overall. Odor and taste not
distinctive.
Basidiospores 8-10 × 3.5-4 µm [xm = 8.9
± 0.6 × 3.9 ± 0.2 µm, Q = 2.0-2.7, Qm = 2.3, n
= 25 spores, s = 1 specimen], narrowly
ellipsoid, slightly curved in profile, smooth,
hyaline, inamyloid, thin-walled. Basidia 25-30
× 5 µm, cylindrical to clavate, 4-spored.
Basidioles fusoid to clavate. Cheilocystidia
abundant, 8-14 × 7-10 µm, broadly clavate to
pyriform or irregularly cylindrical, hyaline,
inamyloid,
thin-walled.
Pleurocystidia
abundant, also scattered along lamellae edge
among the cheilocystidia, 36-46 × 7-10 µm,
fusoid to clavate, usually attenuated and
strangulate at the apex, hyaline, inamyloid,
thin-walled. Pileipellis a hymeniform layer of
Globulares-type cells: 11-28 × 10-15 µm,
clavate to broadly clavate or pyriform, hyaline,
inamyloid, thin-walled. Pileus trama interwoven, dextrinoid. Lamellar trama hyphae 720 µm diam., regular, cylindrical to inflated,
smooth, hyaline, dextrinoid, thin-walled, nongelatinous. Stipitipellis hyphae 3-6 µm diam.,
subparallel, cylindrical, yellow, smooth, weakly
dextrinoid, thin-walled, non-gelatinous. Stipe
trama hyphae 6-9(-11) µm diam., parallel,
cylindrical, hyaline, smooth, dextrinoid, thinwalled, non-gelatinous. Caulocystidia absent.
Clamp connections present in all tissues.
Fungal Diversity
Habit, habitat and known distribution.
Scattered to gregarious on dicotyledonous
leaves, Indonesia, northern Thailand and Sri
Lanka.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mokfa
Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
m., 25 June 2005, N. Wannathes 331 (CMU, SFSU).
Discussion: Diagnostic features of
Marasmius calvus include a sulcate to plicate,
greyish orange to light brown pileus, subdistant
(14-19), broad lamellae, a pallid, glabrous stipe,
narrowly ellipsoid basidiospores with mean 8.9
× 3.9 µm, numerous cheilocystidia and
pleurocystidia, and an absence of caulocystidia.
The Thai specimen matches the material
described from Indonesia and Sri Lanka quite
well. Absent from the description of the
Indonesian specimen (Desjardin et al., 2000)
are the sterile lamellae edge composed of short,
broadly clavate cheilocystidia as reported
herein. Re-examination of Retnowati 145
(SFSU) confirmed their presence. Marasmius
calvus is the only known member of sect.
Globulares from Thailand that forms distinct
pleurocystidia.
19. Marasmius albimyceliosus Corner, Beih.
Nova. Hedwigia. 111: 28. 1996
(Figs 23, 30-2)
Pileus 15-38 mm diam., convex when
young, plano-convex to plano-concave and
subumbilicate in age, reticulate, glabrous, dull,
disc and striae brown, elsewhere pale cream to
grayish brown. Context light brownish cream,
thin. Lamellae adnate, subdistant (12-16) with
3-4 series of lamellulae, narrow (2-3 mm),
cream, non-marginate, strong intervenose to
poroid-reticulate. Stipe 23-53 × 1-4 mm,
central, cylindrical or tapering downward to a
subbulbous base, hollow, glabrous, noninsititious, apex light greyish brown, base
brown to reddish brown. Odor and taste not
distinctive.
Basidiospores 6-7(-8) × (3-)3.5-4(-4.5)
µm [xm = 6.8 ± 0.5 × 4 ± 0.3 µm, Q = 1.5 - 2.0,
Qm = 1.7, n = 25 spores, s = 1 specimen],
ellipsoid, smooth, hyaline, inamyloid, thinwalled. Basidia not observed. Basidioles
cylindrical. Cheilocystidia poorly differentiated,
Fig. 23. Marasmius albimyceliosus (N. Wannathes 422).
1. Basidiomes 2. Basidioles 3. Basidiospores 4.
Pileipellis, Scale bar 1 = 10 mm, 2-4 = 10 µm
basidiole-like, clavate cells. Pleurocystidia
absent. Pileipellis a hymeniform layer of
Globulares-type cells: 15-30 × 9-18 µm,
clavate to broadly clavate or pyriform, hyaline
to light brown, inamyloid, thin-walled. Pileus
trama interwoven, dextrinoid. Lamellar trama
hyphae 4-10 (-16) µm diam., regular to
interwoven, cylindrical to inflated, smooth,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Stipitipellis hyphae 4-8 µm diam., subparallel,
cylindrical, brown to dark brown, smooth,
inamyloid, thick-walled (up to 2 µm), nongelatinous. Stipe trama hyphae 7-12 µm diam.,
subparallel, cylindrical, pale yellow to yellow,
smooth, inamyloid, thin- to slightly thickwalled (up to 0.5 µm), non-gelatinous.
Caulocystidia absent. Clamp connections
present in all tissues.
Habit, habitat and known distribution:
Solitary on dicotyledonous leaves, northern
Thailand and Singapore.
Material examined: Thailand. Phrae Province,
Muang District, Pa Dang, Nong Cham Village, 18
August 2005, N. Wannathes 422 (CMU, SFSU).
Discussion: Marasmius albimyceliosus is
distinguished by a reticulate, pale cream to
greyish brown pileus with darker disc and
striae, subdistant, poroid-reticulate lamellae, a
glabrous, brown to reddish brown stipe, small
basidiospores with mean 6.8 × 4 µm, poorly
differentiated cheilocystidia, and an absence of
pleurocystidia and caulocystidia. The Thai
material matches quite well with the protologue
based on specimens collected from Singapore
(Corner, 1996). Marasmius albimyceliosus is
similar to M. cohortalis Berk. described from
South America, but the latter differs in forming
well differentiated, subglobose to pyriform
cheilocystidia.
241
20.
Marasmius pellucidus Berk. & Broome,
J. Linn. Soc., Bot. 14: 35. 1873.
(Figs 24,
30-3)
= Cantharellus elegans Berk. & Broome, J. Linn.
Soc., Bot. 14: 33. 1873.
= Marasmius papyraceus Massee, Bull. Misc.
Inform. 10: 358. 1914.
Pileus 15-50(-98) mm diam, obtusely
conical to convex or campanulate when young,
expanding to broadly convex, broadly campanulate, plano-campanulate or nearly plane in age,
often with a shallow central depression; disc
smooth to rugulose or reticulate; margin striate
to rugulose-striate or rugulose-sulcate, pellucid,
initially decurved, in age decurved to upturned,
wavy or undulate; surface hygropha-nous,
glabrous, moist to dry, dull; disc ivory, cream
(4A3), pale orange white (5A2-3), or pale
brownish grey (6C3); margin white, ivory, buff,
pale yellowish white (4A2) or cream (4A3), in
age entire pileus often dingy white and pellucid.
Context thin (0.5-1 mm), pliant, white.
Lamellae adnate to shallowly adnexed, noncollariate, close to subdistant or distant (1220(-26)) with 3-5 series of lamellulae, narrow
(0.5-2 mm, rarely up to 4 mm broad),
sometimes not reaching pileus margin, often
anastomosing and intervenose, white to ivory,
pale yellowish white (4A2) or cream (4A3);
edges even, non-marginate. Stipe (20-)40-112 ×
(1-)2-3 mm, central, terete or sometimes oncecleft, ±equal, sometimes wavy, cartilaginous or
pliant, tough, fistulose, sometimes twistedfibrous, dull, dry, apex minutely pruinose, base
pruinose or furfuraceous to appressed-fibrillose,
non-insititious, arising from dense creamy
white mycelium; apex white to pale yellowish
white (4A2), centrally greyish orange (5B4),
brownish yellow (5C5-6) or brownish orange
(6C5-7), base brown (7D-E5-8), reddish brown
(8D-E4-8), or dark brown (7-8F5-8). Odor
mild, somewhat sweet, like Marasmius oreades.
Taste not distinctive.
Basidiospores (6-)6.5-8(-8.5) × (2.5-)3-4
µm [xmr = 6.9-7.3 × 3-3.6 µm, xmm = 7.1 ± 0.1
× 3.4 ± 0.2 µm, Qmr = 1.7-2.3, Qmm = 2.1 ± 0.2,
n = 20-25 spores, s = 11 specimens], subfusoid
to ellipsoid or amygdaliform, with a prominent
hilar appendix, smooth, hyaline, inamyloid,
thin-walled. Basidia 16-30 × 4.5-7 µm, clavate,
4-spored. Basidioles clavate to subfusoid.
Cheilocystidia abundant, lamellar edge sterile,
242
Fig. 24. Marasmius pellucidus (N. Wannathes 022). 1.
Basidiomes 2. Basidia and basidioles 3. Basidiospores 4.
Cheilocystidia 5. Pileipellis 6. Caulocystidia, Scale bar 1
= 10 mm, 2-6 = 10 µm
12-30(-40) × 4.5-8(-12) µm, versiform, ranging
from irregularly cylindrical to fusoid,
ventricose, clavate or boot-shaped, hyaline,
inamyloid, thin-walled. Pleurocystidia absent.
Pileipellis a hymeniform layer of Globularestype cells: cells 10-25(-32) × (6-)8-18(-25) µm,
subglobose to pyriform, clavate or broadly
clavate, sometimes in short chains, hyaline,
inamyloid, thin-walled; arising from a subcutis
up to 30 µm thick formed of irregularly
isodiametric or puzzle-shaped cells 5-12 µm
diam that are tightly packed, dextrinoid, thinwalled. Pileus trama of loosely interwoven
hyphae 2.5-8(-12) µm diam, cylindrical to
irregularly cylindrical, thin-walled or thickwalled (up to 0.5 µm), hyaline, dextrinoid, nongelatinous. Lamellar trama regular; hyphae
2.5-10 µm diam, cylindrical, hyaline,
dextrinoid,
thin-walled,
non-gelatinous.
Stipitipellis hyphae 5-10 µm diam, parallel,
cylindrical, smooth, non-incrusted, yellow to
tawny in H2O, pale yellowish green in 3%KOH,
strongly dextrinoid, thick-walled (up to 0.5
µm), non-gelatinous. Stipe trama hyphae 516(-20) µm diam, subparallel, cylindrical,
hyaline, dextrinoid thin-walled. Caulocystidia
15-60(-85) × (4-)6-14 µm, versiform, ranging
from irregularly cylindrical to fusoid,
ventricose, clavate or irregular in outline,
sometimes geniculate, hyaline, inamyloid to
dextrinoid, thin-walled to thick-walled (0.5-1.5
µm). Clamp connections present in all tissues.
Habit, habitat and known distribution:
Gregarious to densely gregarious, typically in
Fungal Diversity
cespitose clusters on leaf mulch or on woody
debris or rotten logs of various dicotyledonous
plants, sometimes amongst bamboo leaves.
Borneo, Java, Malaysia, New Caledonia,
Singapore, Sri Lanka and Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Inthanon National Park, Mae Chem jct.,
elev. 1600 m, N18°31.420’, E98°29.216’, 26 June 2003,
N. Wannathes 007 (CMU, SFSU); Chiang Mai Province,
Pathummikaram temple on Hwy 1095, 13 Jul. 2003, N.
Wannathes 022 (CMU, SFSU); Chiang Mai Province,
Mushroom Research Centre, 27 km marker on Hwy
1095, 2 Jul. 2003, N. Wannathes 016 (CMU, SFSU);
Chiang Mai Province, Doi Suthep National Park, Sangra
Sabhasri Lane to Huai Kok Ma Village, elev. 1145 m,
N18°48.402’, E98°54.617’, 14 Aug. 2003, N. Wannathes
109 (CMU, SFSU); same location, 24 June 2005, N.
Wannathes 321 (CMU, SFSU); Chiang Mai Province,
Mae Taeng District, Mae Sae Village, Pang Sa Ded
Water Conservation Area, N 19º 14.6', E 98º 38.5' alt.
962 m., 26 june 2005, N. Wannathes 342 (CMU, SFSU);
Doi Inthanon National Park, Mae Chem jct., elev. 1600
m, N18°31.420’, E98°29.216’, 27 June 2005, N.
Wannathes 352 (CMU, SFSU).
Discussion: Marasmius pellucidus is
recognized easily in the field because of its
relatively large, paper-thin, white, pellucidstriate pileus, very narrow and intervenose
lamellae, and cespitose, reddish brown to
brown stipes. It forms large, dense clusters on
leaf mulch, woody debris or on rotten logs.
Microscopically, diagnostic features include: a
hymeniform pileipellis of subglobose to
broadly clavate, non-setulose cells; versiform,
non-setulose cheilocystidia and caulocystidia; a
lack of pleurocystidia; relatively small,
subfusoid basidiospores; and dextrinoid tissues.
Collectively, these characters indicate that M.
pellucidus belongs in sect. Globulares, where it
is allied with a number of species from Africa
and South America. A redescription of M.
pellucidus based on examination of the type
specimens of M. pellucidus, M. papyraceus and
C. elegans, plus numerous newly collected
specimens throughout Southeast Asia was
published by Wannathes et al. (2004).
21. Marasmius
mokfaensis
Wannathes,
Desjardin & Lumyong, Fungal Diversity 36:
160. 2009.
(Figs 25, 30-4)
MycoBank: MB511934
Pileus 30-90 mm diam., obtusely conical
when young, convex to campanulate in age,
dull, dry, glabrous, rugulose-plicate, hygrophanous, blackish blue (19F4-6) to purplish
grey (14F3-4) or dark brown to dark greyish
brown (7-9F3-5) overall when young,
becoming greyish magenta (14D3) to pale
brownish grey (8-9D-E3) on disc and plicae in
age with the ridges paler to pale greyish brown
(10E3) or pale violet brown. Context pale
greyish white to white, thin. Lamellae subfree
to adnexed, distant (13-18) with 0-1 series of
lamellulae, broad (4-10 mm), greyish white
(9B1) to grey (9C-D2), greyish magenta (14D3)
or brownish grey (8D2), non-marginate, nonintervenose. Stipe 85-180 × 3-7 mm, central,
cylindrical with a narrowly clavate base,
twisted-fibrous,
Fig. 25. Marasmius mokfaensis (D.E. Desjardin 7726). 1.
Basidiomes 2. Basidia 3. Cheilocystidia 4. Pileipellis 5.
Basidiospores, Scale bar 1 = 10 mm, 2-6 = 10 µm
hollow, glabrous, base tomentose, apex greyish
violet (17D4-6) to greyish white, base greyish
yellow (4B3-4) to brownish grey (6-8D2-3);
stipe context yellow in age. Odor faintly
raphanoid to rancid. Taste unpleasant.
Basidiospores 27-33 × 5-6 µm [xmr =
28.6-30.7 × 5.0-5.6 µm, xmm = 30.0 ± 1.0 × 5.3
± 0.3 µm, Q = 4.7-6.6, Qmm = 5.7 ± 0.2, Qmr =
5.5-5.9, n = 25 spores, s = 4 specimens],
clavate to subfusoid, often curved in profile,
hyaline, inamyloid, thin- walled. Basidia 41-45
× 11-12 µm, clavate, 4-spored. Basidioles
cylindrical to clavate. Cheilocystidia common,
15-33 × (3-)5-11(-17) µm, irregularly clavate
to ventricose, hyaline, inamyloid, thin-walled.
Pleurocystidia absent. Pileipellis a hymeniform
layer of Globulares-type cells, 11-31(-39) × 715 µm, clavate to broadly clavate or pyriform,
hyaline, inamyloid, thin-walled. Pileus trama
interwoven, dextrinoid. Lamellar trama hyphae
4-8 µm diam., interwoven, cylindrical to
inflated, smooth, hyaline, dextrinoid, thinwalled, non-gelatinous. Stipitipellis hyphae 5243
15 µm diam., parallel, cylindrical, smooth,
brown to dark greenish brown, inamyloid to
weakly dextrinoid, thin- to thick-walled (up to
1 µm), non-gelatinous. Stipe trama hyphae 3-8
µm diam., parallel, cylindrical, smooth, hyaline,
dextrinoid,
thin-walled,
non-gelatinous.
Caulocystidia absent. Clamp connections
present in all tissues.
Habit, habitat and known distribution:
Solitary on bamboo and dicotyledonous leaves,
Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mokfa
Waterfall, on Hwy 1095, N19° 6.581', E98°46.353',
1014 m alt., 28 June 2003, D.E. Desjardin 7592 (CMU,
SFSU); same location, 3 July 2003, D. E. Desjardin 7606
(CMU, SFSU); same location, 8 July 2003, N.
Wannathes 020 (CMU, SFSU); same location, 29 June
2004, D.E. Desjardin 7726 (CMU: holotype; SFSU:
isotype); same location, 25 June 2005, N. Wannathes
328 (CMU, SFSU).
Discussion: Diagnostic features of
Marasmius mokfaensis include: a very large,
plicate pileus 30-90 mm diam., colored
blackish blue to purplish grey with paler ridges;
distant, broad greyish brown lamellae; a large
(85-180 × 3-7 mm), pale colored stipe; clavate
basidiospores with mean 30 x 5.3 µm and mean
Q = 5.7; irregularly clavate to ventricose
cheilocystidia, and absence of pleurocystidia
and caulocystidia. This is the largest known
Marasmius in Thailand and is phenetically
similar to two large African species.
Marasmius zenkeri Henn. differs in forming
pilei with more lilac-violet to lilac-pink pilei,
smaller basidiospores (15.5-27 × 4.2-5.6 µm
with mean Q = 4.0), and more consistently
clavate
cheilocystidia
(Antonín
2007).
Marasmius bekolacongoli Beeli differs in
forming pilei that in age have dull red to
greyish red disc and sulcae with yellowish
white to lemon yellow ridges, smaller
basidiospores (17.5-26 × 3.8-5.4 µm with mean
Q = 4.8), and more consistently clavate
cheilocystidia (Antonín 2007).
22. Marasmius purpureostriatus Hongo, J.
Jap. Bot. 33: 344. 1958.
(Figs 26, 30-8)
Pileus 13-20 mm diam., obtusely conical
to convex with a small rugulose umbo, sulcate
to plicate, glabrous, dull, striped, disc dark
violet (18F8), sulcae greyish violet (18E6),
greyish yellow (2B4) elsewhere. Context
greyish violet (18E6), thin. Lamellae adnexed,
244
subdistant (9-13) with 0-1 series of lamellulae,
broad (1-3 mm), pale yellow (2A3), nonmarginate, non-intervenose. Stipe 52-103 ×
0.5-1.5 mm, central, cylindrical with or without
a subbulbous base, hollow, glabrous, noninsititious, apex purplish grey (16D2), base
brown to reddish brown (E8E). Odor and taste
not distinctive.
Fig. 26. Marasmius purpureostriatus (N. Wannathes
318). 1. Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5. Pileipellis, Scale bar 1 = 10 mm, 2-6 =
10 µm
Basidiospores (19-)21-30 × (4-)5-6(-7)
µm [xmr = 21.3-28.2 × 4.6-6.2 µm, xmm = 23.8
± 3.9 × 5.3 ±0.8 µm, Qmr = 3.6-6.2, Qmm = 4.7,
n = 25 spores, s = 8 specimens], clavate to
fusoid, often curved in profile, smooth, hyaline,
inamyloid, thin-walled. Basidia not observed.
Basidioles clavate to cylindrical. Cheilocystidia
abundant, 10-31 × 7-15 µm, cylindrical to
broadly clavate or pyriform, hyaline, inamyloid,
thin-walled. Pleurocystidia absent. Pileipellis a
hymeniform layer of Globulares-type cells, 1227 × 8-14 µm, broadly clavate to pyriform,
hyaline to light brown, inamyloid, thin- to
thick-walled (up to 0.5 µm). Pileus trama
interwoven, dextrinoid. Lamellar trama hyphae
3-10(-16) µm diam., interwoven, cylindrical to
inflated, smooth, hyaline, dextrinoid, thin- to
thick-walled (up to 1.5 µm), non-gelatinous.
Stipitipellis hyphae 4-11 µm diam., subparallel
to parallel, cylindrical, yellowish brown to light
brown, smooth, weakly dextrinoid to inamyloid,
thin- to thick-walled (up to 1µm), nongelatinous. Stipe trama hyphae 3-8(-12) µm
diam., subparallel, cylindrical, hyaline, smooth,
dextrinoid to weakly dextrinoid, thin-walled,
non-gelatinous. Caulocystidia absent. Clamp
connections present in all tissues.
Habit, habitat and known distribution:
Solitary to gregarious on dicotyledonous leaves,
Fungal Diversity
Japan, Malaysia, Thailand and Papua New
Guinea.
Material examined: Thailand. Chiang Mai
Province, Mae Taeng District, Tung Joaw Village, N 19º
8.07’ E 98º 38.9’ alt. 1423 m., 13 June 2004, N.
Wannathes 158 (CMU, SFSU), Chiang Mai Province,
Doi Suthep-Pui National Park, Medicinal Plant garden,
16 June 2005, N. Wannthes 317and 318 (CMU, SFSU),
Chiang Mai Province, Doi Inthanon National Park, At 25
km marker on Highway 1,009, N 18º 32.5’ E 98º 33.5’
alt. 1,076 m. 27 June 2005, N. Wannathes 363 (CMU,
SFSU).
Discussion: Marasmius purpureostriatus is
distinguished by a relatively large, plicate and
striped pileus with greyish violet disc and
plicae and greyish yellow ridges, subdistant,
broad lamellae, a purplish grey to reddish
brown stipe, large clavate basidiospores with
mean range of 21.3-28.2 × 4.6-6.2 µm, and an
absence of pleurocystidia and caulocystidia.
The Thai material matches quite closely with
the descriptions of specimens collected from
Papua New Guinea (Desjardin & Horak, 1997)
except that the Thai specimens formed smaller
basidiomes (pilei 13-20 mm diam. vs 15-50
mm diam.).
23. Marasmius
pseudopurpureostriatus
Wannathes, Desjardin & Lumyong, Fungal
Diversity 36: 162. 2009.
(Figs 27, 30-36)
MycoBank: MB511935
Pileus 14-38 mm broad, convex to
depressed, dull, glabrous, plicate, disc and
plicae dark purple (14F5) to dark magenta
(13F7), striped with pale yellow (3A3) to
greyish yellow (4B4) ridges. Context greyish
yellow (4B4), thin. Lamellae adnexed, distant
(9-11) with 0-1 series of lamellulae, narrow,
greyish yellow (4B4), non-marginate, nonintervenose. Stipe 62-80 × 1.5-3 mm, central,
cylindrical to tapering upwards, hollow,
glabrous, non-insititious, apex greyish magenta
(14E5), base brown (7E7). Odor and taste not
distinctive.
Basidiospores 20-25 × 5-6.2 µm [xm =
22.8 ± 2.4 × 5.6 ± 0.6, Q = 3.9-4.4, Qm = 4.1, n
= 4 spores, s = 1 specimen], clavate to
subfusoid, often curved in profile, hyaline,
inamyloid, thin-walled. Basidia not observed.
Basidioles cylindrical to clavate. Cheilocystidia
common, 12-25 × 6-12 µm, clavate to broadly
clavate, hyaline, inamyloid, thin-walled.
Fig. 27. Marasmius pseudopurpureostriatus (N.
Wannathes 286). 1. Basidiomes 2. Basidioles 3.
Pileipellis 4. Cheilocystidia 5. Basidiospores, Scale bar 1
= 10 mm, 2-6 = 10 µm
Pleurocystidia absent. Pileipellis a hymeniform
layer of Globulares-type cells, 14-23 × 10-11
µm, clavate to broadly clavate, hyaline,
inamyloid, thin- to thick-walled (up to 1 µm).
Pileus trama interwoven, dextrinoid. Lamellar
trama hyphae 4-10 µm diam., interwoven,
cylindrical to inflated, hyaline, smooth,
dextrinoid,
thin-walled,
non-gelatinous.
Stipitipellis hyphae (3-) 5-10 µm diam.,
subparallel, cylindrical, brown to light brown,
smooth, dextrinoid, thin- to thick-walled (up to
1 µm), non-gelatinous. Stipe trama hyphae 512 µm diam., subparallel, cylindrical, hyaline,
smooth, dextrinoid, thin-walled. Caulocystidia
absent. Clamp connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, Northern Thailand.
Material examined: Thailand, Chiang Rai
Province, Weing Papoa District, Khun Chae National
Park, N19°4.405', E99°23.543', 963 m alt., 10 June 2005,
N. Wannathes 286 (CMU: holotype; SFSU: isotype).
Discussion: Basidiomes of Marasmius
pseudopurpureostriatus look like a robust form
of M. purpureostriatus Hongo, and the two
species may be confused in the field. M.
purpureostriatus differs from the new species
in forming smaller pilei (13-20 mm diam.) with
darker violet disc and plicae and more whitish
ridges, has a thinner stipe (< 1.5 mm), and
slightly longer basidiospores (21-30 µm with
mean length about 24 µm). Marasmius
pseudopurpureostriatus is also similar to M.
bekolacongoli, a species widespread in tropical
Africa, but the latter differs in forming pilei
that are violaceous brown to dull red or greyish
red with lemon yellow stripes, and a much
larger stipe (50-150 x 2.5-10 mm).
245
24. Marasmius grandiviridis Wannathes,
Desjardin & Lumyong, Fungal Diversity 36:
156. 2009.
(Figs 28, 30-7)
MycoBank: MB511932
Pileus 37-88 mm diam., conical to
hemispherical when young, broadly conical
with a slight depression in age, glabrous, dull,
plicate, yellowish green with olive green plicae.
Context yellowish green, thin. Lamellae
narrowly adnate, distant (10-13) with 1 series
of lamellulae, broad (2-9 mm), yellowish green,
Fig. 28. Marasmius grandiviridis (N. Wannathes 152). 1.
Basidiomes 2. Basidioles 3. Cheilocystidia 4.
Basidiospores 5. Pileipellis, Scale bar 1 = 10 mm, 2-6 =
10 µm
non-marginate, non-intervenose. Stipe 133-180
× 3-7 mm, central, cylindrical, hollow,
glabrous, non-insititious, apex yellowish light
brown, base brownish green to brown. Odor and
taste not distinctive.
Basidiospores (23-)26-30 × 4-5 µm [xm =
26.7 ± 1.9 × 4.4 ± 0.5 µm, Q = 4.6-7.5, Qm =
6.1, n = 25 spores, s = 1 specimen], clavate to
subfusoid, often curve in profile, smooth,
hyaline, inamyloid, thin-walled. Basidia not
observed. Basidioles cylindrical to clavate.
Cheilocystidia abundant, lamellae edge sterile,
18-43 × 5-12 µm, irregularly cylindrical to
clavate, lageniform with a broad mucro, rarely
forked, hyaline, inamyloid, thin-walled.
Pleurocystidia absent. Pileipellis a hymeniform
layer of Globulares-type cells, 15-42 × 11-15
µm, clavate to broadly clavate or pyriform,
hyaline to pale yellow, inamyloid, thin-walled.
Pileus trama interwoven, dextrinoid. Lamellar
trama hyphae (3-) 5-10 µm diam., regular to
interwoven, cylindrical to inflated, smooth,
hyaline, weakly dextrinoid to inamyloid, thinwalled, non-gelatinous. Stipitipellis hyphae 4-8
(-10) µm diam., subparallel, cylindrical, yellow
to greenish yellow, smooth, weakly dextrinoid
246
to dextrinoid, thick-walled (up to 2 µm), nongelatinous. Stipe trama hyphae 4-12 µm diam.,
parallel, cylindrical, hyaline, smooth, dextrinoid, thin-walled, non-gelatinous. Caulocystidia absent. Clamp connections present in
all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves; Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Inthanon National Park, junction of Hwy
1009 and road to Mae Chem, N19°31.58', E 98°29.64',
1700 m alt.,10 June 2004, N. Wannathes 152 (CMU:
holotype; SFSU: isotype); same location, 27 June 2005,
N. Wannathes 349 (CMU, SFSU).
Discussion: Distinctive features of M.
grandiviridis include a large, plicate pileus (3788 mm diam) colored yellowish green with
darker olive-green plicae, yellowish green,
distant (10-13) lamellae, a large (133-180 × 3-7
mm), yellow to brownish green, glabrous stipe,
clavate basidiospores with mean 26.7 × 4.4 µm,
irregularly clavate to lageniform-mucronate
cheilocystidia, and an absence of pleurocystidia
and caulocystidia. It is similar to M. viridis
Desjardin & E. Horak, described from Papua
New Guinea, but the latter differs in forming
smaller basidiomes (pilei 10-15 mm diam, stipe
30-50 × 1 mm) with more lamellae (12-16), a
dark reddish brown stipe, smaller basidiospores
(20-25 × 4-5 µm) and cheilocystidia that are
more regularly clavate (Desjardin & Horak
1997).
25. Marasmius
laticlavatus
Wannathes,
Desjardin & Lumyong, Fungal Diversity 36:
158. 2009.
(Figs 29, 30-5)
MycoBank: MB511933
Pileus 15-19(-33) mm diam., convex to
plano-convex or plano-infundibuliform with a
wrinkled umbo, dull, glabrous, sulcate to
plicate, disc brown, margin yellowish grey to
greyish cream. Context greyish yellow, thin.
Lamellae adnexed to adnate, distant (10-13)
with 1-2 series of lamellulae, broad (2-6 mm),
buff to cream or white, non-marginate, nonintervenose. Stipe 30-80 × 1-2 mm, central,
cylindrical with a narrow subbulbous at base,
hollow, glabrous, apex yellowish grey to buff,
base brownish orange to reddish brown. Odor
and taste not distinctive.
Basidiospores 26-35 × 5-6(-7) µm [xmr =
28.7-32 × 5.1-6 µm, xmm = 30.4 ± 1.9 × 5.6 ±
Fungal Diversity
0.5 µm, Q = 4.1-6.9, Qmr = 5.0-6.0, Qmm = 5.5
± 0.5, n = 25 spores, s = 4 specimens], clavate
to subfusoid, often curved in profile, hyaline,
inamyloid, thin-walled. Basidia not observed.
Fig. 29. Marasmius laticlavatus (N. Wannathes 231). 1.
Basidiomes 2. Basidioles 3. Cheilocystidia 4. Pileipellis
5. Basidiospores, Scale bar 1 = 10 mm, 2-6 = 10 µm
Basidioles cylindrical to clavate. Cheilocystidia
abundant, 13-30 × 9-16 µm, clavate to broadly
clavate, hyaline, inamyloid, thin-walled.
Pleurocystidia absent. Pileipellis a hymeniform
layer of Globulares-type cells, 15-33 × 9-17(23) µm, broadly clavate to pyriform, hyaline,
inamyloid,
thin-walled.
Pileus
trama
interwoven, dextrinoid. Lamellar trama hyphae
3-10 µm diam., interwoven, cylindrical,
smooth, hyaline, weakly dextrinoid to
dextrinoid, thin-walled, non-gelatinous. Stipitipellis hyphae 3-8 µm diam., subparallel,
cylindrical, smooth, yellow to light brown,
dextrinoid, thick-walled (up to 2 µm), nongelatinous. Stipe trama hyphae 5-10 µm diam.,
parallel, cylindrical, smooth, hyaline, weakly
dextrinoid, thin-walled, non-gelatinous. Caulocystidia absent. Clamp connections present in
all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on bamboo or
dicotyledonous leaves, Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, 22 km marker on Hwy 1095, N19°07.57',
E98°45.65', 750 m alt., 5 July 2004, T. Y. Shin 312
(CMU, SFSU); Chiang Mai Province, Mae Rim District,
Huai Tung Taow Reservoir, 23 July 2004, N.Wannathes
231(CMU: holotype; SFSU: isotype); Chiang Rai
Province, Muang District, Pong Prabath Waterfall, 11
June 2005, N.Wannathes 293(CMU, SFSU); Phrae
Province, Muang District, Cherng Thong Waterfall, 16
August 2005, N. Wannathes 412 (CMU, SFSU).
Discussion: The new Thai species is
characterized by the following features:
relatively small, plicate pilei with brown disc
and pallid greyish cream margin; distant,
cream-colored lamellae; a thin, glabrous stipe
colored yellowish grey on the apex and reddish
brown at the base; large clavate basidiospores
with mean 30.4 × 5.6 µm and mean Q = 5.5;
broadly clavate cheilocystidia; and an absence
of pleurocystidia and caulocystidia. Marasmius
laticlavatus is most phenetically similar to
several pale-colored African species that also
lack pleurocystidia, as recently documented by
Antonín (2007) Marasmius camerunensis
Antonín & Mossebo differs in forming much
larger pilei (40-70 µm diam.) with violaceous
brown disc, broader lamellae (8-10 mm), a
much thicker stipe (4-6 mm), smaller
basidiospores (21-30 × 5.5-6.5 µm with mean
Q = 4.2), and growth on wood. Marasmius
tshopoensis Antonín differs in forming larger
pilei (up to 60 mm diam.), strongly intervenose
lamellae, smaller basidiospores (19-26 × 4.8-6
µm with mean Q = 4.3), and variably-shaped
cheilocystidia some of which may be
irregularly lobed, but not consistently broadly
clavate. Marasmius brunneolus (Beeli) Singer
differs in forming larger pilei (30-80 mm diam.)
that are more evenly pigmented brown to
reddish brown or striped white, a larger stipe
(70-180 × 3-6 mm), and smaller basidiospores
(15.5-25.5 × 3.8-5.4 µm with mean Q = 4.5).
Section Neosessiles Singer, Mycologia 50:
104. 1958.
Type: Marasmius neosessilis Singer.
26. Marasmius tenuissimus (Jungh.) Singer,
Fl. Neotrop. Monogr. 17: 258. 1976.
(Figs 31, 38-1)
≡ 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. &
Br.) Petch apud Petch, in & Bisby, Peradeniya Manual 6:
59. 1950.
247
Fig. 30. Basidiomata of Marasmius section Globulares. 1. M. calvus (N. Wannathes 331) 2. M. albimyceliosus (N.
Wannathes 422) 3. M. pellucidus (N. Wannathes 321) 4. M. mokfaensis (D.E. Desjardin 7726) 5. M. laticlavatus (N.
Wannathes 412) 6. M. pseudopurpureostriatus (N. Wannathes 286) 7. M. grandiviridis (N. Wannathes 152) 8. M.
purpureostriatus (N. Wannathes 318), Scale bars = 20 mm
248
Fungal Diversity
Fig. 31. Marasmius tenuissimus (N. Wannathes 199). 1.
Basidiomes 2. Basidium 3. Basidiospores 4.
Cheilocystidia 5. Pileipellis 6. Caulocystidia, Scale bar 1
= 20 mm, 2-6 = 10 µm
Pileus 7-22 mm diam., convex to planoconvex, striate to sulcate, often subreticulate,
glabrous, dull to shiny, greyish orange (6B5) to
golden brown (6C5) or light brown (6D5).
Context cream, thin. Lamellae adnate, remote
(5-9) with 0-1 series of lamellulae, narrow,
reticulate, intervenose, cream, non-marginate.
Stipe 0.5-4 × 0.3-1 mm, eccentric to sublateral,
cylindrical to tapering downwards, hollow,
pubescent to velutinous, insititious, dark brown
(7F4) overall. Odor and taste not distinctive.
Basidiospores (7-)9-10(-12) × 4-6(-7) µm
[xmr = 8.1-10.1 × 4.5-6.1 µm, xmm = 9.4 ± 0.8 ×
5.3 ± 0.6 µm, Qmr = 1.7-1.9, Qmm = 1.8 ± 0.1, n
= 25 spores, s = 5 specimens], ellipsoid,
smooth, hyaline, inamyloid, thin-walled.
Basidia 22-40 × 8-11 µm, cylindrical to clavate,
4-spored. Basidioles cylindrical to clavate.
Cheilocystidia of Siccus-type broom cells;
main body 11-25 × 5-14 µm, cylindrical to
clavate or pyriform, hyaline, inamyloid, thinwalled; apical setulae 3-8 × 1-2 µm, conical to
cylindrical, obtuse to subacute, hyaline, thinwalled. Pleurocystidia absent. Pileipellis
hymeniform, mottled, composed of Siccus-type
broom cells; main body 13-28(-35) × 7-11 µm,
clavate to broadly clavate or pyriform, usually
forked at apex, yellow to hyaline, inamyloid,
thin- to thick-walled; apical setulae 3-7 × 1-1.5
µm, conical to cylindrical, obtuse to subacute,
yellow to light brown, thick-walled. Pileus
trama interwoven, inamyloid. Lamellar trama
hyphae 3-7 µm diam., interwoven, cylindrical
to inflated, smooth, hyaline, inamyloid, thinwalled, non-gelatinous. Stipitipellis hyphae 4-9
µm diam., subparallel, cylindrical, brown to
yellowish brown, smooth, inamyloid, thin- to
thick-walled (up to 2 µm), non-gelatinous.
Stipe trama hyphae 3-6(-8) µm diam.,
subparallel, cylindrical, hyaline to pale yellow,
smooth, dextrinoid, thick-walled (up to 2 µm),
non-gelatinous. Caulocystidia common, 27-100
× 5-12 µm, cylindrical to clavate or irregular in
outline, often apically branched, hyaline to pale
yellow, inamyloid, thin- to thick-walled. Clamp
connections present in all tissues.
Habit, habitat and known distribution:
Gregarious to densely gregarious, typically in
cespitose clusters on wood, Malaysia, Northern
Thailand and South America (the latter
reported by Singer, 1976).
Material examined: Thailand. Chiang Mai
Province, Mushroom Research Centre, 27 km marker on
Hwy 1095, 28 July 2003, N. Wannathes 050 (CMU,
SFSU); Chiang Mai Province, Mae Taeng District, Tung
Joaw Village, N 19º 8.07’ E 98º 38.9’ alt. 1423 m., 31
July 2003, N. Wannathes 065 (CMU, SFSU); same
location, 30 June 2004, N. Wannathes 192 (CMU,
SFSU); Chiang Mai Province, Mae Taeng District, New
Waterfall, on 36 km. marker of Hwy1095, 2 July 2004,
N. Wannathes 199 (CMU, SFSU); same location, 20
August 2004, N. Wannathes 261 (CMU, SFSU).
Discussion: Marasmius tenuissimus is
characterized by a greyish orange to golden
brown or light brown, plicate-reticulate pileus,
remote, reticulate and intervenose lamellae, a
short, eccentric to sublateral stipe, small
basidiospores with mean 9.4 × 5.3 µm, and
growth on woody debris. Interestingly, the ITS
data indicate that M. tenuissimus is sister to M.
leveilleanus, the only known species in sect.
Leveilleani, and surprisingly distant from
members of sect. Sicci where its morphology
suggests it should be allied.
249
Section Leveilleani, Singer, Bull. Jard. Bot.
Bruxelles 34: 362: 1964.
Type: Marasmius leveilleanus (Berk.) Pat.
27. Marasmius leveilleanus (Berk.) Pat., Bull.
Soc. Mycol. Fr. 33: 35. 1917.
(Figs 32, 38-2)
= Heliomyces leveilleanus Berk., Lond. J. Bot. 6:
490. 1847.
= Marasmius umbraculum Berk. & Broome, J.
Linn. Soc. Bot. 14: 36. 1873.
Pileus 14-43 mm diam., hemispherical to
convex when young, convex to depressed in
age, umbonate, striate to sulcate, glabrous, dull
to shiny, disc dark brown (7F8) to brown (6E78), margin brown (7E8, 6D-E7) to brownish
orange (6D6). Context cream, thin. Lamellae
free, subdistant to close (17-25) with 0-1 series
of lamellulae, broad (up to 5 mm), cream, nonmarginate, non-intervenose. Stipe 22-90 × 0.51 mm, central, cylindrical, wiry, tough, hollow,
glabrous, insititious, brown to dark brown
overall; rhizomorphs sometimes present. Odor
and taste not distinctive.
Basidiospores (8-)9-11(-12) × (3-)4(-5)
µm [xmr = 9.0-10.7 × 3.9-4.7 µm, xmm = 9.6 ±
0.7 × 4.2 ± 0.3 µm, Qmr = 2.1-2.6, Qmm = 2.3 ±
0.2, n = 25 spores, s = 6 specimens], ellipsoid,
smooth, hyaline, inamyloid, thin-walled.
Basidia 24-28 × 6-9 µm, cylindrical to clavate,
4-spored. Basidioles cylindrical to clavate.
Cheilocystidia of Siccus-type broom cells;
main body 12-28(-37) × 4-15 µm, cylindrical to
clavate, hyaline, inamyloid, thin-walled; apical
setulae 1-5 × 1-2(-3) µm, conical to cylindrical,
obtuse, hyaline, thin- to thick-walled.
Pleurocystidia absent. Pileipellis hymeniform,
mottled, composed of Siccus-type broom cells;
main body 9-24(-36) × 5-11 µm, clavate to
broadly clavate or pyriform, yellow to hyaline,
inamyloid, thin- to thick walled; apical setulae
3-6 × 1.5-3 µm, conical to cylindrical, obtuse,
brown to light brown, thick-walled. Pileus
trama interwoven, inamyloid. Lamellar trama
hyphae (3-)5-12(-18) µm diam., interwoven,
cylindrical to inflated, smooth, hyaline,
inamyloid, thin-walled, non-gelatinous. Stipitipellis hyphae 3-8(-10) µm diam., subparallel,
cylindrical, brown to dark brown, smooth,
weakly dextrinoid, thin- to thick- walled, nongelatinous. Stipe trama hyphae (3-)5-8(-10) µm
diam., subparallel, cylindrical, hyaline to pale
yellow, smooth, weakly dextrinoid, thin-walled,
250
Fig. 32. Marasmius leveilleanus (N. Wannathes 268). 1.
Basidiomes 2. Basidia 3. Basidiospores 4. Cheilocystidia
5. Pileipellis, Scale bar 1 = 20 mm, 2-5 = 10 µm
non-gelatinous. Caulocystidia absent. Clamp
connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on wood, Africa,
Mexico, Malaysia and Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mokfa
Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
m., 28 August 2003, N. Wannathes 137 (CMU, SFSU);
same location, 29 June 2004, Y. S. Tan 276 (CMU,
SFSU); Chiang Mai Province, Chiang Mai University, 5
August 2004, N. Wannathes 248 (CMU, SFSU); Chiang
Mai Province, Mae Taeng District, New Waterfall, on 36
km. marker of Hwy1095, 20 August 2004, N. Wannathes
264 (CMU, SFSU); same location, 30 June 2005, N.
Wannathes 381 (CMU, SFSU); Chiang Mai Province,
Doi Suthep-Pui National Park, Mokfa Waterfall, on Hwy
1095, N19º 6.5’ E 98º 46.3’ alt. 1014 m., 21 August
2004, N. Wannathes 268 (CMU, SFSU); Chiang Rai
Province, Muang District, Khun Con Waterfall, 12 June
2005, N. Wannathes 308, (CMU, SFSU).
Discussion: Marasmius leveilleanus is
distinguished by a relatively large sulcate
pileus colored brown to brownish orange, free,
non-collariate lamellae, a wiry, tough, dark
brown, insititious stipe, Siccus-type broom
cells, moderately sized basidiospores with
mean 9.6 × 4.2 µm, non-dextrinoid tissues, and
lignicolous habit. Macromorphological features
suggest that the species is allied with members
of sect. Marasmius (e.g., wiry, insititious stipe)
although the lamellae are free and not collariate,
Fungal Diversity
while micromorphological features suggest that
it is allied with members of sect. Sicci (e.g.,
Siccus-type broom cells). Because of the
unusual combination of features and especially
the free lamellae and inamyloid tissues, Singer
(1964) erected the monotypic sect. Leveilleani
to accommodate the species. The ITS data
presented here support its distinction from
members of both sects. Marasmius and Sicci,
and suggest that it is more closely allied with
M. tenuissimus of sect. Neosessiles. It should
be noted that M. leveilleanus is phenetically
very similar to M. nodulocystis Pegler,
described from Africa. A comparison of the
descriptions of both species as reported by
Pegler (1977) and Antonín (2007) indicate that
they differ primarily in M. nodulocystis having
dextrinoid tissues, and M. leveilleanus having
an insititious stipe base and sometimes
rhizomorphs. The stipe basal attachment of M.
nodulocystis was not described although it has
been interpreted as non-insititious because the
species is accepted in sect. Sicci.
Section Sicci Singer, Mycologia 50: 106. 1958.
Type: Marasmius siccus (Schwein.) Fr.
Subsect. Siccini, ser. Spinulosi (Clémençon)
Desjardin, in Antonín & Noordeloos, Lib.
Bot. 8: 179. 1993.
Type: Marasmius cohaerens (Pers.)
Cooke & Quél. [Bas. Agaricus cohaerens
Pers.].
= series Actinopus Singer, pro parte, Fl. Neotrop.
Monogr. 17: 236. 1976.
28. Marasmius
coarctatus
Desjardin & Lumyong, sp. nov.
Wannathes,
(Figs 33, 38-3)
MycoBank: MB512417
Etymology: ‘coarctatus’ (L) = close-set, crowded;
referring to the lamellae spacing.
Pileus 10-27 mm diametro, convexus usque
plano-convexus, saepe subumbonatus, laevis usque
pellucido-striatus, velutinus, hebetatus, brunneus cum
perpaulo coloris olivacei usque brunneo-ravus vel
brunneo-aurantiacus. Contextus laete flavo-brunneus,
tenuis. Lamellae anguste adnatae usque annexae,
coarctatae (33-36) cum 3-5 seriebus lamellularum,
angustae, obcure flavae, haud marginatae vel
brunneomarginatae, haud intervenosae. Stipes 41-60 × 12 mm, centralis, cylindratus, cavus, hispidulus, haud
insititius, apice pallide flavo, basi atrobrunnea. Odor
saporque non propria. Basidiosporae 6-8(-10) × 2.5-3.5
µm, ellipsoideae, laeves, hyalinae, inamyloideae,
tenuitunicatae. Basidia non observata. Basidiolae
clavatae. Cheilocystidia abundantia, 2 typorum: a)
cellulis dispersis typi Sicci 12-23 × 5-7(-10)µm,
cylindratis usque clavatis, hyalinis, inamyloideis,
tenuitunicatis; setulis apicalibus 5-15 × 1-2 µm,
cylindratis, obtusis uque subacutis, hyalinis usque pallide
flavis, tenui- usque crassetunicatis; b) cheilosetis 25-57 ×
4-6 µm, fusoideis usque lanceolatis, subacutis usque
acutis,
hyalinis,
inamyloideis,
tenuiusque
crassetunicatis. Pleurosetae nullae vel rarae, 30-60 × 4-6
µm, fusoideae usque lanceolatae, subacutae usque acutae,
hyalinae, inamyloideae, tenui- usque crassetunicatae.
Pileipellis hymeniformis, leniter maculosus, 3 typorum:
a) cellulis typi Sicci 11-20 × 6-10 µm, cylindratis usque
clavatis, flavis, inamyloideis, tenui- usque crassetunicatis;
setulis apicalibus 3-7(15) × 1-2 µm, coarctatis,
cylindratis usque conicis, obtusis usque subacutis,
brunneis, tenuitunicatis; b) pileosetis 34-60 × 5-8 µm,
vulgaribus, dispersis, lanceolatis usque fusoideis,
interdum lobatis, subacutis usque acutis, hyalinis usque
pallide flavo-brunneis, inamyloideis, crassetunicatis; c)
intermediis inter cellulas scopiformes et setas 24-39 ×711 µm; paucis setularum apicalium 7-37 × 2-4 µm,
crassetunicatis. Trama pilei intertexta, dextrinoidea.
Trama lamellae regularis usque intertexta, hyphis 4-14(20) µm diametro, cylindratis, laevibus, hyalinis,
dextrinoideis,
tenuitunicatis,
haud
gelatinosis.
Stipitipellis subparallelus, hyphis 6-10(-20) µm diametro,
cylindratis, flavis usque laete brunneis, laevibus,
dextrinoideis, tenui- usque crassetunicatis (usque ad 1
µm), haud gelatinosis. Trama stipitis parallela, hyphis
(8-)14-18 µm diametro, cylindratis, hyalinis, laevibus,
dextrinoideis,
tenuitunicatis,
haud
gelatinosis.
Caulocystidia 2 typorum: a) cellulis typi Sicci 6-11 × 5-7
µm, cylindratis vel adumbratim inaequabilibus, flavis,
inamyloideis, tenuitunicatis; 3-5 setulis apicalibus, 5-26
× 1-3 µm, cylindratis, subacutis usque acutis, flavis
usque laete brunneis, tenuitunicatis; b) caulosetis 38-68
× 6-9 µm, lanceolatis usque fusoideis, raro furcatis,
flavis, inamyloideis, crassetunicatis. Fibulae praesentes
in omnibus texturis.
Typus: Chiang Mai Province, Doi Suthep-Pui
National Park, trail opposite with the way to Huai Kok
Ma Village, solitarius in folis plantae dicotyledoneae vel
ligno, 2 July 2005 N. Wannathes 385 (CMU: holotypus;
SFSU: isotypus).
Pileus 10-27 mm diam., convex to planoconvex, often subumbonate, smooth or
pellucid-striate, velutinous, dull, brown with a
hint of olive to brownish beige (6F3) or
brownish orange (5C5). Context light yellowish
brown (5D4), thin. Lamellae narrowly adnate
to adnexed, crowded (33-36) with 3-5 series of
lamellulae, narrow, dull yellow (3B3) with or
without brown edges, non-intervenose. Stipe
41-60 × 1-2 mm, central, cylindrical, hollow,
hispidulous, non-insititious, apex pale yellow
251
Fig. 33. Marasmius coarctatus (N. Wannathes 385). 1.
Basidiomes 2. Basidiole 3. Basidiospores 4a. Siccus-type
cheilocystidia 4b. Cheilosetae 5a. Siccus-type pileipellis
5b. Cells transitional between broom cells and setae 5c.
5c. Pileosetae 6a. Siccus-type caulocystidia 6b.
Caulosetae, Scale bar 1 = 10 mm, 2-6 = 10 µm
(4A3), base dark brown (9F6). Odor and taste
not distinctive.
Basidiospores 6-8(-10) × 2.5-3.5 µm
[xmr = 6.4-6.8 × 3.0 µm, xmm = 6.6 ± 0.4 × 3.0 ±
0.2 µm, Qmr = 2.1, Qmm = 2.1, n = 15-25 spores,
s = 2 specimens], ellipsoid, smooth,
hyaline,inamyloid, thin-walled. Basidia not
observed. Basidioles clavate. Cheilocystidia
abundant, composed of 2 types of cells: a)
scattered Siccus-type broom cells with main
body 12-23 × 5-7(-10) µm, cylindrical to
clavate, hyaline, inamyloid, thin-walled; apical
setulae 5-15 × 1-2 µm, cylindrical, obtuse to
subacute, hyaline to pale-yellow, thin- to thickwalled; b) cheilosetae 25-57 × 4-6 µm, fusoid
to lanceolate, subacute to acute, hyaline,
inamyloid, thin- to thick-walled. Pleurosetae
absent or seldom present, 30-60 × 4-6 µm,
fusoid to lanceolate, subacute to acute, hyaline,
inamyloid, thin- to thick-walled. Pileipellis
hymeniform, weakly mottled, composed of 3
types of cells: a) Siccus-type broom cells with
main body 11-20 × 6-10 µm, cylindrical to
clavate, yellow, inamyloid, thin- to thickwalled; apical setulae 3-7(15) × 1-2 µm,
crowded, cylindrical to conical, obtuse to
subacute, brown, thick-walled; b) pileosetae,
common, scattered, 34-60 × 5-8 µm, lanceolate
252
to fusoid, sometime lobed, subacute to acute,
hyaline to pale yellowish brown, inamyloid,
thick-walled; c) cells transitional between
broom cells and setae, 24-39 ×7-11 µm, with
few apical setulae 7-37 × 2-4 µm, thick-walled.
Pileus trama interwoven, dextrinoid. Lamellar
trama hyphae 4-14(-20) µm diam., regular to
interwoven, cylindrical, smooth, hyaline,
dextrinoid, thin-walled, non-gelatinous. Stipitipellis hyphae 6-10(-20) µm diam., subparallel,
cylindrical, yellow to light brown, smooth,
dextrinoid, thin- to thick-walled (up to 1 µm),
non-gelatinous. Stipe trama hyphae (8-)14-18
µm diam., parallel, cylindrical, hyaline, smooth,
dextrinoid, thin-walled, non-gelatinous. Caulocystidia composed of 2 types of cells: a)
Siccus-type broom cells with main body 6-11 ×
5-7 µm, cylindrical or irregular in outline,
yellow, inamyloid, thin-walled; apical setulae
5-26 × 1-3 µm, ranging from 3-5 setulae per
cell, cylindrical, subacute to acute, yellow to
light brown, thin-walled; b) caulosetae 38-68 ×
6-9 µm, lanceolate to fusoid, seldom forked,
yellow, inamyloid, thick-walled. Clamp
connections present in all tissues.
Habit, habitat and known distribution:
Solitary on dicotyledonous leaves or on wood,
Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National ParkMedicianl plant
garden, 16 June 2005, N. Wannathes 315 (CMU, SFSU);
Chiang Mai Province, Doi Suthep-Pui National Park,
trail opposite with the way to Huai Kok Ma Village, 2
July 2005 N. Wannathes 385 (CMU: holotype; SFSU:
isotype).
Discussion: Marasmius coarctatus is
distiguished by the following features: a
medium size pileus that is smooth or pellucidstriate and coloured brown with a hint of olive;
crowded (33-36), narrow, dull yellow lamellae;
a hispidulous stipe; small basidiospores with
mean 6.6 × 3.0 µm; a lack of pleurocystidia
and pleurosetae (rarely present); and presence
of distinctive pileosetae, cheilosetae and
caulosetae. The new Thai species is similar to
M. cohaerens var. lachnophyllus (Berk.)
Gilliam described from North America, but the
latter species differs in forming larger pilei (1060 mm diam.) that are dark reddish brown,
broader (1-2 mm) lamellae with reddish brown
edges, slightly larger basidiospores (6.5-9 ×
3.2-4.8 µm, xm = 7.6 × 4.1 µm), and brownish
orange to reddish brown, dextrinoid setae up to
Fungal Diversity
20 µm diam. (Desjardin, 1989). Marasmius
coarctatus is also similar to M. coklatus
Desjardin, Retn. & E. Horak, described from
Indonesia, but the latter differs in forming
larger (15-60 mm diam.) chocolate brown pilei,
remote to distant (10-15) lamellae 4-12 mm
broad, and larger basidiospores in the range 1011 × 4.5-6 µm with mean 10.3 × 5.3 µm. Other
micromorphological features are similar except
for occasional dextrinoid setae; they are all
inamyloid in the Thai species (Desjardin et al.,
2000). Two additional phenetically similar
species are M. nexus Desjardin & E. Horak and
M. acanthocheilus Desjardin & E. Horak, both
described from Papua New Guinea (Desjardin
and Horak, 1997), Marasmius nexus differs in
forming Globulares-type cell in the pileipellis,
broader hymenial setae (up to 13 µm), and
grows on wood. Marasmius acanthocheilus
differs in forming a rusty orange pileus, larger
basidiospores (9-10.5 × 4-4.5 µm), and grows
on grass or bamboo leaves.
29. Marasmius trichotus Corner, Beih. Nova
Hedwigia 111: 102. 1996.
(Figs 34, 38-4)
Pileus 6-23 mm diam., broadly conical to
convex when young, plano-convex with or
without an umbo in age, smooth (non-striate),
hispidulous with erect golden hairs, dull, pale
orange (5A5) to orange (6A-B7-8), golden
yellow (5B7-8) or pale brownish orange (5C45), sometimes orangish white (4-5A3) at the
margin. Context yellowish white, thin.
Lamellae adnexed to narrowly adnate,
subdistant to close (12-22) with 2-4 series of
lamellulae, narrow, yellowish white, nonmarginate, non-intervenose. Stipe 15-90 × 0.51 mm, central, cylindrical, hollow, hispid with
brown hairs overall, non-insititious, apex
yellowish white (4-5A2) to pale orange (5A35), base brownish orange (6C5-6), rarely to
brown (7E7-8) or reddish brown (8E-F8). Odor
and taste not distinctive.
Basidiospores (10-)12-15(-16) × 3-5 µm
[xmr = 10.6-14 × 3.7-4.6 µm, xmm = 12.1 ± 0.8
× 4.1 ± 0.16 µm, Qmr = 2.9-3.4, Qmm = 3.1 ±
0.2, n = 25 spores, s = 13 specimens], fusoid to
narrowly ellipsoid, smooth, hyaline, inamyloid,
thin-walled. Basidia 14-16 × 8-9 µm, clavate,
4-spored. Basidioles fusoid to clavate.
Cheilocystidia abundant, composed of Siccus-
type broom cells; main body (9-)15-28 × 6-15
µm, cylindrical to clavate, hyaline, inamyloid,
thin-walled; apical setulae 2-5(-8) × 1(-2) µm,
conical to cylindrical, obtuse to subacute,
yellow, thick-walled. Pleurocystidia absent.
Pileipellis hymeniform, mottled, composed of 2
types of cells: a) Siccus-type broom cells with
main body 14-26(-41) × 6-10 µm, cylindrical to
clavate, seldom branched, hyaline to yellow,
inamyloid, thin- to thick-walled; apical setulae
(2-)4-8(-14) × 1-1.5 µm, crowded, cylindrical,
obtuse to subacute, brownish yellow, thickwalled; b) pileosetae, common, scattered, 60300 × 5-15 µm, lanceolate to fusoid, subacute
to acute, golden to golden brown, inamyloid,
thick-walled (up to 2 µm). Pileus trama hyphae
3-6(-9) µm, interwoven, cylindrical to inflated,
smooth, hyaline, dextrinoid, thin-walled.
Lamellar trama hyphae 3-8(-10) µm diam.,
regular to intervowen, cylindrical to inflated,
smooth, hyaline, dextrinoid, thin-walled, nongelatinous. Stipitipellis hyphae 3-5 µm diam,
subparallel, cylindrical, yellowish brown to
brown, smooth, inamyloid to weakly dextrinoid,
thick-walled (up to 1.5 µm), non-gelatinous.
Stipe trama hyphae 4-7(-11) µm diam.,
subparallel, cylindrical, hyaline, smooth,
dextrinoid,
thin-walled,
non-gelatinous.
Caulosetae abundant, 22-213 × 4-7(-15) µm,
fusoid to lanceolate, sometimes cylindrical,
brown, inamyloid, thick-walled. Clamp
connections present in all tissues.
Fig. 34. Marasmius trichotus (N. Wannathes 262). 1.
Basidiomes 2. Basidia 3. Cheilocystidia 4. Basidiospores
5a. Siccus-type pileipellis 5b. Pileosetae 6. Caulosetae,
Scale bar 1 = 10 mm, 2-6 = 10 µm
253
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, Northern Thailand, Papua New Guinea
and Singapore.
Material examined: Thailand. Chiang Mai
Province, Doi Inthanon National Park Junction of
Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E
98º 29.6’ alt. 1,703 m., 26 June 2003, N. Wannathes 006
(CMU, SFSU); same location, 25 June 2004, N.
Wannathes 178 (CMU, SFSU); same location, 3 July
2004, Y.S. Tan 295 and 296 (CMU, SFSU); same
location, 27 June 2005, N. Wannathes 355 (CMU,
SFSU); Chiang Mai Province, Mushroom Research
Centre, 27 km marker on Hwy 1095, 13 July 2003, N.
Wannathes 024 (CMU, SFSU); same location, 27 July
2003, N. Wannathes 037 and N. Wannathes 039(CMU,
SFSU); same location, 28 July 2003, N. Wannathes 045
(CMU, SFSU); same location, 10 August 2003, N.
Wannathes 098 (CMU, SFSU); Chiang Mai Province,
Doi Suthep-Pui National Park, Mokfa Waterfall, on Hwy
1095, N19º 6.5’.E 98º 46.3’ alt. 1014 m., 1 August 2003,
N. Wannathes 074 (CMU, SFSU), same location, 25
June 2005, N. Wannathes 326 (CMU, SFSU); Chiang
Mai Province, Mae Taeng District, New Waterfall, on 36
km. marker of Hwy1095, 20 August 2004, N. Wannathes
262 and N. Wannathes263 (CMU, SFSU); Phrae
Province, Muang District, Suen Keun, Na Koo Haa
Waterfall, 15 August 2005, N. Wannathes 408 (CMU,
SFSU); Phrae Province, Muang District, Pa Dang, Nong
Kam Village, near Nong Kam reservoir, 18 August 2005,
N. Wannathes 429 (CMU, SFSU).
Discussion: Marasmius trichotus is easily
recognized in the field by the orange to
brownish orange pileus stipe, both covered
with erect golden brown hairs. In addition, the
species has close (16-22) lamellae, narrowly
ellipsoid basidiospores with mean 12.1 × 4.1
µm, and lacks hymenial setae. Thai populations
differ from Papua New Guinea and Singapore
populations only in forming larger pilei (6-23
mm diam. vs 4-12 mm diam.) that are slightly
paler. All pertinent diagnostic features are
indistinguishable.
30. Marasmius nummularius Berk. &
Broome, J. Linn. Soc. Bot. 14: 33. 1873.
(Figs 35, 38-5)
Pileus 6-20 mm diam., broadly conical to
convex when young, plano-convex to
depressed and often umbonate in age, pruinose
to subvelutinous, dull; disc wrinkled, reddish
brown (8E7); margin striate, brown (6-7E6-8).
Context pale orange (6A3), thin. Lamellae
adnexed to subadnate, subdistant (12-18) with
2-3 series of lamellulae, narrow, pale orange
(6A3) with pinkish brown edge, non254
intervenose. Stipe 20-45 × < 1 mm, central,
cylindrical, hollow, velutinous to hispid, noninsititious, apex brownish orange (6C6-8), base
reddish brown (8E7). Odor and taste not
distinctive.
Basidiospores 12-14 × 3-4.5 µm [xmr =
12.9-13.6 × 3.8-4.0 µm, xmm = 13.2 ± 0.5 × 3.9
± 0.1 µm, Qmr = 3.3-3.6, Qmm = 3.4 ± 0.2, n =
25 spores, s = 2 specimens], fusoid to narrowly
ellipsoid, smooth, hyaline, inamyloid, thinwalled. Basidia not observed. Basidioles fusoid
to clavate. Cheilocystidia abundant, composed
of Siccus-type broom cells; main body 14-21 ×
5-13 µm, cylindrical to clavate or broadly
clavate, hyaline, inamyloid, thin-walled; apical
setulae 2-7 × 1 µm, cylindrical, obtuse to
subacute, dark brown to black, thick-walled.
Pleurocystidia absent or rare, of Siccus-type
broom cells like the cheilocystidia; pleurosetae
absent. Pileipellis hymeniform, mottled,
composed of 2 types of cells: a) Siccus-type
broom cells with main body (5-)14-28 × 7-18
µm, cylindrical to clavate or turbinate, hyaline
to yellow, inamyloid, thin- to thick-walled;
apical setulae (2-)3-9(-13) × 1-1.5 µm,
crowded, cylindrical, obtuse to subacute,
brown to reddish brown, thick-walled; b)
pileosetae, absent or rare, scattered, 100-115 ×
8-13 µm lanceolate to fusoid, subacute to acute,
hyaline to reddish brown, inamyloid, thickwalled. Pileus trama interwoven, dextrinoid.
Lamellar trama regular; hyphae 4-9 µm diam.,
cylindrical, smooth, hyaline, dextrinoid, thinwalled, non-gelatinous. Stipitipellis hyphae 3-6
µm diam., subparallel, cylindrical, yellow to
light brown, smooth, inamyloid, thick-walled
(up to 1 µm), non-gelatinous. Stipe trama
hyphae 4-7 µm diam., subparallel, cylindrical,
hyaline, smooth, dextrinoid, thin-walled, nongelatinous. Caulocystidia composed of 2 types
of cells: a) Siccus-type broom cells with main
body 8-18(-36) × 5-10(-15) µm, cylindrical to
clavate or irregular in outline, hyaline to yellow,
inamyloid, thin-walled; apical setuale 2-9(-23)
×1-2 µm, cylindrical, subacute to acute, yellow,
thick-walled; b) caulosetae 45-110 × 9-16 µm,
fusoid to lanceolate, often apically or centrally
setulose (2-4 setules per cell), hyaline to yellow,
inamyloid, thick-walled. Clamp connections
present in all tissues.
Fungal Diversity
Thai material matches nicely with Indonesian
material as described by Desjardin et al. (2000).
Fig. 35. Marasmius nummularius (N. Wannathes 266). 1.
Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5a. Siccus-type pileipellis 5b. Pileosetae
6a. Siccus-type caulocystidia 6b. Caulosetae, Scale bar 1
= 10 mm, 2-6 = 10 µm
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, Sri Lanka, Java and Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mokfa
Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
m., 21 August 2004, N. Wannathes 226, (CMU, SFSU);
Chiang Mai Province, Doi Suthep-Pui National Park,
Medicinal Plant Garden, 3 August 2005, N. Wannathes
396 (CMU, SFSU).
Discussion: Marasmius nummularius is
characterized by a reddish brown rugulosestriate pileus, subdistant (12-18) lamellae with
pinkish brown edges, narrowly ellipsoid
basidiospores with mean 13.2 × 3.9 µm,
scattered pileosetae, no pleurosetae, and a
velutinous to hispid stipe covered with
caulosetae that are often apically setulose. The
31. Marasmius coklatus Desjardin, Retnowati
& E. Horak, Sydowia 52: 146. 2000. (Fig. 36)
Pileus 17-33 mm diam., broadly conical
to campanulate, subumbonate, smooth to
weakly striate, subvelutinous, dull, disc dark
brown (7F8), margin brown (6E6-7) to
yellowish brown (5E7). Context greyish brown,
thin. Lamellae adnate, subdistant (12-15) with
1 series of lamellulae, broad, orangish white
(5A2) with yellowish brown edges, nonintervenose. Stipe 18-35 × 1.5-2 mm, central,
cylindrical, hollow, pruinose, non-insititious,
apex pale yellow (4A3), base brown (6E8).
Odor and taste not distinctive.
Basidiospores (10-)11-12(-13) × (5.5-)67 µm [xm = 11.2 ± 0.6 × 6.1 ± 0.3 µm, Q = 1.62.0, Qm = 1.8, n = 25 spores, s = 1 specimen],
ellipsoid, smooth, hyaline, inamyloid, thinwalled. Basidia not observed. Basidioles fusoid
to clavate. Cheilocystidia abundant, composed
of 2 types of cells: a) Siccus-type broom cells,
scattered with main body (9-)16-20 × 6-9 µm,
cylindrical to clavate or turbinate, hyaline,
inamyloid, thick-walled; apical setulae 4-12 ×
1-3 µm, cylindrical to conical, obtuse to
subacute, brown, thick-walled; b) cheilosetae
20-44 × 4-6 µm, fusoid to lanceolate, often
forked, subacute to acute, brown, inamyloid,
thick-walled. Pleurosetae abundant, 20-44 × 46 µm, fusoid to lanceolate, often forked,
subacute to acute, brown, inamyloid, thickwalled. Pileipellis hymeniform, mottled,
composed of 2 types of cells: a) Siccus-type
broom cells with main body (9-)16-37 × 6-11
µm, cylindrical to clavate, hyaline to yellow,
inamyloid, thick-walled; apical setulae 4-20×
1.5-3 µm, crowded, cylindrical to conical,
subacute, brown to dark brown, thick-walled; b)
pileosetae, common, scattered, 35-38 × 7-9 µm
lanceolate to fusoid, conical to subacute,
yellow to yellowish brown, inamyloid, thickwalled. Pileus trama interwoven, inamyloid.
Lamellar trama hyphae 6-12 µm diam.,
interwoven, cylindrical to inflated, smooth,
hyaline, inamyloid, thin-walled, non- gelatinous.
Stipitipellis hyphae 6-9(-12) µm diam., parallel,
cylindrical, brown to light brown, smooth,
inamyloid, thick-walled (up to 2 µm),
255
pruinose stipe covered with caulosetae,
moderately long and broad basidiospores, and
numerous pileo-, cheilo-, pleuro- and
caulosetae that are often forked.
32. Marasmius araneocephalus Wannathes,
Desjardin & Lumyong, sp. nov.
(Figs 37, 38-6)
MycoBank: MB512418
Fig. 36. Marasmius coklatus (S.Y. Tan.301) 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Hymenial setae 5a. Siccus-type pileipellis 5b. Pileosetae
6a. Siccus-type caulocystidia 6b. Caulosetae, Scale bar 1
= 10 mm, 2-6 = 10 µm
non-gelatinous. Stipe trama hyphae 7- 17 µm
diam., parallel, cylindrical, hyaline, smooth,
weakly dextrinoid to inamyloid, thin-walled,
non-gelatinous. Caulocystidia composed of 2
types of cells: a) Siccus-type broom cells with
main body 18-26 × 7 µm, cylindrical or
irregular in outline, often lobed, yellow,
inamyloid, thick-walled; apical setuale 15-26 ×
2-3 µm, ranging from 2-4 setulae per cell,
cylindrical, subacute to acute, yellow to light
brown, thick-walled; b) caulosetae 39-52 × 6-7
µm, fusoid to lanceolate, seldom lobed, yellow,
inamyloid, thick-walled. Clamp connections
present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, Java and Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Inthanon National Park Junction of
Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E
98º 29.6’ alt. 1,703 m., 3 July 2004, T.Y. Shin 301
(CMU, SFSU).
Discussion:
Marasmius
coklatus,
described originally from material collected in
Indonesia by Desjardin et al. (2000), is
characterized by a dark chocolate brown,
velutinous pileus, distant to subdistant, broad,
lamellae with yellowish brown edges, a brown,
256
Etymology: ‘aranea’ = spider; ‘cephalus’ = head;
referring to the spider-like broom cells on the pileus
surface.
Pileus 20-42 mm diametro, convexus,
depressione exigua ad discum regulosum, pellucidostriatus usque reticulato-venosus, glaber, hebetatus, disco
purpureo-atrobrunneo,
margine
cinereo-brunneo.
Contextus rubro-cinereus, tenuis. Lamellae adnatae,
subdistantes (10-12) cum 3 seriebus lamellularum, latae
(3-5 mm), brunneo-aurantiacae, atropurpureomarginatae,
eminenter intervenosae usque reticulatae. Stipes 27-45 ×
1-2 mm, centralis, cylindratus, cavus, hispidus cum pilis
purpureo-atrobrunneis, haud insititius, apice brunneocinereo, basi laete brunnea. Odor saporque non propria.
Basidiosporae 6-7 (-8) × 3.5-4 µm, ellipsoideae, laeves,
hyalinae, inamyloideae, tenuitunicatae. Basidia non
observata. Basidiolae fusoideae usque clavatae.
Cheilocystidia abundantia, 2 typorum: a) cellulis typi
Sicci 13-22 × 5-7 µm, cylindratis vel adumbratim
inaequabilibus, hyalinis, inamyloideis, tenui- usque
crassetunicatis (usque ad 0.5 µm); 2-4 setulis apicalibus,
3-16 × 1-3 µm, cylindratis usque conicis, obtusis usque
subacutis, flavis, tenui- usque crassetunicatis (usque ad
0.5 µm); b) cheilosetis 19-22 × 4-6 µm, cylindratis vel
adumbratim inaequabilibus, obtusis usque subacutis,
hyalinis, inamyloideis, tenui- usque crassetunicatis
(usque ad 0.5 µm). Pleurosetae vulgares, cheilosetis
similes, 25-38 × 4-6 µm, fusoideae usque ventricosofusoideae, raro clavatae, obtusae usque subacutae,
interdum furcatae, pallide flavae, inamyloideae, tenuiusque crassetunicatae. Pileipellis hymeniformis, haud
maculosus, 2 typorum: a) cellulis typi Sicci 13-18 × 6-10
µm, cylindratis usque clavatis vel late clavatis, hyalinis,
inamyloideis, tenuitunicatis; setulis apicalibus 2-5 (-23)
× 1-2 (-4) µm, coarctatis, cylindratis usque conicis,
obtusis usque subacutis, purpureo-brunneis, tenuitunicatis; b) cellulis araneae similibus, dispersis, 16-38 ×
7-8 µm, cylindratis vel adumbratim inaequabilibus,
flavis usque laete brunneis, inamyloideis, crassetunicatis;
3-6 setulis apicalibus, (4-) 30-70 × 2-3 µm, cylindratis,
subacutis usque acutis, flavis usque laete brunneis,
inamyloideis, crassetunicatis. Trama pilei intertexta,
dextrinoidea. Trama lamellae intertexta, hyphis 2-7 µm
diametro, cylindratis, laevibus, hyalinis, dextrinoideis,
tenuitunicatis, haud gelatinosis. Stipitipellis subparallelus,
hyphis 4-8 µm diametro, cylindratis, flavis usque laete
brunneis, laevibus, dextrinoideis, tenui- usque
crassetunicatis (usque ad 1 µm), haud gelatinosis. Trama
stipitis parallela, hyphis 8-13 µm diametro, cylindratis,
hyalinis, laevibus, dextrinoideis, tenuitunicatis, haud
gelatinosis. Caulocystidia 2 typorum: a) cellulis typi
Fungal Diversity
Sicci 12-31 × 5-6 µm, cylindratis, hyalinis, inamyloideis,
tenui- usque crassetunicatis; 2-3 setulis apicalibus, 10-20
× 1.5-2 µm, cylindratis, obtusis usque subacutis, hyalinis
usque flavis, crassetunicatis; b) caulosetis 40-50 × 4-6
µm, polymorphis, cylindratis usque clavatis vel
adumbratim inaequabilibus, saepae furcatis, hyalinis,
inamyloideis, crassetunicatis. Fibulae praesentes in
omnibus texturis.
Typus: Chiang Mai Province, Doi Inthanon
National Park Junction of Highway 1,009 and road to
Mae Chaem, N 18º 31.6’ E 98º 29.6’ alt. 1,703 m.,
dispersus usque gregarius in folis plantae dicotyledoneae
vel ligno, 27 June 2005, N. Wannathes 358 (CMU:
holotypus; SFSU: isotypus).
Pileus 20-42 mm diam., convex with a
slightly depressed, rugulose disc, pellucidstriate to venose-reticulate, glabrous, dull, disc
dark purplish brown (14F4), margin greyish
brown (7E3). Context reddish grey (12C2), thin.
Lamellae adnate, subdistant (10-12) with 3
series of lamellulae, broad (3-5 mm), brownish
orange (6C3) with dark purple edges, strong
intervenose to reticulate. Stipe 27-45 × 1-2 mm,
central, cylindrical, hollow, hispid with dark
purple (14F4) hairs, non-insititious, apex
brownish grey (11C2), base light brown. Odor
and taste not distinctive.
Basidiospores 6-7 (-8) × 3.5-4 µm [xm =
6.6 ± 0.6 × 3.9 ± 0.3 µm, Q = 1.2 - 2.0, Qm =
1.7, n = 25 spores, s = 1 specimen], ellipsoid,
smooth, hyaline, inamyloid, thin-walled.
Basidia not observed. Basidioles fusoid to
clavate. Cheilocystidia abundant, composed of
2 types of cells: a) Siccus-type broom cells
with main body 13-22 × 5-7 µm, cylindrical or
irregular in outline, hyaline, inamyloid, thin- to
thick-walled (up to 0.5 µm); apical setulae 3-16
× 1-3 µm, cylindrical to conical, ranging from
2-4 setulae per cell, obtuse to subacute, yellow,
thin- to thick-walled (up to 0.5 µm); b)
cheilosetae 19-22 × 4-6 µm, cylindrical or
irregular in outline, obtuse to subacute, hyaline,
inamyloid, thin- to thick-walled (up to 0.5 µm).
Pleurosetae common, like the cheilosetae, 2538 × 4-6 µm, fusoid to fusoid-ventricose, rarely
clavate, obtuse to subacute, sometimes forked,
pale yellow, inamyloid, thin- to thick-walled.
Pileipellis hymeniform, not mottled, composed
of 2 types of cells: a) Siccus-type broom cells
with main body 13-18 × 6-10 µm, cylindrical
to clavate or broadly broom cells, scattered,
with main body 16-38 × 7-8 µm, cylindrical or
irregular in outline, yellow to light brown,
inamyloid, thick-walled; apical setulae(4-) 3070 × 2-3 µm, ranging from 3-6 setulae per cell,
cylindrical, subacute to acute, yellow to light
brown, inamyloid, thick-walled. Pileus trama
interwoven, dextrinoid. Lamellar trama hyphae
2-7 µm diam., interwoven, cylindrical, smooth,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Stipitipellis hyphae 4-8 µm, subparallel,
cylindrical, yellow to light brown,smooth,
dextrinoid, thin- to thick-walled (up to 1 µm),
non-gelatinous. Stipe trama hyphae 8-13 µm,
parallel, cylindrical, hyaline, smooth, dextrioid,
thin-walled, non-gelatinous. Caulocystidia
composed of 2 types of cells: a) Siccus-type
broom cells with main body 12-31 × 5-6 µm,
cylindrical, hyaline, inamyloid, thin- to thickwalled; apical setulae 10-20 × 1.5-2 µm,
ranging from 2-3 setulae per cell, cylindrical,
obtuse to subacute, hyaline to yellow, thickwalled; b) caulosetae 40-50 × 4-6 µm,
polymorphic, cylindrical to clavate or irregular
in outline, often forked, hyaline, inamyloid,
thick-walled. Clamp connections present in all
tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves or on wood, Northern Thailand.
Fig. 37 Marasmius araneocephalus (N. Wannathes358) 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pleurocystidia 5a. Siccus-type pileipellis 5b. Spider-like
pileosetae 6. Caulocystidia, Scale bar 1 = 10 mm, 2-6 =
10 µm
257
Fig. 38. Basidiomata of Marasmius section Neosessiles, Leveilleani and Section Sicci ser. Spinulosi. 1. M. tenuissimus
(N. Wannathes 199) 2. M. leveilleanus (N. Wannathes 248) 3. M. coarctatus (N. Wannathes 385) 4. M. trichotus (N.
Wannathes 326) 5. M. nummularius (N. Wannathes 266) 6. M. araneocephalus (N. Wannathes 358), Scale bars = 20
mm.
Material examined: Thailand. Chiang Mai
Province, Doi Inthanon National Park Junction of
Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E
98º 29.6’ alt. 1,703 m., 27 June 2005, N. Wannathes 358
(CMU: holotype; SFSU: isotype).
Discussion: Marasmius araneocephalus
has the following diagnostic features: a venosereticulate, greyish brown pileus with dark
purplish brown disc; subdistant (10-12),
strongly intervenose to reticulate lamellae; a
light brown stipe covered with dark purple
hairs; small basidiospores with mean 6.6 × 3.9
µm; and scattered spider-like broom cells on
the pileus surface with setulae 30-70 µm long.
Many micromorphological features of M.
araneocephalus are similar to those of M.
258
coklatus, but the latter species differs in
forming larger basidiospores (xm = 11.2 × 6.1
µm), a smooth, dark brown pileus, nonintervenose-reticulate lamellae, and lacks the
distinctive spider-like pileipellis cells. In the
field M. araneocephalus look like a dark form
of M. albimyceliosus, but the latter is a member
of sect. Globulares with distinctly different
micromorphological features.
Section Sicci, subsect. Siccini, ser.
Atrorubentes Desjardin & Horak, Biblio.
Mycol. 168: 27. 1997.
Type: Agaricus atrorubens Berk.,
London J. Bot. 1: 138. 1842.
Fungal Diversity
[= Marasmius atrorubens (Berk.) Berk.,
Hooker's J. Bot. Kew. Gard. Misc. 8: 137.
1856].
33. Marasmius auratus Wannathes, Desjardin
& Lumyong, sp. nov.
(Figs 39, 49-1)
MycoBank: 512419
Etymology: ‘auratus’ = gold; referring to the gold
color of the pileus.
Pileus 5-30 mm diametro, late conicus, saepe
margine erecto, laevis usque striatulatus, pruinosus,
hebetatus, disco auruto, margine aurantiaco usque flavoaurantiaco vel cinereo-flavo. Contextus laete aurantiacus,
tenuis. Lamellae adnatae, arctae (17-24) cum 3-4
seriebus lamellularum, angustae, creameae, haud
marginatae, haud intervenosae. Stipes 35-68 × 1.5-2 mm,
centralis, cylindratus, cavus, hispidulus usque hispidus,
haud insititius, apice flavo-albo, basi rubro-atrobrunnea.
Odor saporque non propria. Basidiosporae 11-14 × 4-5
µm, ellipsoideae, oblique curvae, laeves, hyalinae,
inamyloideae, tenuitunicatae Basidia non observata.
Basidiolae clavatae. Cheilocystidia nulla vel simplicia,
22-28 × 4-6 µm, cylindrata usque clavata vel
adumbratim inaequabilia, raro lobata, hyalina,
inamyloidea, tenuitunicata. Pleurocystidia nulla.
Pileipellis hymeniformis, maculosus, typi Sicci; 9-26 ×
5-11 µm, clavatus usque late clavatus, turbinatus vel
adumbratim inaequabilis, hyalinus, inamyloideus, tenuiusque crassetunicatus; setulis apicalibus 3-9 × 1-2 µm,
coartatis, cylindratis usque adumbratim inaequabilibus,
obtusis usque subacutis, flavis, crassetunicatis. Trama
pilei intertexta, dextrinoidea. Trama lamellae intertexta,
hyphis (2-)4-8(-12) µm diametro, cylindratis usque
inflatis, laevibus, hyalinis, dextrinoideis, tenuitunicatis,
haud gelatinosis. Stipitipellis parallelus, hyphis 5-10(-15)
µm diametro, cylindratis, flavis usque brunneis, laevibus,
leniter dextrinoideis usque dextrinoideis, tenui- usque
crassetunicatis (usque ad 0.5 µm), haud gelatinosis.
Trama stipitis parallela, hyphis (4-)6-9(-11) µm diametro,
cylindratis, hyalinis, laevibus, dextrinoideis, tenuitunicatis, haud gelatinosis. Caulocystidia numerosa, 22-12 ×
5-7 µm, cylindrata usque adumbratim inaequabilia,
hyalina, inamyloidea usque leniter dextrinoidea,
crassetunicata (usque ad 1 µm). Fibulae praesentes in
omnibus texturis.
Typus: Chiang Mai Province, Doi Inthanon
National Park Junction of Highway 1,009 and road to
Mae Chaem, N 18º 31.6’ E 98º 29.6’ alt. 1,703 m.,
dispersus usque gregarius in folis plantae dicotyledoneae,
raro in ligno, 25 June 2004, N. Wannathes 175 (CMU:
holotypus; SFSU: isotypus).
Pileus 5-30 mm diam., broadly conical,
often with upturned margin, smooth to
striatulate, pruinose, dull, disc golden yellow
(5B7), margin orange (5A6-7) to yellowish
orange (4B6-7) or greyish yellow (4B5).
Context light orange (5A4), thin. Lamellae
adnate, close (17-24) with 3-4 series of
lamellulae, narrow, cream, non-marginate, nonintervenose. Stipe 35-68 × 1.5-2 mm, central,
cylindrical, hollow, hispidulose to hispid, noninsititious, apex yellowish white, base dark
reddish brown. Odor and taste not distinctive.
Basidiospores 11-14 × 4-5 µm [xmr =
11.9-12.6 × 4.0-4.3, xmm = 12.2 ± 0.4 × 4.1 ±
0.1 µm, Qmr = 2.8 - 3.1, Qmm = 3.0 ± 0.2, n =
25 spores, s = 3 specimens], ellipsoid, curved
in profile, smooth, hyaline, inamyloid, thinwalled. Basidia not observed. Basidioles
clavate.
Fig. 39. Marasmius auratus (N. Wannathes351) 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pileipellis 5. Caulocystidia, Scale bar 1 = 20 mm, 2-5 =
10 µm
Cheilocystidia absent or simple, 22-28 × 4-6
µm, cylindrical to clavate or irregular in outline,
seldom lobed, hyaline, inamyloid, thin-walled.
Pleurocystidia absent. Pileipellis hymeniform,
mottled, composed of Siccus-type broom cells;
main body 9-26 × 5-11 µm, clavate to broadly
clavate, turbinate or irregular in outline,
hyaline, inamyloid, thin- to thick-walled; apical
setulae 3-9 × 1-2 µm, crowded, cylindrical to
irregular in outline, obtuse to subacute, yellow,
thick-walled. Pileus trama interwoven, dextrinoid. Lamellar trama hyphae (2-)4-8(-12)
µm diam., interwoven, cylindrical to inflated,
259
smooth, hyaline, dextrinoid, thin- walled, nongelatinous. Stipitipellis hyphae 5-10(-15) µm
diam., parallel, cylindrical, yellow to brown,
smooth, weakly dextrinoid to dextrinoid, thinto thick-walled (up to 0.5 µm), non-gelatinous.
Stipe trama hyphae (4-)6-9(-11) µm diam.,
parallel, cylindrical, hyaline, smooth, dextrinoid, thin-walled, non-gelatinous. Caulocystidia numerous, simple, 22-12 × 5-7 µm,
cylindrical to irregular in outline, obtuse,
hyaline, inamyloid to weakly dextrinoid, thickwalled (up to 1 µm). Clamp connections
present in all tissues.
Habit, habitat and known distribution:
Solitary, scattered to gregarious on dicotyledonous leaves, or rarely on wood. Northern
Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Inthanon National Park Junction of
Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E
98º 29.6’ alt. 1,703 m., 2 August 2003, N. Wannathes
076 (CMU, SFSU); same location, 25 June 2004, N.
Wannathes 175 (CMU: holotype; SFSU: isotype); same
location, 27 June 2005, N. Wannathes 351 (CMU,
SFSU).
Discussion: Marasmius auratus is
characterized by the following features: a
golden yellow, smooth to striatulate pileus;
close lamellae; dark reddish brown stipe;
slender, moderately long basidiospores with
mean 12.2 × 4.1 µm; cheilocystidia that are
absent or only simple and basidiole-like;
abundant, simple, cylindrical, obtuse, thickwalled, inamyloid caulocystidia; and growth on
dicotyledonous leaves. This new Thai species
is most phenetically similar to several
Malaysian species (Tan et al., 2009) that have
simple caulocystidia. Marasmius musicolor
Y.S. Tan & Desjardin differs in forming fewer
lamellae (12-14), larger basidiospores with
mean 16.1 × 3.9 µm, has distinct Siccus-type
cheilocystidia, and dextrinoid caulocystidia
formed in clusters. Marasmius ochropoides
Y.S. Tan & Desjardin differs in forming a
smaller pileus (5-6 mm diam.), paler stipe
(orangish white), smaller basidiospores with
mean 10.6 µm, well-developed Siccus-type
cheilocystidia, and dextrinoid caulocystidia.
Marasmius iras Y.S. Tan & Desjardin differs
in forming a brown pileus, slightly longer
basidiospores with mean 14.1 × 3.7 µm, welldeveloped Siccus-type cheilocystidia, and
smaller, lobed or forked caulocystidia.
260
Marasmius auratus is also similar to two
American species that are members of this
series. Marasmius actinopus Mont. sensu
Singer from South America differs in forming
a sulcate pileus colored ochraceus yellow to
dull
olive
brown,
slightly
narrower
basidiospores (3-4 µm diam.), and has Siccustype cheilocystidia (Singer, 1976; see also
Desjardin and Horak, 1997: 27). Marasmius
ciliatomarginatus Desjardin, described from
North America, differs in forming a reddish
brown pileus, larger basidiospores with mean
15.7 × 4.0 µm, and red-marginate lamellae with
cheilocystidia having reddish orange contents
(Desjardin and Petersen, 1989).
34. Marasmius pseudopellucidus Wannathes,
Desjardin & Lumyong, sp. nov.
(Figs 40, 49-3)
MycoBank: MB512420
Etymology: ‘pseudo’ = false; referring to the
basidiomes which look like M. pellucidus.
Pileus 10-40 mm diametro, late conicus usque
convexus cum disco leviter depresso et margine reflexo
ubi vetus, pellucido-striatus, pruinosus, hebetatus, primo
disco pallide flavo et margine albo demum ubique albus.
Contextus albidus, tenuis. Lamellae adnatae, arctae (1922) cum 2-3 seriebus lamellularum, angustae, albae,
haud marginatae, intervenosae ubi vetus. Stipes 16-50 ×
1-2 mm, centralis, cylindratus, cavus, subvelutinus usque
velutinus, haud insititius, apice albo, basi laete
aurantiaca. Odor suavis. Sapor non proprius.
Basidiosporae 10-12 × 3.5-4.5 µm, ellipsoideae, oblique
curvae, laeves, hyalinae, inamyloideae, tenuitunicatae
Basidia 33-35 × 7-8 µm, clavata, 4-spora. Basidiolae
clavatae. Margo lamellae sterilis; cheilocystidia
simplicia, basidiolae similibus, 13-18 × 3.5-10 µm,
cylindrata usque clavata, raro adumbratim inaequabilia,
hyalina, inamyloidea, tenuitunicata. Pleurocystidia nulla.
Pileipellis hymeniformis, maculosus, typi Sicci; (9-)1323 × 7-13µm, clavatus usque late clavatus vel turbinatus,
saepe ramosus, hyalinus, inamyloideus, tenui- usque
crassetunicatus; setulis apicalibus 2-12 × 1-1.5µm,
coartatis, cylindratis, obtusis usque subacutis, flavis,
crassetunicatis. Trama pilei intertexta, dextrinoidea.
Trama lamellae intertexta, hyphis 4-9(-14) µm diametro,
cylindratis
usque
inflatis,
laevibus,
hyalinis,
dextrinoideis,
tenuitunicatis,
haud
gelatinosis.
Stipitipellis parallelus, hyphis (4-)6-10 µm diametro,
cylindratis, flavis usque brunneo-flavis, laevibus,
dextrinoideis, crassetunicatis (usque ad 1 µm), haud
gelatinosis. Trama stipitis parallela, hyphis 6-14 µm
diametro, cylindratis, hyalinis, laevibus, dextrinoideis,
tenuitunicatis, haud gelatinosis. Caulocystidia numerosa,
23-87 × 6-9 µm, cylindrata usque adumbratim
inaequabilia, raro lobata, hyalina usque pallide flava,
leniter dextrinoidea, tenui- usque crassetunicata (usque
ad 2 µm). Fibulae praesentes in omnibus texturis.
Fungal Diversity
Typus: Chiang Rai Province, Muang District,
Khun Con Waterfall, dispersus usque gregarius, saepe
caespitosus in putredine e bambusa, raro in folis plantae
dicotyledoneae vel putridine ligneus , 12 June 2005, N.
Wannathes 305 (CMU: holotypus; SFSU: isotypus).
Pileus 10-40 mm diam., broadly conical
to convex with a slightly depressed disc and
reflexed margin in age, pellucid-striate,
pruinose, dull, disc pale yellow, margin white,
fading to white overall. Context off-white, thin.
Lamellae adnate, close (19-22) with 2-3 series
of lamellulae, narrow, intervenose in age, white,
non-marginate. Stipe 16-50 × 1-2 mm, central,
cylindrical, hollow, subvelutinous to velutinous,
non-insititious, apex white, base light orange
(5B5). Odor sweet. Taste not distinctive.
Basidiospores 10-12 × 3.5-4.5 µm [xmr =
11.0-11.1 × 4.0-4.2 µm, xmm = 11.0 ± 0.0 × 4.1
± 0.1 µm, Qmr = 2.6-2.8, Qmm = 2.7 ± 0.1, n =
25 spores, s = 2 specimens], ellipsoid, curved
in profile, smooth, hyaline, inamyloid, thinwalled. Basidia 33-35 × 7-8 µm, clavate, 4spored. Basidioles clavate. Lamellar edge
sterile; cheilocystidia simple, basidiole-like,
13-18 × 3.5-10 µm, cylindrical to clavate,
seldom irregular in outline, hyaline, inamyloid,
thin-walled. Pleurocystidia absent. Pileipellis
hymeniform, mottled, composed of Siccus-type
broom cells; main body (9-)13-23 × 7-13 µm,
clavate to broadly clavate or turbinate, often
branched, hyaline, inamyloid, thin- to thickwalled; apical setulae 2-12 × 1-1.5 µm,
crowded, cylindrical, obtuse to subacute,
yellow, thick-walled. Pileus trama interwoven,
dextrinoid. Lamellar trama hyphae 4-9(-14)
µm diam., interwoven, cylindrical to inflated,
smooth, hyaline, dextrinoid, thin-walled, nongelatinous. Stipitipellis hyphae (4-)6-10 µm
diam., parallel, cylindrical, yellow to brownish
yellow, smooth, dextrinoid, thick-walled (up to
1 µm), non-gelatinous. Stipe trama hyphae 614 µm diam., parallel, cylindrical, hyaline,
smooth, dextrinoid, thin-walled, non-gelatinous.
Caulocystidia numerous, 23-87 × 6-9 µm,
cylindrical to irregular in outline, rarely lobed,
hyaline to pale yellow, weakly dextrinoid, thinto thick-walled (up to 2 µm). Clamp
connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious, often in cespitose
clusters on bamboo debris or seldom on
dicotyledonous leaves
Northern Thailand.
or
woody
debris,
Material examined: Thailand. Chiang Mai
Province, Huai Nam Dang National Park, North 19º
18.3’ E 98º 35.8’ alt. 1,538 m., 28 June 2004, N.
Wannathes 186 (CMU, SFSU); Chiang Rai Province,
Muang District, Khun Con Waterfall, 12 June 2005, N.
Wannathes 305 (CMU: holotype; SFSU: isotype).
Fig. 40. Marasmius pseudopellucidus (N. Wannathes305)
1. Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pileipellis 5. Caulocystidia, Scale bar 1 = 20 mm, 2-5 =
10 µm
Discussion: Basidiomes of Marasmius
pseudopellucidus look like M. pellucidus, and
the two species may be confused in the field.
Marasmius pellucidus is a member of sect.
Globulares and is characterized by having
smooth, clavate to subglobose cells forming the
pileipellis, rather than Siccus-type broom cells
as in M. pseudopellucidus. The new Thai
species is most similar to two white African
species. Marasmius haediniformis Singer
differs in forming fewer lamellae (12-18 vs 1922), a stipe colored deep brown at the base,
numerous Siccus-type cheilocystidia, and lacks
caulocytidia (Antonín, 2007). Marasmius
subarborescens Singer differs in forming
smaller basidiospores (xm = 6.3 × 3 µm vs 11 ×
4.1 µm), Siccus-type cheilocystidia, and a
chestnut brown base stipe (Pegler, 1977;
Antonín, 2007). Marasmius pseudopellucidus
also phenetically similar to two white species
261
from the neotropics. Marasmius personatus
Berk. & M.A. Curtis differs in forming a
smaller pileus (6-10 mm diam. vs 5-40 mm
diam.), a fulvous or dark brown stipe, and has
pleurocystidia and Siccus-type cheilocystidia
(Singer, 1976). Marasmius pseudoniveus
Singer differs in forming a sulcate pileus,
darker stipe (chestnut brown to brown at the
base), smaller basidiospores (8-10.3 × 2.7-4
µm), Siccus-type cheilocystidia, and three
types of pileipellis cells (Singer, 1976).
35. Marasmius ochroleucus Desjardin & E.
Horak, Biblio. Mycol. 168: 35. 1997.
(Fig. 41)
Pileus 11-18 mm diam., convex to planoconvex, umbonate, smooth to striate, pruinose,
dull, hygrophanous, disc light yellow (4A2) to
cream (4A3-4), margin paler. Context off-white,
thin. Lamellae adnexed, close to crowded (2024) with 3-4 series of lamellulae, narrow,
intervenose, off-white to cream, non-marginate.
Stipe 35-58 × 1-1.2 mm, central, cylindrical,
hollow, hispidulous, non-insititious, apex offwhite to light yellow (4A4), base yellowish
orange (4A6) to light brown. Odor sweet. Taste
not distinctive.
Basidiospores 8-12 × 3.5-4 µm [xm = 9.7
± 1.0 × 3.9 ± 0.2 µm, Q = 2.0-3.1, Qm = 2.5, n
= 25 spores, s = 1 specimen], ellipsoid, curved
in profile, smooth, hyaline, inamyloid, thinwalled. Basidia not observed. Basidioles fusoid
to clavate. Cheilocystidia common, of Siccustype broom cells; main body 12-27 × 4-6 µm,
cylindrical or irregular in outline, hyaline,
inamyloid, thin-walled; apical setulae 3-10 × 12 µm, cylindrical, seldom branched, obtuse to
subacute, hyaline, thin-walled. Pleurocystidia
absent. Pileipellis hymeniform, weakly mottled,
composed of Siccus-type broom cells; main
body 13-20 × (3-)8-9 µm, clavate to broadly
clavate or turbinate, hyaline to pale yellow,
inamyloid, thin- to thick-walled; apical setulae
2-7 × 1 µm, crowded, cylindrical to irregular in
outline, obtuse to subacute, yellow, thick-walled.
Pileus trama interwoven, dextrinoid. Lamellar
trama hyphae 3-6 µm diam., interwoven,
cylindrical to inflated, smooth, hyaline,
dextrinoid,
thin-walled,
non-gelatinous.
Stipitipellis hyphae 4-10 µm diam., parallel,
cylindrical, yellow, smooth, weakly dextrinoid,
262
Fig. 41. Marasmius ochroleucus (N. Wannathes 299) 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pileipellis 5. Caulocystidia, Scale bar 1 = 20 mm, 2-5 =
10 µm
thin- to thick-walled (up to 1 µm), nongelatinous. Stipe trama hyphae 6-8(-13) µm
diam., parallel, cylindrical, hyaline, smooth,
dextrinoid,
thin-walled,
non-gelatinous,
Caulocystidia numerous, 14-53 × 5-12 µm,
cylindrical to fusoid-ventricose or irregular in
outline, obtuse, hyaline, inamyloid, thin- to
thick-walled. Clamp connections present in all
tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, New Caledonia and Northern Thailand.
Material examined: Thailand. Chiang Rai
Province, Muang District, Khun Con Waterfall, 12 June
2005, N. Wannathes 299 (CMU, SFSU).
Discussion: Marasmius ochroleucus,
originally described from material collected in
New Caledonia, is characterized by a smooth,
hygrophanous pileus colored light yellow to
cream, close to crowded, emarginate lamellae,
a pale yellowish orange to light brown
hispidulous stipe, numerous irregularly
cylindrical, obtuse, thin-walled caulocystidia,
and an absence of pleurocystidia. The Thai
material is nearly indistinguishable from the
protologue (differing only in forming fewer
lamellae) and the species is most phenetically
similar to M. auratus (described above) and M.
musicolor Y.S.Tan & Desjardin from Malaysia.
Marasmius auratus differs in forming a darker,
golden yellow pileus, larger basidiospores (xm
Fungal Diversity
= 12.2 × 4.1 µm vs 9.7 × 3.9 µm µm), and
lacks Siccus-type cheilocystidia. Marasmius
musicolor
differs
in
forming
larger
basidiospores with mean 16.1 × 3.9 µm, fewer
lamellae (12-14 vs 20-24), and has clustered
caulocystidia.
36. Marasmius luteolus Berk. & M.A. Curtis,
Proc. Amer. Acad. Arts 4: 119. 1860.
(Figs 42, 49-2)
Pileus 4-42 mm diam., convex to planoconvex, expanding to plano-depressed, striate
to sulcate, pruinose to minutely velutinous, dull,
disc bright brownish orange to golden orange,
margin pale cream-orange or paler to yellowish
orange. Context pale yellow, thin. Lamellae
adnate, subdistant (8-14) with 2-3 series of
lamellulae, narrow (< 4 mm), white to buff,
non-marginate, non-intervenose. Stipe 10-35 ×
1 mm, central, cylindrical with a subbulbous
base, hollow, pruinose, non-insititious, apex
yellowish white to buff, base brown. Odor and
taste not distinctive.
Basidiospores 9-12× 3.5-5 µm [xmr = 9.711 × 3.7-4.3 µm, xmm = 10.2 ± 0.5 × 4.1 ± 0.2
µm, Qmr = 2.4-3.0, Qmm = 2.6 ± 0.2, n = 25
spores, s = 6 specimens], ellipsoid, curved in
profile, smooth, hyaline, inamyloid, thinwalled. Basidia not observed. Basidioles fusoid
to clavate. Cheilocystidia and pleurocystidia
similar, common on the sides and edges of
lamellae, composed of Siccus-type broom cells;
main body (9-)12-33 × (5-)7-10 µm, cylindrical
to clavate or broadly clavate, hyaline,
inamyloid, thin- to thick-walled; apical setulae
3-12 × 1-2 µm, cylindrical to conical, wavy in
outline, subacute, pale yellow to yellow, thickwalled. Pileipellis hymeniform, mottled,
composed of Siccus-type broom cells; main
body 10-20 × 6-10µm, clavate to broadly
clavate, hyaline, inamyloid, thin-to thickwalled; apical setulae 3-11 × 1-2 µm, crowded,
cylindrical to conical, wavy in outline,
subacute, pale yellow to yellow, thick-walled.
Pileus trama interwoven, dextrinoid. Lamellar
trama hyphae 3-8(-10) µm diam., interwoven,
cylindrical, smooth, hyaline, dextrinoid, thinwalled, non-gelatinous. Stipitipellis hyphae 312 µm diam., parallel, cylindrical, yellowish
brown, smooth, dextrinoid, thick-walled (up to
2 µm), non-gelatinous. Stipe trama hyphae 4-
Fig. 42. Marasmius luteolus (N. Wannathes 138) 1.
Basidiomes 2. Basidiospores 3. Hymenial cystidia 4.
Pileipellis 5a. Non-setulose caulocystidia 5b. Siccus-type
caulocystidia, Scale bar 1 = 20 mm, 2-5 = 10 µm
15 µm diam., parallel, cylindrical, hyaline,
smooth, dextrinoid, thin-walled, non-gelatinous.
Caulocystidia composed of two types of cells:
a) Siccus-type broom cells with main body 818 × 6-7 µm, scattered, uncommon, cylindrical
or irregular in outline, hyaline; apical setulae 26 × 1 µm, cylindrical to conical, wavy in
outline, subacute, hyaline to pale-yellow, thickwalled; b) non-setulose cells, common, 11-60 ×
5-10 µm, polymorphic, cylindrical to clavate or
irregular in outline, often lobed, obtuse, hyaline,
inamyloid, thin- to thick-walled (up to 1 µm).
Clamp connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on bamboo debris or on
dicotyledonous leaves. Japan (Bonin Islands),
Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mokfa
Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
m., 28 June 2003, N. Wannathes 008 (CMU, SFSU);
same location, 28 August 2003, N. Wannathes 138(CMU,
SFSU); Chiang Mai Province, Doi Suthep-Pui National
Park, Sangasabhasri Land on the way to Huai Kok Ma
Village, N 18º 48.4’ E 98º 54.6’ alt. 1,146 m., 29 August
2003, N. Wannathes 143 and N. Wannathes 144 (CMU,
SFSU); Chiang Mai Province, Huai Nam Dang National
Park, North 19º 18.3’ E 98º 35.8’ alt. 1,538 m., 28 June
2004, N. Wannathes 185 (CMU, SFSU); Chiang Rai
Province, Muang District, Khun Con Waterfall, 12 June
2005, N. Wannathes 304 (CMU, SFSU).
263
Discussion: Marasmius luteolus is
characterized in forming striate to sulcate pilei
colored pale cream-orange to yellowish orange
with bright brownish orange to golden orange
disc, subdistant, narrow lamellae, a relatively
short (10-35 mm), pruinose stipe covered with
two types of caulocystidia (Siccus-type broom
cells and non-setulose cells), medium-sized
basidiospores, and growth on monocotyledonous and dicotyledonous debris. A
distinctive feature of M. luteolus are
pleurocystidia of Siccus-type broom cells, a
feature rare in Marasmius. The type was
collected on palm leaves in the Bonin Islands
(isotype FH!) and this is the first report of the
species outside of Japan. Marasmius luteolus is
similar to M. setulosifolius Singer, another
species with Siccus-type pleurocystidia, but the
latter species differs in lacking simple, nonsetulose caulocystidia (Singer, 1976).
flavo-brunneis usque brunneis, crassetunicatis. Trama
pilei intertexta, fortiter dextrinoidea. Trama lamellae
intertexta, hyphis (4-)8-15(-25) µm diametro, cylindratis
usque inflatis, laevibus, hyalinis, fortiter dextrinoideis,
tenuitunicatis, haud gelatinosis. Stipitipellis parallelus,
hyphis (3-)5-8 µm diametro, cylindratis, flavo-brunneis
usque brunneis, laevibus, dextrinoideis, crassetunicatis
(usque ad 1 µm), haud gelatinosis. Trama stipitis
parallela, hyphis 4-8 µm diametro, cylindratis, hyalinis,
laevibus, dextrinoideis, tenuitunicatis, haud gelatinosis.
Caulocystidia 2 typorum: a) cellulis typi Sicci (5-)15-25
× 4-7 µm, dispersis, raris, cylindratis vel adumbratim
inaequabilibus, hyalinis; setulis apicalibus (2-)4-11 × 11.5 µm, cylindratis usque conicis vel undulatis, pallide
flavis, tenui- usque crassetunicatis; b) cellulis haud
setulosis, abundantia, 11-42 × 4-10 µm, cylindratis vel
adumbratim inaequabilibus, raro ramosis, obtusis usque
subacutis, hyalinis, inamyloideis, tenui- usque
crassetunicatis (usque ad 1 µm). Fibulae praesentes in
omnibus texturis.
Typus: Chiang Mai Province, Doi Suthep-Pui
National Park, Mokfa Waterfall, on Hwy 1095, N19º
6.5’.E 98º 46.3’ alt. 1014 m., dispersus usque gregarius
in folis plantae dicotyledoneae vel ligno, 1 August 2003,
N. Wannathes 067 (CMU: holotypus; SFSU: isotypus).
37. Marasmius jasminodorus Wannathes,
Desjardin & Lumyong, sp. nov.
(Figs 43, 49-4)
MycoBank: 512421
Pileus 10-40 mm diam., broadly convex
to campanulate, sometimes with a small umbo,
pruinose, dull; disc rugulose, dark reddish
brown (7-8D6-8); margin rugulose-striate, light
brown to brownish orange (7C6-8). Context
yellowish white, thin. Lamellae narrowly
adnexed, subdistant (12-17) with 3-4 series of
lamellulae, broad (2-5 mm), pale yellowish
white, non-marginate or with pale brownish
orange edges, non-intervenose. Stipe 20-60 ×
1-2 mm, central, cylindrical, tough, hollow,
glabrous to minutely velutinous, non-insititious,
base with strigose, brownish orange mycelium;
apex pale yellowish white (4A2), base brown
(7E7-8) to reddish brown (8E7-8) or dark
reddish brown (8F5-8). Odor strong, sweet and
fragrant, like jasmine tea. Taste mildly bitter.
Basidiospores 9-12(-14) × 3-4(-4.5) µm
[xmr = 9.4-11.5 × 3.1-4.1 µm, xmm = 10.1 ± 0.5
× 3.6 ± 0.3 µm, Qmr = 2.6-3.4, Qmm = 2.8 ± 0.1,
n = 25 spores, s = 18 specimens], ellipsoid,
curved in profile, smooth, hyaline, inamyloid,
thin-walled. Basidia 23-25 × 5-6 µm,
cylindrical to clavate, 4-spored. Basidioles
fusoid to clavate. Cheilocystidia common, of
Siccus-type broom cells; main body 9-19(-26)
× 6-10 µm, cylindrical to clavate, hyaline to
pale yellow, inamyloid, thin- to thick-walled;
apical setulae (2-)4-9(-11) × 1-2 µm, cylindrical or irregular in outline, obtuse to subacute,
yellow to brownish yellow, thick-walled.
Etymology: ‘jasmin’ = jasmine, ‘odorus’ = odor,
referring to the jasmine tea odor of the basidiomes.
Pileus 10-40 mm diametro, late convexus usque
campanulatus, interdum umbone parvo, pruinosus,
hebetatus, disco ruguloso, rubro-atrobrunneo, margine
rugoloso-striato, laete brunneo usque brunneo-aurantiaco.
Contextus flavo-albus, tenuis. Lamellae annexae,
subdistantes (12-17) cum 3-4 seriebus lamellularum,
latae (2-5 mm), pallide flavo-albae, haud marginatae vel
pallide brunneo-aurantiacomarginatae, haud intervenosae.
Stipes 20-60 × 1-2 mm, centralis, cylindratus, tenax,
cavus, glaber usque minute velutinus, haud insititius,
mycelio strigosum et brunneo-aurantiacum ad basim,
apice pallide flavo-albo, basi brunnea usque rubrobrunnea vel rubro-atrobrunnea. Odor fortis, suavis et
fragrans, iasmino sicco similis. Sapor leviter amarus.
Basidiosporae 9-12(-14) × 3-4(-4.5) µm, ellipsoideae,
oblique curvae, laeves, hyalinae, inamyloideae,
tenuitunicatae Basidia 23-25 × 5-6 µm, cylindrata usque
clavata, 4-spora. Basidiolae fusoideae usque clavatae.
Cheilocystidia vulgaria, typi Sicci; 9-19(26) × 6-10 µm,
cylindrata usque clavata, hyalina usque pallide flava,
inamyloidea, tenui- usque crassetunicata; setulis
apicalibus (2-)4-9(-11) × 1-2 µm, cylindratis vel
adumbratim inaequabilibus, obtusis usque subacutis,
flavis usque brunneo-flavis, crassetunicatis. Pleurocystidia nulla. Pileipellis hymeniformis, maculosus, typi
Sicci; 10-23(-29) × 6-9(-15) µm, clavatus usque late
clavatus, saepe ramosus, hyalinus, inamyloideus, tenuiusque crassetunicatus; setulis apicalibus (2-)4-9(-15) ×
1-2(-3) µm, coartatis, cylindratis vel adumbratim
inaequabilibus, saepe ramosis, obtusis usque subacutis,
264
Fungal Diversity
Pleurocystidia absent. Pileipellis hymeniform,
mottled, composed of Siccus-type broom cells;
main body 10-23(-29) × 6-9(-15) µm, clavate
to broadly clavate, often branched, hyaline,
inamyloid, thin- to thick-walled; apical setulae
(2-)4-9(-15) × 1-2(-3) µm, crowded, cylindrical
to irregular in outline, often branched, obtuse
to subacute, yellowish brown to brown, thickwalled. Pileus trama interwoven, strongly
dextrinoid. Lamellar trama hyphae (4-)8-15(25) µm diam., interwoven, cylindrical to
inflated, smooth, hyaline, strongly dextrinoid,
thin-walled,
non-gelatinous.
Stipitipellis
hyphae (3-)5-8 µm diam., parallel, cylindrical,
yellowish brown to brown, smooth, dextrinoid,
thick-walled (up to 1 µm), non-gelatinous.
Stipe trama hyphae 4-8 µm diam., parallel,
cylindrical, hyaline, smooth, dextrinoid, thinwalled, non-gelatinous. Caulocystidia composed of two types of cells: a) Siccus-type
broom cells with main body (5-)15-25 × 4-7
µm, scattered, uncommon, cylindrical or
irregular in outline, hyaline; apical setulae (2)4-11 × 1-1.5 µm, cylindrical to conical or
wavy, pale yellow, thin- to thick-walled; b)
abundant non-setulose cells, 11-42 × 4-10 µm,
cylindrical or irregular in outline, seldom
brunched, obtuse to subacute, hyaline,
inamyloid, thin- to thick-walled (up to 1 µm).
Clamp connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves or on wood, Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mokfa
Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
m., 28 June 2003, D.E. Desjardin 7589 (CMU, SFSU);
same location, 25 July 2003, N. Wannathes 028(CMU,
SFSU); same location, 1 August 2003, N. Wannathes
067 (CMU: holotype; SFSU: isotype); same location, 29
June 2004, Y.S. Tan 283(CMU, SFSU); same location, 2
August 2004, N. Wannathes 219 (CMU, SFSU); Chiang
Mai Province, Mae Taeng District, “Water Conservation
Area” Highway 1,095 at 22 km marker, N. 19º 7.5’ E 98º
45.7’ alt. 724 m., 2 July 2003, D.E. Desjardin 7604
(CMU, SFSU); same location, 4 August 2003, N.
Wannathes 084 (CMU, SFSU); same location, 2 June
2004, N. Wannathes 169 (CMU, SFSU); Chiang Mai
Province, Mae Taeng District, Mushroom Research
Centre, 27 km marker on Hwy 1095, 9 June 2004, N.
Wannathes 151 (CMU, SFSU); Chiang Mai Province,
Doi Inthanon National Park, At 25 km marker on
Highway 1,009, N 18º 32.5’ E 98º 33.5’ alt. 1,076 m., 10
June 2004, N. Wannathes 156 (CMU, SFSU); same
Fig. 43 Marasmius jasminodorus (N. Wannathes067) 1.
Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5. Pileipellis 6a. Siccus-type caulocystidia
6b. Caulocystidia, Scale bar 1 = 20 mm, 2-6 = 10 µm
location,25 June 2004, N. Wannathes 172 (CMU, SFSU);
same location, 27 June 2005, N. Wannathes 365 (CMU,
SFSU); Chiang Mai Province, Doi Inthanon National
Park Junction of Highway 1,009 and road to Mae Chaem,
N 18º 31.6’ E 98º 29.6’ alt. 1,703 m., 3 July 2004, N.
Wannathes 210 (CMU, SFSU); Chiang Mai Province,
Doi Suthep-Pui National Park, trial to 9 channel station,
24 July 2004, N. Wannathes 238 (CMU, SFSU); Chiang
Rai Province, Khun Chae National Park, N 19º 4.4' E 99º
23.5' alt. 963 m., 10 June 2005, N. Wannathes 287
(CMU, SFSU); Chiang Rai Province, Muang District,
Doo Village, Pong Prabath Waterfall, N 19º 53.2' E 100º
07.3' alt. 408 m., 11 Jume 2005, N. Wannathes 294
(CMU, SFSU);Chiang Mai Province, Doi Suthep-Pui
National Park, the trail opposite with the way to Huai
Kok Ma Village, 2 July 2005, N. Wannathes 387 (CMU,
SFSU); Chiang Mai Province, Mae Taeng District, Sri
Lanna National Park, near Mae Ngad Dam, 9 August
2005 , N. Wannathes 404 (CMU, SFSU); Phrae Province,
Muang District, Cherng Thong Waterfall, 16 August
2005, N. Wannathes 414 (CMU, SFSU).
Discussion: Distinctive features of this
new species include the following: a rugulosestriate pileus colored light brown to brownish
265
orange on the margin with dark reddish brown
disc; subdistant (12-17), broad, pale yellowish
white lamellae; basidiospores with mean 10.1 ×
3.6 µm; and two types of caulocystidia, Siccustype broom cells and simple cylindrical nonsetulose cells. A most remarkable feature is the
strong, sweet and fragrant odor like jasmine tea,
unknown in any other Marasmius species.
Marasmius jasminodorus is very similar to M.
araucariae Singer described from material
collected in Argentina, but the latter species
differs in forming longer subcylindrical
caulocystidia, more numerous lamellae (16-24)
and lacks Siccus-type broom cells on the stipe
surface and lacks a fragrant odor (Singer, 1976).
Marasmius jasminodorus is also phenetically
similar to M. araucariae var. siccipes described
from Java, but the latter species differs in
forming a more brownish orange pileus that is
not rugulose overall, darker coloured lamellae
(brownish orange to greyish brown) and lacks a
fragrant odor (Desjardin et al,. 2000).
38A. Marasmius araucariae Singer var.
araucariae, Sydowia 18: 333. 1965. (Fig. 44)
Pileus 9-34 mm diam., convex to planoconvex, slightly depressed in age, pruinose,
dull; disc rugulose, reddish brown; margin
striate, brownish orange. Context off-white,
thin. Lamellae narrowly adnate, subdistant (1415) with 3-4 series of lamellulae, broad (1-3
mm), cream, non-marginate, non-intervenose.
Stipe 30-45 × 1-1.2 mm, central, cylindrical,
hollow, subvelutinous, non-insititious, apex
off-white, base brown. Odor and taste not
distinctive.
Basidiospores 9-11(-12) × 3-4 µm [xm =
10.5 ± 0.8 × 3.9 ± 0.3 µm, Q = 2.3-3.1, Qm =
2.7, n = 25 spores, s = 1 specimen], ellipsoid,
curved in profile, smooth, hyaline, inamyloid,
thin-walled. Basidia not observed. Basidioles
fusoid to clavate. Cheilocystidia common, of
Siccus-type broom cells; main body 11-15 × 57 µm, subcylindrical to clavate or irregular in
outline, seldom lobed, hyaline, inamyloid, thinwalled; apical setulae 3-8 × 1-2 µm, cylindrical
to conical, sometimes wavy in outline, obtuse
to subacute, yellow to brownish yellow, thinwalled. Pleurocystidia absent. Pileipellis
hymeniform, mottled, composed of Siccus-type
broom cells; main body 12-21 × 7-12 µm,
266
Fig. 44. Marasmius araucariae var. araucariae (N.
Wannathes 300) 1. Basidiomes 2. Basidiospores 3.
Cheilocystidia 4. Pileipellis 5. Caulocystidia, Scale bar 1
= 20 mm, 2-5 = 10 µm
clavate to broadly clavate, turbinate or irregular
in outline, hyaline, inamyloid, thin- to thickwalled; apical setulae 3-13 × 1-1.5 µm,
crowded, cylindrical to irregular in outline,
obtuse to subacute, brownish yellow, thickwalled. Pileus trama interwoven, dextrinoid.
Lamellar trama hyphae 4-10(-16) µm diam.,
interwoven, cylindrical, smooth, hyaline,
dextrinoid,
thin-walled,
non-gelatinous.
Stipitipellis hyphae 4-8 µm diam., parallel,
cylindrical,
yellowish
brown,
smooth,
dextrinoid, thin- to thick-walled (up to 1 µm),
non-gelatinous. Stipe trama hyphae 6-13 µm
diam., parallel, cylindrical, hyaline, smooth,
dextrinoid,
thin-walled,
non-gelatinous.
Caulocystidia numerous, 24-60 × 5-8 µm,
polymorphic, cylindrical to clavate or irregular
in outline, obtuse, hyaline, inamyloid, thin- to
thick-walled (up to 1.5 µm); Siccus-type broom
cells absent on stipe surface. Clamp
connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious, often in cespitose
clusters on dicotyledonous leaves. Argentina,
Northern Thailand, Papua New Guinea.
Fungal Diversity
Material examined: Thailand. Chiang Rai
Province, Muang District, Khun Con Waterfall, 12 June
2005, N. Wannathes 300 (CMU, SFSU).
Discussion: Marasmius araucariae is
charaterized by rugulose pilei coloured reddish
brown to brownish orange, subdistant (14-15),
broad, cream-coloured and non-marginate
lamellae, a brown, subvelutinous stipe that
often develops in cespitose clusters, basidiospores with mean 10.5 × 3.9 µm, simple,
cylindrical caulocystidia, and an absence of
pleurocystidia. The Thai specimen matches
nicely with the material that was reported from
Papua New Guinea (Desjardin & Horak, 1997).
It is similar to M. araucariae var. siccipes (see
below) except for lacking Siccus-type broom
cells on the stipe surface. Marasmius
araucariae is also phenetically similar to M.
iras Y.S. Tan & Desjardin described from
Malaysia, but the latter species differs in
forming larger basidiospores with mean 14.1 ×
3.7 µm, and broader caulocystidia (6-10 µm
diam.).
38B. Marasmius araucariae var. siccipes
Desjardin, Retn. & E. Horak, Sydowia 52: 173.
2000.
(Figs 45, 49-4)
Pileus 10-33 mm diam., convex to
campanulate when young, expanding to planoconvex with upturned margin in age, striate to
subsulcate, pruinose, dull; disc dark brown
(7F7-9F4), margin brown (6E8) to brownish
orange (6C6). Context thin, greyish orange
(6B4) to light brown (6D5). Lamellae adnexed
to adnate, subdistant to close (13-20) with 3-4
series of lamellulae, broad (2-4 mm), brownish
orange (5C4) to greyish brown, non-marginate,
non-intervenose. Stipe 13-82 × 1-2 mm, central,
cylindrical, hollow, pruinose to subglabrous,
non-insititious, brown to dark brown overall.
Odor and taste not distinctive.
Basidiospores (8-)9-12 × 3.5-4 µm [xmr =
9.8-11.3 × 3.3-4.0 µm, xmm = 10.4 ± 0.8 × 3.6 ±
0.4 µm, Qmr = 2.8 - 3.1, Qmm = 2.9 ± 0.2, n =
25 spores, s = 3 specimens], ellipsoid, curved
in profile, smooth, hyaline, inamyloid, thinwalled. Basidia not observed. Basidioles fusoid
to clavate. Cheilocystidia common, of Siccustype broom cells; main body 11-18(-28) × 5-9
µm, subcylindrical to clavate or irregular in
outline, seldom lobed, hyaline, inamyloid, thinwalled; apical setulae 3-8 × 1-2 µm, cylindrical
Fig. 45. Marasmius araucariae var. siccipes (N.
Wannathes 364) 1. Basidiomes 2. Basidiospores 3.
Cheilocystidia 4. Pleurocystidia 5. Pileipellis 6a. Siccustype caulocystidia 6b. Caulocystidia, Scale bar 1 = 20
mm, 2-6 = 10µm
to conical, sometimes wavy in outline, obtuse
to subacute, light brown to brownish yellow,
thin-walled. Pleurocystidia absent. Pileipellis
hymeniform, mottled, composed of Siccus-type
broom cells; main body (9-)14-26 × 7-10 µm,
clavate to broadly clavate, turbinate or irregular
in outline, hyaline, inamyloid, thin- to thickwalled; apical setulae (2-)4-10 × 1-2 µm,
crowded, cylindrical to irregular in outline,
obtuse to subacute, brown to dark brown,
thick-walled. Pileus trama interwoven,
dextrinoid. Lamellar trama hyphae (3-)5-12
µm diam., interwoven, cylindrical, smooth,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Stipitipellis hyphae 4-10 µm diam., parallel,
cylindrical, yellowish brown to brown, smooth,
dextrinoid, thick-walled (up to 1.5 µm), nongelatinous. Stipe trama hyphae 6-14 (-20) µm
diam., parallel, cylindrical, hyaline, smooth,
dextrinoid,
thin-walled,
non-gelatinous.
Caulocystidia composed of two types of cells:
a) Siccus-type broom cells with main body 312 × 4-7 µm, scattered, uncommon, cylindrical
or irregular in outline, hyaline; apical setulae 411 × 1-1.5 µm, cylindrical to conical or wavy,
pale-yellow, thin-walled; b) abundant non267
setulose cells, 13-40 × 4-6 µm, polymorphic,
cylindrical to clavate or irregular in outline,
sometimes branched, obtuse, hyaline, inamyloid, thin- to thick-walled (up to 1 µm). Clamp
connections present in all tissues.
Habit, habitat and known distribution:
Scattered to solitary on dicotyledonous leaves,
Java, Malaysia and Northern Thailand.
Material examined: Thailand, Chiang Mai
Province, Mae Taeng District, Tung Joaw Village, N 19º
8.07’ E 98º 38.9’ alt. 1423 m, 30 August 2003, N.
Wannathes 146 (CMU, SFSU); Chiang Mai Province,
Doi Inthanon National Park, At 25 km marker on
Highway 1,009, N 18º 32.5’ E 98º 33.5’ alt. 1,076 m, 25
June 2004, N. Wannathes 173 (CMU, SFSU); same
location, 27 June 2005, N. Wannathes 364 (CMU,
SFSU).
cylindrical to clavate, broadly clavate or
irregular in outline, often lobed, hyaline,
inamyloid, thin-walled; apical setulae 3-10 × 11.5 µm, cylindrical, obtuse to subacute, pale
yellow to yellow, thick-walled. Pleurocystidia
absent. Pileipellis hymeniform, weakly mottled
to non-mottled, composed of Siccus-type
broom cells; main body (9-)12-24 × 5-11µm,
clavate to broadly clavate, often branched,
hyaline, inamyloid, thin-walled; apical setulae
2-5 × 1-1.5 µm, crowded, cylindrical, obtuse to
subacute, yellowish brown, thick-walled.
Pileus trama interwoven, dextrinoid. Lamellar
trama hyphae 3-10 µm diam., interwoven,
Discussion: This variety described by
Desjardin et al. (2000) from Java differs from
the type variety in forming numerous Siccustype broom cells on the stipe surface amongst
the simple, cylindrical caulocystidia. The Thai
populations are micromorphologically indistinguishable from the populations in Malaysia
and Java, and differ only subtly in forming
darker brown pilei with brown orange to
greyish brown lamellae. At present, these color
difference do not appear significant enough to
recognize the Thai entity as a distinct taxon.
39. Marasmius xestocephalus Singer, Bull.
Jard. Bot. Etat Brux. 34: 367. 1964. (Fig. 46)
Pileus 5-23 mm diam., convex to planoconvex, umbonate, striate to pellucid-striate,
pruinose, dull, disc yellowish brown (6D8),
margin greyish yellow (4B4-5) to yellowish
orange (4B7). Context pale yellow, thin.
Lamellae adnexed to adnate, subdistant (15-18)
with 3-4 series of lamellulae, narrow, pale
yellow with or without light orangish brown
edges, non-intervenose. Stipe 18-35 × 0.5-1
mm, central, cylindrical, hollow, pubescent to
subvelutinous, non-insititious, apex light
yellow (3A4), base light brown (6D6) to
reddish brown (8E7). Odor and taste not
distinctive.
Basidiospores (10-)11-14(-15) × 3-4 µm
[xmr = 11.8-13 × 3.5-3.8 µm, xmm = 12.5 ± 0.6
× 3.7 ± 0.2 µm, Qmr = 3.4, Qmm = 3.4 ± 0.0, n =
25 spores, s = 3 specimens], ellipsoid to fusoid,
smooth, hyaline, inamyloid, thin-walled.
Basidia not observed. Basidioles fusoid to
clavate. Cheilocystidia abundant, of Siccustype broom cells; main body 10-18 × 6-9 µm,
268
Fig. 46. Marasmius xestocephalus (N. Wannathes 344) 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pileipellis 5. Caulocystidia, Scale bar 1 = 20 mm, 2-5 =
10 µm
cylindrical, smooth, hyaline, dextrinoid, thinwalled, non-gelatinous. Stipitipellis hyphae
(2-)4-8 µm diam., subparallel, cylindrical,
brown to yellowish brown, smooth, weakly
dextrinoid, thick-walled (up to 1 µm), nongelatinous. Stipe trama hyphae 3-8(-10) µm
diam., subparallel, cylindrical, hyaline, smooth,
dextrinoid, thin-walled, non-gelatinous. Caulocystidia numerous, usually apically forked;
Fungal Diversity
main body 25-54 × 6-10 µm, cylindrical to
irregular in outline, hyaline to pale yellow,
weakly dextrinoid, thin- to thick-walled (up to
1 µm); apical fork 10-20 × 4-7 µm, cylindrical,
obtuse, hyaline to pale yellow, thin- to thickwalled; Siccus-type broom cells absent on stipe
surface. Clamp connections present in all
tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, Africa and Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Mae Taeng District, “Water Conservation
Area” Highway 1,095 at 22 km marker, N. 19º 7.5’ E 98º
45.7’ alt. 724 m., 5 July 2004, Y.S. Tan 313 (CMU,
SFSU); Chiang Mai Province, Doi Suthep-Pui National
Park, Mokfa Waterfall, on Hwy 1095, N19º 6.5’.E 98º
46.3’ alt. 1014 m., 7 July 2004, J.F. Kerekes 69 (CMU,
SFSU); Chiang Mai Province, Mae Taeng District, Mae
Sae Village, Pang Sa Ded Water Conservation Area, N
19º 14.6', E 98º 38.5' alt. 962 m., 26 June 2005, N.
Wannathes 344 (CMU, SFSU).
Discussion: Diagnostic features of
Marasmius xestocephalus include: a greyish
yellow to yellowish orange pileus with
yellowish brown disc; subdistant (15-18),
narrow, pale yellow lamellae; a pubescent to
subvelutinous stipe; fusoid basidiospores with
mean 12.5 × 3.7 µm; and apically forked
caulocystidia.
The
Thai
material
is
indistinguishable from the protologue (Singer,
1964) and redescription of M. xestocephalus by
Pegler, (1977), and differs only slightly from
the recent description provided by Antonín
(2007) who reported broader basidiospores
(4.5-5.5 µm diam.). Until further material of
this species is collected, sequenced and
compared with Thai specimens, we tentatively
recognize our species as M. xestocephalus.
40. Marasmius iras Y.S. Tan & Desjardin
Fungal Diversity 37: 61, 2009 (Figs 47, 49-6)
Pileus 5-20 mm diam., convex when
young, expanding to plano-convex with
slightly depressed disc in age, striate to sulcate,
pruinose, dull, dark yellowish brown (5F8-5E6)
to dark brown (6E8). Context yellowish brown
(5E8), thin. Lamellae adnexed to adnate,
subdistant (12-17) with 3 series of lamellulae,
narrow, cream brown to brownish orange (5C5)
with brown edges, non-intervenose. Stipe 1743 × 0.5-1 mm, central, cylindrical, hollow,
subvelutinous, non-insititious, apex yellowish
white (3A2), base reddish brown (8E7). Odor
and taste not distinctive.
Basidiospores (11-)12-14(-15) × 3-4(-5)
µm [xmr = 12.6-13.4 × 3.4-4.2, xmm = 12.8 ± 0.5
× 3.7 ± 0.4 µm, Qmr = 3.0-3.7, Qmm = 3.5 ± 0.4,
n = 25 spores, s = 3 specimens], fusiod, smooth,
hyaline, inamyloid, thin-walled. Basidia not
observed. Basidioles fusoid to clavate.
Cheilocystidia common, of Siccus-type broom
cells; main body (8-)13-21 × 5-7(-10) µm,
cylindrical to clavate or irregular in outline,
seldom branched, hyaline, inamyloid, thinwalled; apical setulae (2-)4-9(-11) × 1-1.5(-2)
µm, cylindrical, obtuse to subacute, yellow to
brown, thick-walled. Pleurocystidia absent.
Pileipellis hymeniform, mottled to weakly
mottled, composed of Siccus-type broom cells;
main body 12-25 × 5-9 µm, cylindrical to
clavate or turbinate, seldom branched, hyaline,
inamyloid, thin-walled; apical setulae 3-8 × 1
µm, crowded, cylindrical to irregular in outline,
obtuse to subacute, dark brown to black, thickwalled. Pileus trama interwoven, dextrinoid.
Lamellar trama hyphae 3-8(-10) µm diam.,
interwoven, cylindrical, smooth, hyaline,
dextrinoid, thin-walled, non-gelatinous. Stipitipellis hyphae 3-7(-10) µm diam., subparallel,
cylindrical, yellow to brown, smooth,
inamyloid to weakly dextrinoid, thick-walled
Fig. 47. Marasmius iras (N. Wannathes 276) 1.
Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5. Pileipellis 6. Caulocystidia, Scale bar 1
= 20 mm, 2-6 = 10 µm
269
(up to 1 µm), non-gelatinous. Stipe trama
hyphae 4-10(-13) µm diam., parallel, cylindrical, hyaline, smooth, dextrinoid, thin-walled,
non-gelatinous. Caulocystidia numerous, 23-43
× 6-10 µm, cylindrical to clavate or irregular in
outline, rarely lobed, hyaline, inamyloid, thinwalled; Siccus-type broom cells absent on stipe
surface. Clamp connections present in all
tissues.
Habit, habitat and known distribution:
Scattered to solitary on dicotyledonous leaves,
Malaysia and Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Mae Taeng District, Tung Joaw Village, N 19º
8.07’ E 98º 38.9’ alt. 1423 m., 30 August 2003, N.
Wannathes 147 (CMU, SFSU); same location, 22
August 2004, N. Wannathes 276 (CMU, SFSU); Chiang
Mai Province, Huai Nam Dang National Park, North 19º
18.3’ E 98º 35.8’ alt. 1,538 m., 29 June 2005, N.
Wannathes 375 (CMU, SFSU).
Discussion: Marasmius iras, described
from specimens collected in Malaysia, is
characterized by a yellowish brown pileus with
dark brown disc, subdistant, yellowish white
lamellae, a subvelutinous stipe, fusoid
basidiospores, non-setulose caulocystidia, and
an absence of pleurocystidia and Siccus-type
broom cells on the stipe surface. The Thai
specimens match quite closely the Malaysian
type except for having slightly smaller
basidiospores with mean 12.8 × 3.7 µm vs.
14.1 × 3.7 µm, and longer setulae of
cheilocystidia (4-11µm vs 4-7 µm long).
Marasmius iras is phenetically similar to M.
xestocephalus (see above), but the latter
species differs in forming pilei with yellow
tones and apically forked, thicker-walled
caulocystidia.
41. Marasmius inthanonensis Wannathes,
Desjardin & Lumyong, sp. nov.
(Figs 48, 49-7)
MycoBank: 512422
Etymology: ‘inthanon’ referring to Doi Inthanon
National Park, the site where the holotype specimen was
collected.
Pileus 7-42 mm diametro, convexus ubi iuvenis,
expansus et plano-convexus, disco leviter depresso et
margine reflexo ubi vetus, laevis usque striatus,
pruinosus, hebetatus, primo olivaceo-brunneus demum
brunneo-aurantiacus usque flavo-brunneus. Contextus
pallide aurantiacus usque cinereo-brunneus, tenuis.
Lamellae annexae usque adnatae, arctae (12-20) cum 3-4
seriebus lamellularum, latae (3-6 mm), pallide flavae
usque flavo-cinereae, laete brunneomarginatae, haud
intervenosae. Stipes 40-102 × 1-3 mm, centralis,
270
cylindratus, cavus, subvelutinus usque hispidulus, haud
insititius, apice flavo-albo, basi rubro-atrobrunnea. Odor
saporque non propria. Basidiosporae 8-12 × 3-4 µm,
ellipsoideae usque fusoideae, oblique curvae, laeves,
hyalinae, inamyloideae, tenuitunicatae Basidia non
observata. Basidiolae fusoideae usque clavatae.
Cheilocystidia vulgaria, typi Sicci; 13-23 × 4-9 µm,
subcylindrata usque clavata vel adumbratim inaequabilia,
raro ramosa, hyalina, inamyloidea, tenuitunicata; setulis
apicalibus 2-6 × 1-1.5 µm, cylindratis usque conicis, raro
ramosis, obtusis usque subacutis, hyalinis usque flavis,
tenuitunicatis.
Pleurocystidia
nulla.
Pileipellis
hymeniformis, maculosus, typi Sicci; 15-23 × 6-11 µm,
clavatus usque late clavatus, turbinatus vel adumbratim
inaequabilis, hyalinus, inamyloideus, tenui- usque
crassetunicatus; setulis apicalibus 3-5 × 1 µm, coartatis,
cylindratis vel adumbratim inaequabilibus, obtusis usque
subacutis, flavis usque brunneo-flavis, crassetunicatis.
Trama pilei intertexta, dextrinoidea. Trama lamellae
intertexta, hyphis (3-)9-13 µm diametro, cylindratis,
laevibus, hyalinis, dextrinoideis, tenuitunicatis, haud
gelatinosis. Stipitipellis subparallelus, hyphis 4-6(-9) µm
diametro, cylindratis, brunneis usque olivaceo-brunneis,
laevibus, inamyloideis usque leniter dextrinoideis,
crassetunicatis (usque ad 1 µm), haud gelatinosis. Trama
stipitis parallela, hyphis 4-13 µm diametro, cylindratis,
hyalinis, laevibus, dextrinoideis, tenuitunicatis, haud
gelatinosis. Caulocystidia numerosa, 24-48 × 6-8 µm,
cylindrata,
adumbratim
inaequabilia,
hyalina,
inamyloidea, tenui- usque crassetunicata; sine cellulis
type Sicci super superficiem stipitis. Fibulae praesentes
in omnibus texturis.
Typus: Chiang Mai Province, Doi Inthanon
National Park Junction of Highway 1,009 and road to
Mae Chaem, N 18º 31.6’ E 98º 29.6’ alt. 1,703 m.,
dispersus usque gregarius in folis plantae dicotyledoneae,
27 June 2005, N. Wannathes 353 (CMU: holotypus;
SFSU: isotypus).
Pileus 7-42 mm diam., convex when
young, expanding to plano-convex with a
slightly depressed disc and reflexed margin in
age, smooth to striate, pruinose, dull, at first
olive brown overall, fading through brownish
orange (5C4-5) to yellowish brown (5E4-5) in
age. Context pale orange (5A3) to greyish
brown (5E3), thin. Lamellae adnexed to adnate,
close (12-20) with 3-4 series of lamellulae,
broad (3-6 mm), pale-yellow to yellowish grey
(3B2) with light brown edges, non-intervenose.
Stipe 40-102 × 1-3 mm, central, cylindrical,
hollow, subvelutinous to hispidulous, noninsititious, apex yellowish white, base dark
reddish brown. Odor and taste not distinctive.
Basidiospores 8-12 × 3-4 µm [xmr = 9.810.9 × 3.9-4.0, xmm = 10.3 ± 0.8 × 3.9 ± 0.1 µm,
Qmr = 2.5 - 2.7, Qmm = 2.6 ± 0.1, n = 25 spores,
s = 2 specimens], ellipsoid to fusoid, curved in
profile, smooth, hyaline, inamyloid, thin-
Fungal Diversity
walled. Basidia not observed. Basidioles fusoid
to clavate. Cheilocystidia common, of Siccustype broom cells; main body 13-23 × 4-9 µm,
subcylindrical to clavate or irregular in outline,
seldom branched, hyaline, inamyloid, thinwalled; apical setulae 2-6 × 1-1.5 µm,
cylindrical to conical, seldom branched, obtuse
to subacute, hyaline to yellow, thin-
merous, 24-48 × 6-8 µm, cylindrical, irregular
in outline, hyaline, inamyloid, thin- to thickwalled; Siccus-type broom cells absent on stipe
surface. Clamp connections present in all
tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, Northern Thailand.
Fig. 48. Marasmius inthanonensis (N. Wannathes 353) 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pileipellis 5. Caulocystidia, Scale bar 1 = 20 mm, 2-5 =
10 µm
Discussion: Diagnostic features of
Marasmius inthanonensis include: a smooth
pileus colored olive brown when young and
fading thru brownish orange to yellowish
brown in age; close (12-20), broad (3-6 mm),
yellowish grey lamellae with light brown edges;
a subvelutinous to hispidulous stipe;
basidiospores with mean 10.3 × 3.9 µm; and
relatively short subcylindrical caulocystidia (up
to 48 µm long) and an absence of Siccus-type
broom cells on the stipe surface. This new
species is distinct from all other described
species in ser. Atrorubentes in having a pileus
with olive tones. Phenetically similar species
include M. auratus (No. 33), M. ochroleucus
(No. 35), and M. araucariae var. siccipes (No.
38B). Marasmius auratus differs in forming a
golden pileus that lacks olive tones, slightly
larger basidiospores with mean 12.2 × 4.1 µm,
only cylindrical cheilocystidia, and longer
caulocystidia (up to 112 µm long). Marasmius
ochroleucus differs primarily in forming a
paler pileus that is light yellow to cream and
lacks olive tones; basidiospores size and
cysitidia are indistinguishable. Marasmius
araucariae var. siccipes differs in lacking olive
tones on the pileus, and forms brown lamellae
and two types of caulocystidia. ITS sequence
data confirm the distinction of these four
species.
walled. Pleurocystidia absent. Pileipellis
hymeniform, mottled, composed of Siccus-type
broom cells; main body 15-23 × 6-11 µm,
clavate, to broadly clavate, turbinate or
irregular in outline, hyaline, inamyloid, thin- to
thick-walled; apical setulae 3-5 × 1 µm,
crowded, cylindrical to irregular in outline,
obtuse to subacute, yellow to brownish yellow,
thick-walled. Pileus trama interwoven,
dextrinoid. Lamellar trama hyphae (3-)9-13
µm diam., interwoven, cylindrical, smooth,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Stipitipellis hyphae 4-6(-9) µm diam.,
subparallel, cylindrical, brown to olive brown,
smooth, inamyloid to weakly dextrinoid, thickwalled (up to 1 µm), non-gelatinous. Stipe
trama hyphae 4-13 µm diam., parallel,
cylindrical, hyaline, smooth, dextrinoid, thinwalled, non-gelatinous. Caulocystidia nu-
Material examined: Thailand. Chiang Mai
Province, Doi Inthanon National Park Junction of
Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E
98º 29.6’ alt. 1,703 m., 25 June 2004, N. Wannathes 177
(CMU, SFSU); same location, 27 June 2005, N.
Wannathes 353 (CMU: holotype; SFSU: isotype).
Section Sicci, subsect. Siccini, ser. Leonini
Singer, Fl. Neotrop. Monogr. 17: 160. 1976.
Type: Marasmius leoninus Berk.
42. Marasmius cremeus Wannathes, Desjardin
& Lumyong, Mycol. Res. 111: 993. 2007.
(Figs 50, 57-1)
271
Fig. 49. Basidiomata of Marasmius section Sicci ser. Atrorubentes. 1. M. auratus (N. Wannathes 351) 2. M. luteolus (N.
Wannathes 304) 3. M. pseudopellucidus (N. Wannathes 305) 4. M. jasminodorus (N. Wannathes 414) 5. M. araucariae
var. siccipes (N. Wannathes 364) 6. M. iras (N. Wannathes276) 7. M. inthanonensis (N. Wannathes 353), Scale bars =
20 mm
272
Fungal Diversity
Pileus 3-14 mm diam., obtusely conical
when young, expanding to convex, planoconvex or applanate with or without a broad,
rugulose umbo in age; margin striate to sulcate;
surface dull, dry, glabrous; cream (4A3) overall
when young, in age disc cream (4A3) to light
yellow (4A2), margin white,
Fig. 50. Marasmius cremeus (N. Wannathes 118). 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pileipellis, Scale bar 1 = 20 mm, 2-4 = 10 µm
discoloring pale grayish orange in splotches.
Context thin, white. Lamellae adnate,
subdistant (8-15) with 0-1 series of lamellulae,
non-intervenose, narrow to moderately broad
(0.5-1.0 mm), light buff to cream-colored, nonmarginate. Stipe 11-30 × 0.2-0.5 mm, central,
cylindrical, terete, filiform, tough, pliant,
glabrous, non-insititious, base with short, tufted,
white to cream (4A3) basal tomentum;
yellowish white to dark brownish orange (7C57) at the apex, dark brown (7-8F4-6) to nearly
black at the base; rhizomorphs absent. Odor
and taste not distinctive.
Basidiospores (11-)12-14(-16) × 3-4(-5)
μm [xmr = 11.7-13.5 × 3.1-4.6 μm, xmm = 12.7 ±
0.6 × 3.8 ± 0.5 μm, Q = 1.8-5.0, Qmr = 2.7-4.0,
Qmm = 3.4 ± 0.5, n = 25 spores, s = 8
specimens], narrowly ellipsoid to fusoid,
smooth, hyaline, inamyloid, thin-walled.
Basidia 20-26 × 5-7 μm, clavate to cylindrical,
4-spored. Basidioles clavate to subfusoid.
Cheilocystidia common, of Siccus-type broom
cells; main body 9-19 × 6-12 μm, cylindrical to
pyriform, clavate, broadly clavate or irregular
in outline, hyaline, thin-walled; apical setulae
2-8(-11)× 0.5-1.0 μm, crowded, narrowly
cylindrical to cylindrical or irregular in outline,
rarely fork, subacute to acute, hyaline to pale
yellow, inamyloid, thin-walled. Pleurocystidia
absent. Pileipellis hymeniform, weakly mottled
to mottled, composed of Siccus-type broom
cells; main body (8-)12-21 × 7-16 μm, clavate
to broadly clavate, cylindrical, subglobose,
pyriform or turbinate, sometimes irregular in
outline, hyaline, inamyloid, thin- to thickwalled; apical setulae (2-)3-6(-11) × 0.5-1(-1.5)
μm, narrowly cylindrical to conical or irregular
in outline, often wavy, conical to subacute,
hyaline to pale yellow, inamyloid, thick-walled.
Pileus trama interwoven, lamellar trama
regular to interwoven; hyphae 3-8(-11) μm
diam, cylindrical to inflated, smooth, hyaline,
weakly dextrinoid to dextrionid, thin-walled,
non-gelatinous. Stipitipellis hyphae 2-5(-7) μm
diam, parallel, cylindrical, smooth, yellowish
brown to brown, dextrinoid, thin- to thickwalled (> 1 μm), non-gelatinous. Stipe trama
hyphae (2.5-)4-8(-10) μm diam, parallel,
cylindrical, smooth, hyaline, inamyloid to weak
dextrinoid, thin-walled. Caulocystidia absent.
Clamp connections present in all tissues.
Habit, habitat, and known distribution:
Scattered to gregarious, seldom solitary, on
undetermined dicotyledonous leaves. Northern
Thailand.
Material examined. Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mok Fa
waterfall, on Hwy 1095, N19º 6.581’, E98º 46.353’,
1014 meters, 1 Aug. 2003, N. Wannathes 069 (CMU,
SFSU); Chiang Mai Province, Mae Taeng District,
“Water Conservation Area” Highway 1,095 at 22 km
marker, N. 19º 7.5’ E 98º 45.7’ alt. 724 m., 4 August
2003, N. Wannathes 085 (CMU); same location, 13 June
2004, N. Wannathes 171 (CMU); Chiang Mai Province,
Doi Suthep-Pui National Park, Montathan Falls trail, 650
meters, 4 Jul. 2002, D.E. Desjardin 7455 (CMU, SFSU);
same location, 6 Jul. 2004, Y. S. Tan 320 (CMU, SFSU);
Chiang Mai Province, Doi Suthep-Pui National Park,
Sangasabhasri Land on the way to Huai Kok Ma Village,
N 18º 48.4’ E 98º 54.6’ alt. 1,146 m., 2 July 2005, N.
Wannathes 383 (CMU, SFSU); Chiang Mai Province,
Doi Inthanon National Park, at 25 km marker on
Highway 1009, N18º 32.54’, E98º 33.51’, 1076 meters,
10 Jun. 2004, N. Wannathes154 (CMU, SFSU), same
location, 27 June 2005, N. Wannathes 366 (CMU,
SFSU); Chiang Mai Province, Mae Taeng District, Pa
Deng Village, Pathummikaram temple, on Hwy 1095,
N19º 06' 28.8", E98º 44' 47.3", 1050 meters, 22 Aug.
2003, N. Wannathes 118 (CMU: holotype; SFSU:
isotype); same location, 7 Aug. 2003, N. Wannathes 093
(CMU, SFSU); Chiang Mai Province, Mushroom
Research Centre, 27 km marker on Hwy 1095, 27 Jul.
273
2003, N. Wannathes 042(CMU, SFSU); same location,
10 Aug. 2003, N. Wannathes 099 (CMU, SFSU).
Discussion: Marasmius cremeus is
distinguished by the following features: a
relatively small (3–14 mm diam), sulcate,
cream-colored pileus; subdistant (8–15),
narrow, cream-colored, non-collariate, nonmarginate lamellae; a filiform (0.2–0.5 mm
diam), glabrous, non-insititious, dark brown
stipe with white to cream-colored basal
tomentum; clavate spores with mean range
11.7–13.5 × 3.1–4.6 µm; no pleurocystidia and
no caulocystidia; Siccus-type cheilocystidia and
pileipellis cells with pale yellow setulae; and
growth on dicotyledonous leaves. The new
Thai species is most closely allied M. similis
Berk. & M.A. Curtis, a species described from
South Carolina, USA in 1849 and one that is
known only from the type specimen (K!;
isotype FH!). Marasmius similis is nearly
indistinguishable from M. cremeus, differing
only in forming a white pileus (not creamcolored) and in growing on twigs. A study of
the type specimen of M. similis shows no
differences
in
micromorphology
when
compared to the Thai specimens. However,
because M. similis is known from only a single
specimen collected on twigs in a temperate
forest in eastern North America, until more
material from that region is discovered and
compared with the Thai material, we choose to
recognize the Asian organism as a distinct
species. We conclude that M. cremeus is what
van Overeem and van Overeem-de Haas (1922)
reported as M. similis from Java (cf. Desjardin
et al. 2000:190).
Marasmius cremeus is also phenetically
similar to M. bellus Berk., a cream-colored to
yellow species from the neotropics, and to M.
haediniformis Singer, a pure white species
from Africa and Ecuador. The latter two
species differ from M. cremeus in forming
larger pilei (13–37 mm diam), thicker stipes
(1.0–1.5
mm),
and
slightly
smaller
basidiospores on average, ca 11.3 × 3.3 µm
(Singer 1976; Pegler 1977, 1983).
A bipolar mating system was reported for
M. cremeus by Wannathes et al. (2007).
43. Marasmius hypochroides Berk. &
Broome, J. Linn. Soc., Bot. 14: 35. 1873.
(Figs 51, 57-2)
274
Pileus 17-56 mm diam., convex to planoconvex or applanate, often subumbonate,
rugulose, striate to sulcate, glabrous to
subvelutinous, dull, disc brown (7E8) to dark
brown (7F7-8), margin brownish orange (5C5)
to yellowish brown (5E6). Context yellowish
white to cream, thin. Lamellae free to adnexed,
distant (12-16) with 3-4 series of lamellulae,
broad (2-10 mm), often intervenose, cream
(4B2) with brownish orange edges. Stipe 38-95
× 1-3 mm, central, cylindrical, wiry, hollow,
glabrous, non-insititious, apex buff, base
yellowish brown (5D8) to reddish brown (8E7).
Odor and taste not distinctive.
Basidiospores (8-)9-12(-13) × 5-6(-8)µm
[xmr = 8.9-10.8 × 5.1-6.2 µm, xmm = 9.9 ± 0.7 ×
5.6 ± 0.4 µm, Qmr = 1.6-2.0, Qmm = 1.8 ± 0.1, n
= 25 spores, s = 6 specimens], ellipsoid,
smooth, hyaline, inamyloid, thin-walled.
Basidia not observed. Basidioles cylindrical to
clavate. Cheilocystidia common, of Siccus-type
broom cells; main body (8-)10-25 × 4-10 µm,
cylindrical to clavate or pyriform, hyaline,
inamyloid, thin-walled; apical setulae 4-23(-32)
× 1-1.5 µm, cylindrical, inamyloid, subacute,
brown to light brown, thin- to thick-walled.
Pleurocystidia absent. Pileipellis hymeniform,
mottled, composed of Siccus-type broom cells;
main body 8-19(-26) × 5-11 µm, clavate to
broadly clavate or pyriform, often branched,
hyaline, inamyloid, thin- to thick walled; apical
setulae 3-11(-18) × 1-2 µm, crowded,
cylindrical, subacute to acute, brown to dark
brown, thick-walled. Pileus trama interwoven,
dextrinoid to weakly dextrinoid. regular to
interwoven, cylindrical, smooth, hyaline,
weakly dextrinoid to dextrinoid, thin-walled,
non-gelatinous. Stipitipellis hyphae 3-6 µm
diam., subparallel, cylindrical, light brown to
brown, smooth, dextrinoid, thin- to thickwalled, non-gelatinous. Stipe trama hyphae 712(-18) µm diam., subparallel, cylindrical,
hyaline, smooth, dextrinoid, thin-walled, nongelatinous. Caulocystidia absent. Clamp
connections present in all tissues.
Habit, habitat and known distribution:
Scattered to solitary on dicotyledonous leaves
or on wood. Java, Malaysia, Northern Thailand
and Sri Lanka.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mokfa
Fungal Diversity
MycoBank: MB512423
Fig. 51. Marasmius hypochoides (N. Wannathes 136). 1.
Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5. Pileipellis, Scale bar 1 = 20 mm, 2-5 =
10 µm
Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
m., 28 August 2003, N. Wannathes 136 (CMU, SFSU);
same location, 14 June 2004, N. Wannathes 160 (CMU,
SFSU); same location, 25 June 2005, N. Wannathes 335
(CMU, SFSU); Chiang Mai Province, Mae Taeng
District, Tung Joaw Village, N 19º 8.07’ E 98º 38.9’ alt.
1423 m., 30 June 2004, N. Wannathes 190 (CMU,
SFSU); Chiang Mai Province, Mushroom Research
Centre, 27 km marker on Hwy 1095, 28 July 2004, N.
Wannathes 247 (CMU); Chiang Mai Province, Mae
Taeng District, Sri Lanna National Park, near Mae Ngad
Dam, 9 August 2005, N. Wannathes 405 (CMU, SFSU).
Discussion: Marasmius hypochroides is
characterized by a relatively large, rugulose,
striate to sulcate pileus colored brownish
orange to yellowish brown with dark brown
disc, distant (12-16), broad (2-10 mm) lamellae,
moderately broad basidiospores with mean 9.9
× 5.6 µm, and mean Q = 1.8, cheilocystidia
with very long setulae (4-32 µm), and an
absence of pleurocystidia and caulocystidia.
The Thai specimens match nicely those from
Java reported by Desjardin et al. (2000). For a
comparison with similar species, refer to
Desjardin et al. (2000).
44.Marasmius makok Wannathes, Desjardin &
Lumyong, sp. nov.
(Figs 52, 57-3)
Etymology: ‘makok’ (Thai) = olive; referring to
the olive colour of the pileus margin.
Pileus 3-8 mm diametro, conicus, striatus,
pruinosus, hebetatus, disco cinereo-atrobrunneo, margine
olivacea. Contextus olivaceo-cinereus, tenuis. Lamellae
liberae, distantes (12), latae (1-2 mm), pallide cinereoalbae, haud marginatae, haud intervenosae. Stipes 28-45
× 0.3-0.5 mm, centralis, cylindratus, filo metallico
similis, glaber, haud insititius, apice flavo-albo, basi
atrorubinea. Odor saporque non propria. Basidiosporae
(14-)15-17 × 3-4 µm, anguste fusoideae usque clavatae,
oblique curvae, laeves, hyalinae, inamyloideae,
tenuitunicatae Basidia non observata. Basidiolae
fusoideae. Cheilocystidia vulgaria, typi Sicci; 13-22 × 79 µm, cylindrata usque clavata vel adumbratim
inaequabilia, hyalina, inamyloidea, tenuitunicata; setulis
apicalibus 2-4 × 1 µm, cylindratis usque conicis,
subacutis, hyalinis, tenuitunicatis. Pleurocystidia nulla.
Pileipellis hymeniformis, haud maculosus, typi Sicci; 1322 × 6-9 µm, cylindratus usque clavatus vel adumbratim
inaequabilis, hyalinus, inamyloideus, tenui- usque
crassetunicatus; setulis apicalibus 2-5 × 1 µm, coartatis,
cylindratis, subacutis, brunneis, crassetunicatis. Trama
pilei intertexta, dextrinoidea. Trama lamellae intertexta,
hyphis 3-8 µm diametro, cylindratis, laevibus, hyalinis,
dextrinoideis,
tenuitunicatis,
haud
gelatinosis.
Stipitipellis subparallelus, hyphis 3-4 µm diametro,
cylindratis,
brunneis,
laevibus,
inamyloideis,
crassetunicatis (usque ad 1.5 µm), haud gelatinosis.
Trama stipitis parallela, hyphis 3-7 µm diametro,
cylindratis,
hyalinis,
laevibus,
dextrinoideis,
tenuitunicatis, haud gelatinosis. Caulocystidia nulla.
Fibulae praesentes in omnibus texturis.
Typus: Chiang Mai Province, Doi Inthanon
National Park Junction of Highway 1,009 and road to
Mae Chaem, N 18º 31.6’ E 98º 29.6’ alt. 1,703 m.,
dispersus usque gregarius in folis plantae dicotyledoneae,
3 July 2004, N. Wannathes 201 (CMU: holotypus;SFSU:
isotypus).
Pileus 3-8 mm diam., conical, striate,
pruinose, dull, disc dark grayish brown (5F3),
margin olive (3E-F3-5). Context olive grey,
thin. Lamellae free, distant (12), broad (1-2
mm), pale greyish white, non-marginate, nonintervenose. Stipe 28-45 × 0.3-0.5 mm, central,
cylindrical, wiry, glabrous, non-insititious,
apex yellowish white, base dark ruby (12F4).
Odor and taste not distinctive.
Basidiospores (14-)15-17 × 3-4 µm [x =
15.8 ± 0.9 × 3.4 ± 0.4 µm, Q = 3.8-5.7, Qm =
4.7, n = 25 spores, s = 1 specimen], narrowly
fusoid to clavate, curved in profile, smooth,
hyaline, inamyloid, thin-walled. Basidia not
observed. Basidioles fusoid. Cheilocystidia
common, of Siccus-type broom cells; main
body 13-22 × 7-9 µm, cylindrical to clavate or
275
Fig. 52. Marasmius makok (N. Wannathes 201). 1.
Basidiomes 2. Basidioles 3. Basidiospores 4. Cheilocystidia
5. Pileipellis, Scale bar 1 = 20 mm, 2-5 = 10 µm
irregular in outline, hyaline, inamyloid, thinwalled; apical setulae 2-4 × 1 µm, cylindrical
to conical, subacute, hyaline, thin-walled.
Pleurocystidia absent. Pileipellis hymeniform,
not mottled, composed of Siccus-type broom
cells; main body 13-22 × 6-9 µm, cylindrical to
clavate or irregular in outline, hyaline,
inamyloid, thin- to thick-walled; apical setulae
2-5 × 1 µm, crowded, cylindrical, subacute,
brown, thick-walled. Pileus trama interwoven,
dextrinoid. Lamellar trama hyphae 3-8 µm
diam., interwoven, cylindrical, smooth, hyaline,
dextrinoid,
thin-walled,
non-gelatinous.
Stipitipellis hyphae 3-4 µm diam., subparallel,
cylindrical, brown, smooth, inamyloid, thickwalled (up to 1.5 µm), non-gelatinous. Stipe
trama hyphae 3-7 µm diam., parallel,
cylindrical, hyaline, smooth, dextrinoid, thinwalled, non-gelatinous. Caulocystidia absent.
Clamp connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Inthanon National Park Junction of
Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E
98º 29.6’ alt. 1,703 m., 3 July 2004, N. Wannathes 201
(CMU: holotype; SFSU: isotype)
276
Discussion: Marasmius makok is
distinguished by a conical, primarily olive
pigmented pileus with dark grayish brown disc,
distant, broad, pale greyish white lamellae, a
dark ruby red stipe, clavate basidiospores with
mean 15.8 × 3.4 µm, and an absence of
pleurocystidia and caulocystidia. The Thai new
species is phenetically similar to two olivecoloured neotropical species. Marasmius
digilioi Singer, described from Argentina,
differs in forming a larger, convex pileus,
subclose to subdistant lamellae, and smaller
basidiospores in the range 6-9 × 3.5-4.5 µm
(Singer, 1976). Marasmius trinitatis Dennis,
common throughout the neotropics and
reported from Papua New Guinea by Desjardin
and Horak (1997), differs in forming a
campanulate-convex pileus, more lamellae (1620), a brown to fulvous stipe (not ruby red),
grows on woody debris, and has smaller
basidiospores in the range 8.3-12.5 × 2.7-4 µm
(Singer, 1976). Marasmius makok is sister to M.
bambusiniformis Singer (No. 48 below) in the
ITS phylogenetic tree, and the latter species
differs only in pileus shape and color
(campanulate-convex with reddish brown and
brownish orange tones), and a brown stipe.
45. Marasmius cupreostipes
Desjardin & Lumyong, sp. nov.
MycoBank: MB512424
Wannathes,
(Fig. 53)
Etymology: ‘cupreo’ = copper, ‘stipes’ = stipe;
referring to the copper coloured stipe of dried
basidiomes.
Pileus 3-17 mm diametro, campanulatus, sulcatus
usque plicatus, glaber, hebetatus, disco aurantiacobrunneo, margine laete aurantiaco-brunneo. Contextus
creameus, tenuis. Lamellae annexae, distantes (10-12)
sine lamellula, latae (1-2 mm), creameae, haud
marginatae, haud intervenosae. Stipes 67-250 × 0.5-0.8
mm, centralis, cylindratus, filo metallico similis, glaber,
insititius usque haud insititius, apice aurantiaco, basi
cuprea, ubique cuprea in sicco. Odor saporque non
propria. Basidiosporae (17-)18-21(-26) × 4-5(7-) µm,
cylindratae usque fusoideae, oblique curvae, laeves,
hyalinae, inamyloideae, tenuitunicatae Basidia non
observata. Basidiolae fusoideae usque clavatae.
Cheilocystidia vulgaria, typi Sicci; 13-20 × 6-11 µm,
cylindrata usque clavata vel adumbratim inaequabilia,
plerumque ramosa, hyalina, inamyloidea, tenuitunicata;
setulis apicalibus 2-6 × 1 µm, cylindratis usque conicis,
plerumque undulatis, subacutis, pallide flavis, tenuiusque crassetunicatis. Pleurocystidia nulla. Pileipellis
hymeniformis, haud maculosus, typi Sicci; 10-18(-30) ×
6-15 µm, clavatus usque pyriformis vel adumbratim
Fungal Diversity
inaequabilis, saepe ramosus, hyalinus usque pallide
flavus, inamyloideus, tenuitunicatus; setulis apicalibus 24 × 1-1.5 µm, coartatis, cylindratis, subacutis, brunneis
usque flavo-brunneis, tenui- usque crassetunicatis.
Trama pilei intertexta, dextrinoidea usque leniter
dextrinoidea. Trama lamellae intertexta, hyphis 5-8(-12)
µm diametro, cylindratis usque inflatis, laevibus,
hyalinis, dextrinoidea usque leniter dextrinoidea,
tenuitunicatis, haud gelatinosis. Stipitipellis subparallelus,
hyphis 3-5 µm diametro, cylindratis, brunneis, laevibus,
inamyloideis, tenuitunicatis, haud gelatinosis. Trama
stipitis parallela, hyphis 6-10 µm diametro, cylindratis,
hyalinis, laevibus, dextrinoideis, tenuitunicatis, haud
gelatinosis. Caulocystidia nulla. Fibulae praesentes in
omnibus texturis.
Typus: Chiang Mai Province, Mushroom
Research Centre, 27 km marker on Hwy 1095, dispersus
usque gregarius in folis plantae dicotyledoneae, 3 June
2004, N. Wannathes 150 (CMU: holotypus; SFSU:
isotypus).
Pileus 3-17 mm diam., campanulate,
sulcate to plicate, glabrous, dull, disc orangish
brown, margin light orangish brown. Context
cream, thin. Lamellae adnexed, distant (10-12)
with no lamellula, broad (1-2 mm), cream, nonmarginate, non-intervenose. Stipe 67-250 ×
0.5-0.8 mm, central, cylindrical, wiry, glabrous,
insititious to non-insititious, apex orange, base
copper-colored, copper-colored overall when
dried. Odor and taste not distinctive.
Basidiospores (17-)18-21(-26) × 4-5(7-)
µm [x = 19.4 ± 1.9 × 5.0 ± 0.6 µm, Q = 2.6-5.8,
Qm = 4.0, n = 25 spores, s = 1 specimen],
cylindrical to fusoid, curved in profile, smooth,
hyaline, inamyloid, thin-walled. Basidia not
observed. Basidioles fusoid to clavate.
Cheilocystidia common, of Siccus-type broom
cells; main body 13-20 × 6-11 µm, cylindrical
to clavate or irregular in outline, usually
branched, hyaline, inamyloid, thin-walled;
apical setulae 2-6 × 1 µm, cylindrical to conical,
usually wavy, subacute, pale yellow, thin- to
thick-walled. Pleurocystidia absent. Pileipellis
hymeniform, not mottled, composed of Siccustype broom cells; main body 10-18(-30) × 6-15
µm, clavate to pyriform or irregular in outline,
often branched, hyaline to pale yellow,
inamyloid, thin-walled; apical setulae 2-4 × 11.5 µm, crowded, cylindrical, subacute, brown
to yellowish brown, thin- to thick-walled.
Pileus trama interwoven, dextrinoid to weakly
dextrinoid. Lamellar trama hyphae 5-8(-12)
µm diam., interwoven, cylindrical to inflated,
smooth, hyaline, dextrinoid to weakly
dextrinoid,
thin-walled,
non-gelatinous.
Stipitipellis hyphae 3-5 µm diam., subparallel,
cylindrical, brown, smooth, inamyloid, thinwalled, non-gelatinous. Stipe trama hyphae 610 µm diam., parallel, cylindrical, hyaline,
smooth, dextrinoid, thin-walled, non-gelatinous.
Caulocystidia absent. Clamp connections
present in all tissues.
Fig. 53. Marasmius cupreostipes (N. Wannathes 150). 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pileipellis, Scale bar 1 = 20 mm, 2-4 = 10 µm
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Mushroom Research Centre, 27 km marker on
Hwy 1095, 3 June 2004, N. Wannathes 150 (CMU:
holotype;SFSU: isotype).
Discussion: This new species is
characterized by a campanulate, plicate,
orangish brown pileus, distant, cream-coloured
lamellae, clavate basidiospores with mean 19.4
× 5.0 µm, and an absence of pleurocystidia and
caulocystidia. The most distinctive feature is its
very long (up to 250 mm) copper-colored stipe.
Marasmius aciebrunneus Corner, described
from Singapore, differs in forming brownmarginate lamellae, a fuscous brown stipe only
20-30 mm long, and longer but narrower
basidiospores with mean 24.8 × 4.3 µm
(Holotype E!; Corner 1996). Marasmius
selangorensis Y.S. Tan & Desjardin, described
277
recently from Malaysia, differs in forming a
more brown to yellowish brown pileus, more
numerous lamellae (12-18), a stipe only up to
43 mm long coloured brownish orange to dark
brown (brown when dried), and acerose apical
setulae on broom cells that are 4-15 µm long
(Tan et al., 2007). Marasmius longistipitatus
Antonín, described from Africa, differs in
forming paler pilei (pale greyish orange to
orange white), and distinctive pleurocystidia.
In all other characters the two species are
similar (Antonín, 2004, 2007).
46. Marasmius corneri Wannathes, Desjardin
& Lumyong, Mycol. Res. 111: 992. 2007.
(Figs 54)
≡ Marasmius incarnatus Corner, Beih. Nova
Hedwigia 111: 60. 1996., nom. illegit., non Marasmius
incarnatus Quél., Enchir. Fung.:142. 1886.
Pileus 10-43 mm diam, obtusely conical
to convex or broadly convex, with a shallow
broad umbo, disc occasionally rugulose,
margin striate to sulcate, dull, dry, minutely
velutinous, disc brownish orange (7C8) to light
brown (7D8), orange (6B8) or brownish yellow
(5C7-8), margin light orange (5A4), brownish
orange (6C7), orange (5A6) or grayish orange
(5B5-6). Context thin, yellowish white.
Lamellae adnexed to subfree, subdistant (12-18)
with 1-4 series of lamellulae, non-intervenose,
narrow, yellowish white (4A2-3), nonmarginate. Stipe 23-90 × 0.8-3 mm, central,
terete, cylindrical, with or without a
subbulbous base, hollow, glabrous, noninsititious, base covered with brownish orange
mycelium, apex yellowish white (3-4A2) to
light orange (5A4), base brownish orange (6C8)
to dark brown (6F8) or dark reddish brown,
rhizomorphs absent. Odor and taste not
distinctive.
Basidiospores (16-)18-20(-22) × 4-5 μm
[xmr = 18.3-19.6 × 4.3-4.7 μm, xmm = 19.0 ± 0.6
× 4.5 ± 0.2 μm, Q = 3.2-5.5, Qmr = 4.0-4.5, Qmm
= 4.2 ± 0.2, n = 25 spores, s = 4 specimens],
cylindrical to fusoid, often curved in profile,
smooth, hyaline, inamyloid, thin-walled.
Basidia not observed. Basidioles clavate to
cylindrical. Cheilocystidia common, of Siccustype broom cells; main body 15-25 × 4-9 μm,
cylindrical to clavate, broadly clavate or
278
irregular in outline, hyaline, thin-walled; apical
setulae (2-)3-10 × 0.5-1.5(-2) μm, narrowly
conical to cylindrical or irregular in outline,
subacute to acute, pale yellow, inamyloid, thickwalled. Pleurocystidia absent. Pileipellis
hymeniform, not mottled, composed of Siccustype broom cells; main body (7-)14-25(-30) ×
5-7(-10) μm, clavate to broadly clavate,
cylindrical, pyriform or sometimes irregular in
outline, hyaline, inamyloid, thin- to thickwalled; apical setulae (2-)3-10 × 0.8-1.5 μm,
cylindrical to conical or irregular in outline,
subacute to acute, pale yellow to tawny,
inamyloid, thick-walled. Pileus trama and
lamellar trama interwoven; hyphae (2-)4-8(-11)
μm diam, cylindrical, smooth, hyaline,
dextrinoid to strongly dextrinoid, thin-walled,
non-gelatinous. Stipitipellis hyphae 5-6(-10)
μm diam, parallel, cylindrical, smooth, hyaline
to yellowish brown, dextrinoid to strong
dextrinoid, thin- to thick-walled (>1.5 μm),
non-gelatinous. Stipe trama hyphae 3-7(-12)
μm diam, parallel, cylindrical, smooth, hyaline,
dextrinoid to strongly dextrinoid, thin- to thickwalled, non-gelatinous. Caulocystidia absent.
Clamp connections present in all tissues.
Fig. 54 Marasmius corneri (N. Wannathes 066). 1.
Basidiomes 2. Basidioles 3. Basidiospores 4. Cheilocystidia 5.
Pileipellis, Scale bar 1 = 20 mm, 2-5 = 10 µm
Fungal Diversity
Habit, habitat, and known distribution:
Scattered to gregarious on undetermined
dicotyledonous leaves or wood, Singapore and
Northern Thailand.
Material examined: Singapore. Gardens Jungle, 8
Aug. 1940, Corner s.n. (E: holotype). Thailand. Chiang
Mai Province, Doi Suthep-Pui National Park, Mok Fa
waterfall on Hwy 1095, N19º 6.581’, E98º 46.353’, 1014
meters, 25 Jul. 2003, N. Wannathes 026 (CMU, SFSU);
same location, 1 Aug. 2003, N. Wannathes 066 (CMU,
SFSU); same location, 13 Aug. 2003, N. Wannathes 103
(CMU, SFSU); same location, 29 June 2004, Y.S. Tan
274 (CMU, SFSU); same location, 7 Jul. 2004, Y.S. Tan
325 and N. Wannathes 218 (CMU, SFSU); same
location, 21 Aug. 2004, N. Wannathes 269 (CMU,
SFSU).
Discussion: The name used by Corner
(1996) for this new species described recently
from Singapore is a homonym (non Quélet
1886) and a new name was proposed by
Wannathes et al. (2007). Their choice of
epithet honors E.J.H. Corner who has
published many new species of Marasmius
from Malaysia. The material from northern
Thailand differs only slightly from the
protologue in having pilei that are more deeply
pigmented, ranging from greyish orange to
brownish yellow. The micromorphological
characters are indistinguishable from those of
the holotype specimen (E!).
A bipolar mating system was reported for
M. corneri by Wannathes et al. (2007) based on
Thai material.
inamyloidea, tenuitunicata; setulis apicalibus 3-9 × 1 µm,
cylindratis, plerumque undulatis, saepe furcatis,
inamyloideis, subacutis, hyalinis usque pallide flavis,
crassetunicatis.
Pleurocystidia
nulla.
Pileipellis
hymeniformis, leniter maculosus, typi Sicci; 10-18 × 79(-12) µm, clavatus usque late clavatus vel pyriformis,
hyalinus, inamyloideus, tenui- usque crassetunicatus;
setulis apicalibus 3-5(-8) × 1-1.5 µm, coartatis,
cylindratis, plerumque undulatis, subacutis, brunneis
usque atrobrunneis, crassetunicatis. Trama pilei
intertexta, leniter dextrinoidea usque inamyloidea.
Trama lamellae intertexta, hyphis 3-6(-8) µm diametro,
cylindratis usque inflatis, laevibus, hyalinis, leniter
dextrinoidea usque inamyloidea, tenuitunicatis, haud
gelatinosis. Stipitipellis subparallelus, hyphis (2-)3-5(-6)
µm diametro, cylindratis, brunneis usque atrobrunneis,
laevibus, inamyloideis usque leniter dextrinoideis,
crassetunicatis, haud gelatinosis. Trama stipitis
subparallela, hyphis 3-5(-8) µm diametro, cylindratis,
hyalinis, laevibus, inamyloideis usque leniter
dextrinoideis,
tenuitunicatis,
haud
gelatinosis.
Caulocystidia nulla. Fibulae praesentes in omnibus
texturis.
Typus: Phrae Province, Muang District, Pa Dang,
Nong Kam Village, near Nong Kam reservoir, dispersus
usque gregarius in folis plantae dicotyledoneae vel ligno,
18 August 2005, N. Wannathes 423 (CMU: holotypus;
SFSU: isotypus).
47. Marasmius imitarius Wannathes, Desjardin
& Lumyong, sp. nov.
(Figs 55, 57-4)
MycoBank: MB 512425
Etymology: ‘imitari’ = to imitate; referring to the
micromorphology of the species being indistinguishable
from that of a number of other species.
Pileus 2-15 mm diametro, obtuse conicus usque
convexus ubi iuvenis, expansus et plano-convexus et
saepe leviter depressus ubi vetus, umbone rugoso,
striatus usque sulcatus, pruinosus, hebetatus, disco
brunneo usque rubro-brunneo, margine pallidiore usque
laete brunnea. Contextus flavo-albus, tenuis. Lamellae
annexae, distantes (10-12), angustae, creameae,
brunneomarginatae, haud intervenosae. Stipes 8-23(-80)
× < 0.5 mm, centralis, cylindratus, filo metallico similis,
glaber, insititius usque haud insititius, apice albido, basi
brunnea usque atrobrunnea. Odor saporque non propria.
Basidiosporae 17-20(-21) × 4-5 µm, cylindratae usque
fusoideae,
oblique
curvae,
laeves,
hyalinae,
inamyloideae, tenuitunicatae Basidia non observata.
Basidiolae cylindratae usque clavatae. Cheilocystidia
vulgaria, typi Sicci; 8-17(-23) × (3-)6-10(-15) µm,
cylindrata usque clavata vel pyriformia, hyalina,
Fig. 55. Marasmius imitarius (N. Wannathes 423). 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pileipellis, Scale bar 1 = 20 mm, 2-4 = 10 µm
Pileus 2-15 mm diam., obtusely conical
to convex when young, expanding to convex
and often slightly depressed in age, with a
rugulose umbo, striate to sulcate, pruinose, dull,
disc brown (6-7E5-8) to reddish brown (8E6-7),
margin paler to light brown (7D5-6). Context
yellowish white, thin. Lamellae adnexed,
279
distant (10-12), narrow, cream with brown
edges, non-intervenose. Stipe 8-23(-80) × < 0.5
mm, central, cylindrical, wiry, glabrous,
insititious to non-insititious, apex off-white,
base brown to dark brown. Odor and taste not
distinctive.
Basidiospores 17-20(-21) × 4-5 µm [xmr
= 17.8-19.2 × 4.1-4.7 µm, xmm = 18.6 ± 0.7 ×
4.4 ± 0.3 µm, Qmr = 4.1-4.4, Qmm = 4.4 ± 0.3, n
= 25 spores, s = 3 specimens], cylindrical to
fusoid, curved in profile, smooth, hyaline,
inamyloid, thin-walled. Basidia not observed.
Basidioles cylindrical to clavate. Cheilocystidia
common, of Siccus-type broom cells; main
body 8-17(-23) × (3-)6-10(-15) µm, cylindrical
to clavate or pyriform, hyaline, inamyloid,
thin-walled; apical setulae 3-9 × 1 µm,
cylindrical, usually wavy, often forked,
inamyloid, subacute, hyaline to pale yellow,
thick-walled. Pleurocystidia absent. Pileipellis
hymeniform, weakly mottled, composed of
Siccus-type broom cells; main body 10-18 × 79(-12) µm, clavate to broadly clavate or
pyriform, hyaline, inamyloid, thin- to thickwalled; apical setulae 3-5(-8) × 1-1.5 µm,
crowded, cylindrical, usually wavy, subacute,
brown to dark brown, thick-walled. Pileus
trama interwoven, weakly dextrinoid to
inamyloid. Lamellar trama hyphae 3-6(-8) µm
diam., interwoven, cylindrical to inflated,
smooth, hyaline, weakly dextrinoid to
inamyloid,
thin-walled,
non-gelatinous.
Stipitipellis hyphae (2-)3-5(-6) µm diam,
subparallel, cylindrical, brown to dark brown,
smooth, inamyloid to weakly dextrinoid, thickwalled, non-gelatinous. Stipe trama hyphae 35(-8) µm diam., subparallel, cylindrical,
hyaline, smooth, inamyloid to weakly
dextrinoid,
thin-walled,
non-gelatinous.
Caulocystidia absent. Clamp connections
present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves or on wood, Northern Thailand.
Material examined: Thailand. Chiang Rai
Province, Muang District, Khun Con Waterfall, 12 June
2005, N. Wannathes 297 (CMU, SFSU); Phrae Province,
Muang District, Pa Dang, Nong Kam Village, near Nong
Kam reservoir, 18 August 2005, N. Wannathes 423
(CMU: holotype; SFSU: isotype) and N. Wannathes 425
(CMU, SFSU).
Discussion: Marasmius imitarius is
characterized by a light brown, sulcate pileus
280
with darker brown to reddish brown disc,
distant (10-12), cream-coloured lamellae with
brown edges, clavate basidiospores with mean
18.6 × 4.4 µm, an absence of pleurocystidia
and caulocystidia, and growth mostly on
woody sticks. The micromorphology of this
new species is very similar to that of many
other species, hence our choice of epithets.
Marasmius kanchingensis Y.S. Tan &
Desjardin, described recently from Malaysia,
differs in forming a striped pileus with more
brownish orange to greyish orange tones, more
lamellae (14-16), slightly longer basidiospores
with mean range 18.5-20.5 µm long, and
growth on leaves. Marasmius mazatecus Singer,
an orange-ferruginous to deep rusty-red species
from Mexico, M. striaepileus Antonín, an
orangish brown species from Burundi, and M.
sierraleonis Beeli, a dull yellowish to rusty
brown species from Africa, differ primarily in
forming larger, more brightly pigmented pilei.
Marasmius bambusiniformis Singer (see below)
differs in forming a brownish orange to greyish
orange pileus, more lamellae (10-18), has
slightly smaller basidiospores with mean 16.2 x
3.7 µm, and grows on dicot leaves. In the ITS
phylogenetic analyses, M. imitarius is not sister
to M. bambusiniformis.
48. Marasmius bambusiniformis Singer, Fl.
Neotrop. Monogr. 17: 167. 1976.
(Figs 56, 57-5)
Pileus 3-10 mm diam., obtusely conical,
usually umbonate, striate to sulcate, pruinose,
dull, disc reddish brown (9E8) to brownish
orange (6C8), margin brown (7D8) to greyish
orange (6B4). Context thin, cream. Lamellae
adnexed, subdistant (10-18) with 0-1 series of
lamellulae, narrow, cream with brown edges,
non- intervenose. Stipe 13-52 × 0.2-0.4 mm,
central, cylindrical, wiry, glabrous, noninsititious, apex yellowish white, base brown.
Odor and taste not distinctive.
Basidiospores (15-)16-18(-19) × 3.5-4
µm [xmr = 15.7-17 × 3.6-4.0 µm, xmm = 16.2 ±
0.7 × 3.7 ± 0.2 µm, Qmr = 4.3-4.5, Qmm = 4.4 ±
0.1, n = 25 spores, s = 3 specimens], narrowly
fusoid, curved in profile, smooth, hyaline,
inamyloid, thin-walled. Basidia not observed.
Basidioles fusoid to clavate. Cheilocystidia
common, of Siccus-type broom cells; main
body (4-)10-18 × 6-9 µm, cylindrical to clavate,
Fungal Diversity
Koo Haa Waterfall, 15 August 2005, N. Wannathes 410
(CMU, SFSU).
Discussion: Diagnostic features of M.
bambusiniformis, include: a small, striate to
sulcate pileus with reddish brown to brownish
orange disc and greyish ornge margin;
subdistant (10-18), cream-coloured lamellae
with brown edges; a wiry, glabrous, noninsititious stipe; basidiospores with mean 16.2
x 3.7 µm,; an absence of pleurocystidia and
caulocystidia; and growth on dicot leaves. The
Thai specimens match nicely those reported
from Papua New Guinea by Desjardin and E.
Horak (1997), but differ from the holotype
collected in Ecuador in forming more lamellae
(10-18 vs 7-10).
Fig. 56. Marasmius bambusiniformis (N. Wannathes
410). 1. Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pileipellis, Scale bar 1 = 20 mm, 2-4 = 10 µm
hyaline, inamyloid, thin-walled; apical setulae
2-5 × 1 µm, cylindrical to conical, subacute,
yellow, thick-walled. Pleurocystidia absent.
Pileipellis hymeniform, weakly mottled,
composed of Siccus-type broom cells; main
body 10-20 × (4-)7-13 µm, cylindrical to
clavate or irregular in outline, seldom branched,
hyaline, inamyloid, thin- to thick-walled; apical
setulae 2-5(7-) × 1 µm, crowded, cylindrical,
subacute, brownish yellow to light brown, thick
walled. Pileus trama interwoven, dextrinoid.
Lamellar trama hyphae (3-)4-10(-15) µm diam.,
interwoven, cylindrical, smooth, hyaline,
dextrinoid, thin-walled, non-gelatinous. Stipitipellis hyphae (2-)4-7 µm diam., subparallel,
cylindrical, yellowish brown to light brown,
smooth, dextrinoid, thick-walled (up to 1.5 µm),
non-gelatinous. Stipe trama hyphae 3-10 µm
diam, parallel, cylindrical, hyaline, smooth,
dextrinoid, thin-walled, non-gelatinous. Caulocystidia absent. Clamp connections present in
all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, Malaysia, neotropics, Northern
Thailand and Papua New Guinea.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mokfa
Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
m., 25 June 2005, N. Wannathes 329 (CMU, SFSU);
Chiang Mai Province, Doi Inthanon National Park, At 25
km marker on Highway 1,009, N 18º 32.5’ E 98º 33.5’
alt. 1,076 m., 27 June 2005, N. Wannathes 368 (CMU,
SFSU); Phrae Province, Muang District, Suen Keun, Na
Section Sicci, subsect. Siccini, ser.
Haematocephali Singer, Fl. Neotrop.
Monogr. 17: 201. 1976.
Type: Marasmius haematocephalus
(Mont.) Fr.
49. Marasmius confertus Berk. & Broome, J.
Linn. Soc., Bot. 14: 34. 1873. (Figs 57-7, 58)
= Marasmius chondripes Berk. & Broome, J.
Linn. Soc., Bot. 14: 36. 1873.
= Marasmius hemibaphus Berk. & Broome, J.
Linn. Soc., Bot. 14: 39. 1873.
Pileus 18-24(-40) mm diam., convex to
broadly convex, subumbonate, smooth to
striatulate, glabrous, dull, disc brown to dark
brown (7E-F8) or reddish brown (8E-F8),
margin brownish orange (5C4-5) to yellowish
brown (5E8), hygrophanous, disc fading to
light brown (6D6). Context thin, orangish grey
(5B3). Lamellae adnexed to adnate, close (1518) with 3-4 series of lamellulae, broad (2-3
mm), pale greyish yellow, non-marginate or
slightly pale brown-marginate, non-intervenose.
Stipe 53-83 × 1-2 mm, central, cylindrical, wiry,
hollow, glabrous, non-insititious, light brown
(5D6) overall. Odor of coconut juice. Taste not
distinctive.
Basidiospores 8-10(-12) × 4-5(-6) µm
[xmr = 9.3-9.8 × 4.3-5.1 µm, xmm = 9.6 ± 0.3 ×
4.8 ± 0.4 µm, Qmr = 1.9-2.2, Qmm = 2.1 ± 0.1, n
= 25 spores, s = 3 specimens], ellipsoid,
smooth, hyaline, inamyloid, thin-walled.
Basidia not observed. Basidioles cylindrical to
clavate. Cheilocystidia scattered, of Siccus-type
broom cells; main body 9-16 × 4-7 µm,
cylindrical to fusoid, hyaline, inamyloid,
281
Fig. 57. Basidiomata of Marasmius section Sicci ser. Leonini. and ser. Haematocephali 1. M. cremeus (N. Wannathes
154) 2. M. hypochroides (N. Wannathes 335) 3. M. makok (N. Wannathes 201) 4. M. imitarius (N. Wannathes 423) 5. M.
bambusiniformis (N. Wannathes 410) 6. M. brunneoolivascens (N. Wannathes 277) 7. M. confertus (Y.S. Tan 294) 8. M.
suthepensis (N. Wannathes 309).
282
Fungal Diversity
thin-walled; apical setulae 3-7(-11) × 1(-2) µm,
cylindrical, obtuse to subacute, hyaline, thinwalled. Pleurocystidia common, 38-52 × 7-13
µm, cylindrical to fusoid or irregular in outline,
wavy to constricted at the apex, hyaline,
inamyloid, thin-walled. Pileipellis hymeniform,
mottled, composed of Siccus-type broom cells;
main body (5-)8-14 × 4-7 µm, cylindrical to
clavate or irregular in outline, often branched,
light brown, inamyloid, thin- to thick-walled;
apical setulae 4-11(-20) × 1-1.5(-2) µm,
crowded, cylindrical, obtuse to subacute,
brown, thick-walled. Pileus trama interwoven,
dextrinoid.
dextrinoid, thin-walled, non-gelatinous. Caulocystidia absent. Clamp connections present in
all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves. South America (Bolivia, Brazil, Peru,
Venezuela), Africa (Kenya, Uganda), Sri
Lanka, and Northern Thailand.
Material examined. Thailand. Chiang Mai
Province, Doi Inthanon National Park, Junction of
Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E
98º 29.6’ alt. 1,703 m., 3 July 2004, Y. S. Tan 294 and
300 (CMU, SFSU); same location, 27 June 2005, N.
Wannathes 359 (CMU).
Discussion: Marasmius confertus is
characterized by a smooth to striatulate pileus
colored brownish orange to yellowish brown
with brown to reddish brown disc, close (15-18)
and broad lamaellae, a glabrous light brown
stipe, relatively small basidiospores with mean
9.6 × 4.8 µm, strangulate, cylindrical
pleurocystidia, and growth on dicot leaves. The
Thai specimens match quite closely with the
Sri Lankan specimens reported by Pegler
(1986), except for forming larger pilei (up to
40 mm vs up to 20 mm) and having broader
basidiospores (4-5 µm vs 3-4 µm). The species
has a widespread distibution in tropical regions,
and has been recorded from Venezuela (Dennis,
1951), Bolivia, Brazil and Peru (Singer, 1965,
1976), and Kenya and Uganda (Pegler, 1977).
50. Marasmius ganyao Wannathes, Desjardin
& Lumyong, sp. nov.
(Fig. 59)
MycoBank: MB512426
Fig. 58. Marasmius confertus (Y.S. Tan 294). 1.
Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5. Pleurocystidia 6. Pileipellis, Scale bar 1
= 20 mm, 2-6 = 10 µm
Lamellar trama hyphae 3-10(-14) µm diam.,
interwoven to regular, cylindrical to inflated,
smooth, hyaline, dextrinoid to weakly
dextrinoid,
thin-walled,
non-gelatinous.
Stipitipellis hyphae 3-7(-9) µm diam., parallel,
cylindrical, yellowish brown to light brown,
smooth, inamyloid, thick-walled (up to 1 µm),
non-gelatinous. Stipe trama hyphae 7-14 µm
diam., parallel, cylindrical, hyaline, smooth,
Etymology: ‘ganyao’ (Thai) = long stipe;
referring to the exceedingly long stipe.
Pileus 2-5 mm diametro, obtuse conicus, sulcatus
usque plicatus, glaber usque pruinosus, hebetatus, disco
atro usque atrobrunneo, margine brunneo vel sulcis laete
brunneis
usque
brunneo-aurantiacis.
Contextus
aurantiaco-albus, tenuis. Lamellae annexae usque
adnatae, distantes (6-8), angustae, flavo-albae, haud
marginatae, haud intervenosae. Stipes 100-195 × 0.5 mm,
centralis, cylindratus, filo metallico similis, tenax, glaber,
insititius usque haud insititius, apice flavo-albo, basi
brunnea. Odor saporque non propria. Basidiosporae 2532 × 4-4.5 µm, cylindratae usque clavatae, laeves,
hyalinae, inamyloideae, tenuitunicatae Basidia 36-40 ×
9-10 µm, cylindrata usque clavata, 4-spora. Basidiolae
cylindratae usque clavatae. Cheilocystidia vulgaria, typi
Sicci; 9-19 × 4-8(-13) µm, cylindrata usque clavata vel
adumbratim
inaequabilia,
hyalina,
inamyloidea,
tenuitunicata; setulis apicalibus 3-9 × 1-2 µm, cylindratis,
saepe undulatis, raro furcatis, subacutis, pallide flavis
usque flavis, crassetunicatis. Pleurocystidia vulgaria, 51-
283
82 × 7-13 µm, cylindrata usque fusoidea, plerumque
constricta ad apicem, hyalina, inamyloidea, tenuitunicata,
raro crassetunicata ad apicem. Pileipellis hymeniformis,
maculosus, typi Sicci; (5-)10-19 × 5-12 µm, cylindratus
usque clavatus vel adumbratim inaequabilis, saepe
ramosus, pallide flavus, inamyloideus, tenui- usque
crassetunicatus; setulis apicalibus 2-8 × 1-2 µm, coartatis,
cylindratis, undulatis, subacutis usque acutis, brunneis
usque atrobrunneis, crassetunicatis. Trama pilei
intertexta, dextrinoidea. Trama lamellarum intertexta,
hyphis 3-7 µm diametro, cylindratis usque inflatis,
laevibus, hyalinis, leniter dextrinoideis, tenuitunicatis,
haud gelatinosis. Stipitipellis subparallelus, hyphis 2-5
µm diametro, cylindratis, brunneis usque laete brunneis,
laevibus, inamyloideis usque leniter dextrinoideis, tenuiusque crassetunicatis, haud gelatinosis. Trama stipitis
parallela, hyphis 3-8 µm diametro, cylindratis, hyalinis,
laevibus, inamyloideis, tenui- usque crassetunicatis, haud
gelatinosis. Caulocystidia nulla. Fibulae praesentes in
omnibus texturis.
Typus. Chiang Mai Province, Huai Nam Dang
National Park, North 19º 18.3’ E 98º 35.8’ alt. 1,538 m.,
dispersus usque gregarius in folis bambusae, 24 June
2003, N. Wannathes 005 (CMU: holotypus; SFSU:
isotypus).
Pileus 2-5 mm diam., obtusely conical,
sulcate to plicate, glabrous to pruinose, dull,
disc black to dark brown, margin brown or
with light brown to brownish orange sulcae.
Context thin, orangish white. Lamellae
adnexed to adnate, distant (6-8), narrow,
yellowish
white,
non-marginate,
nonintervenose. Stipe 100-195 × 0.5 mm, central,
cylindrical, wiry, tough, glabrous, insititious to
non-insititious, apex yellowish white, base
brown. Odor and taste not distinctive.
Basidiospores 25-32 × 4-4.5 µm [xmr =
28.2-28.3 × 4.1-4.6 µm, xmm = 28.2 ± 0.1 × 4.3
± 0.4 µm, Qmr = 6.2-7.0, Qmm = 6.6 ± 0.6, n =
25 spores, s = 2 specimens], cylindrical to
clavate, smooth, hyaline, inamyloid, thinwalled. Basidia 36-40 × 9-10 µm, cylindrical to
clavate, 4-spored. Basidioles cylindrical to
clavate. Cheilocystidia common, of Siccus-type
broom cells; main body 9-19 × 4-8(-13) µm,
cylindrical to clavate or irregular in outline,
hyaline, inamyloid, thin-walled; apical setulae
3-9 × 1-2 µm, cylindrical, often wavy, seldom
forked, subacute, pale yellow to yellow, thickwalled. Pleurocystidia common, 51-82 × 7-13
µm, cylindrical to fusoid, usually constricted at
the apex, hyaline, inamyloid, thin-walled,
rarely apically thick-walled. Pileipellis hymeniform, mottled, composed of Siccus-type broom
cells; main body (5-) 10-19 × 5-12 µm,
cylindrical to clavate or irregular in outline,
284
often branched, pale yellow, inamyloid, thin- to
thick walled; apical setulae 2-8 × 1-2 µm,
crowded, cylindrical, wavy, subacute to acute,
brown to dark brown, thick-walled. Pileus
trama interwoven, dextrinoid. Lamellar trama
hyphae 3-7 µm diam., interwoven, cylindrical
Fig. 59 Marasmius ganyao (N. Wannathes 183). 1.
Basidiomes 2. Basidia 3. Basidiospores 4. Cheilocystidia
5. Pleurocystidia 6. Pileipellis, Scale bar 1 = 20 mm, 2-6
= 10 µm
to inflated, smooth, hyaline, weakly dextrinoid,
thin-walled, non-gelatinous. Stipitipellis hyphae 2-5 µm diam., subparallel, cylindrical,
brown to light brown, smooth, inamyloid to
weakly dextrinoid, thin- to thick-walled, nongelatinous. Stipe trama hyphae 3-8 µm diam.,
parallel, cylindrical, hyaline, smooth, inamyloid, thin- to thick-walled, non-gelatinous.
Caulocystidia absent. Clamp connections
present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on bamboo leaves,
Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Huai Nam Dang National Park, North 19º
Fungal Diversity
18.3’ E 98º 35.8’ alt. 1,538 m., 24 June 2003, N.
Wannathes 005 (CMU: holotype; SFSU: isotype); same
location, 28 June 2004, N. Wannathes 183 (CMU, SFSU)
Discussion: The new species is
distinguished by a small obtusely conical,
sulcate pileus with a black to dark brown disc
and brown margin with paler sulcae, distant (68), narrow lamellae, a long (up to 195 mm
long ), wiry, brown stipe that is often insititious
and grows on bamboo leaves, long clavate
basidiospores with mean 28.2 × 4.3 µm, and
simple strangulate pleurocystidia. Marasmus
ganyao is similar to the following species.
Marasmius davidii Antonín, a species
previously known as M. brunneolus (Berk. &
Broome) Pegler (non M. brunneolus (Beeli)
Singer; cf. Antonín, 2003) and described from
Sri Lanka, differs in forming larger pilei (711mm vs 2-5 mm), a shorter stipe (30-45 mm),
smaller basidiospores (20-27 × 4-6 µm, xm = 22
× 4.5 µm), and has a lignicolous habit (Pegler,
1986). Marasmius longistipitatus Antonín,
from Africa, differs in forming a pale greyish
orange to orangish white pileus with a distinct
conical papilla, more numerous and
pseudocollariate lamellae (9-12), a reddish
orange to brownish orange stipe, and slightly
smaller basidiospores (20-28 × 4.5-6 µm)
(Antonín, 2007). Marasmius megistosporus
Singer, described from Bolivia, differs in
forming larger pilei (up to 35 mm diam.) with
deep ferruginous to golden brown margin,
possibly longer basidiospores (28-37.5 µm
long), and has a lignicolous habit (Singer, 1965,
1976).
51. Marasmius
graminipes
Desjardin & Lumyong, sp. nov.
MycoBank: MB512427
Wannathes,
(Fig. 60)
Etymology: ‘graminipes’ (Latin) = grass-stem;
referring to the tufts of grass-like caulocystidia on the
stipe surface.
Pileus 8-23 mm diametro, campanulatus, striatus,
glaber, hebetatus, disco atrobrunneo, margine brunneocinereus. Contextus brunneo-creameus, tenuis. Lamellae
annexae, arctae (20-24) cum 3 seriebus lamellularum,
latae, cinereo-albae, haud marginatae, haud intervenosae.
Stipes 35-45 × 1.5-2.5 mm, centralis, cylindratus,
pruinosae, haud insititius, apice brunneo-flavo, basi
brunnea. Odor saporque non propria. Basidiosporae 1821(-22) × 4 µm, anguste clavatae usque cylindratae,
laeves, hyalinae, inamyloideae, tenuitunicatae Basidia
non observata. Basidiolae clavatae. Cheilocystidia
vulgaria, typi Sicci; 10-15 × 5-6 µm, cylindrata usque
clavata, hyalina, inamyloidea, tenuitunicata; setulis
apicalibus 3-12(-16) × 1-1.5(-2) µm, cylindratis, saepe
undulatis, raro furcatis, subacutis, hyalinis, tenuitunicatis.
Pleurocystidia dispersa usque rara, 45-47 × 8-10 µm,
cylindrata usque fusoidea, plerumque undulata vel
constricta ad apicem, hyalina, inamyloidea, tenuitunicata.
Pileipellis hymeniformis, maculosus, typi Sicci; 12-27 ×
8-12 µm, clavatus usque late clavatus vel turbinatus,
hyalinus, inamyloideus, tenui- usque crassetunicatus;
setulis apicalibus 5-18 × 1-2 µm, coartatis, cylindratis,
subacutis, atrobrunneis, crassetunicatis. Trama pilei
intertexta, hyphis 5-10 µm diametro, cylindratis usque
inflatis, laevibus, hyalinis, dextrinoideis, tenuitunicatis.
Trama lamellae regularis usque intertexta, hyphis 5-12
µm diametro, cylindratis, laevibus, hyalinis, leniter
dextrinoideis,
tenuitunicatis,
haud
gelatinosis.
Stipitipellis parallelus, hyphis 4-9(-11) µm diametro,
cylindratis, flavis usque laete brunneis, laevibus,
inamyloideis, tenuitunicatis, haud gelatinosis. Trama
stipitis parallela, hyphis 6-8 µm diametro, cylindratis,
hyalinis, laevibus, dextrinoideis, tenuitunicatis, haud
gelatinosis. Caulocystidia abundantia, plerumque
caespitosa, typi Sicci; 6-28 × 7-8 µm, cylindrata, hyalina
usque pallide flava, inamyloidea, tenuitunicata; setulis
apicalibus 5-40 × 1.5-2 µm, cylindratis, saepe furcatis,
conicis usque subacutis, flavis, crassetunicatis (usque ad
1 µm). Fibulae praesentes in omnibus texturis.
Typus: Chiang Mai Province, Doi Inthanon
National Park, Junction of Highway 1,009 and road to
Mae Chaem, N 18º 31.6’ E 98º 29.6’ alt. 1,703 m.,
dispersus usque gregarius in ligno vel folis plantae
dicotyledoneae, 2 August 2003, N. Wannathes 078
(CMU: holotypus; SFSU: isotypus).
Pileus 8-23 mm diam., campanulate,
striate, glabrous, dull, disc dark brown, margin
brownish grey. Context thin, brownish cream.
Lamellae adnexed, close (20-24) with 3 series
of lamellulae, broad, greyish white, nonmarginate, non-intervenose. Stipe 35-45 × 1.52.5 mm, central, cylindrical, pruinose, noninsititious, apex brownish yellow, base brown.
Odor and taste not distinctive.
Basidiospores 18-21(-22) × 4 µm [x =
19.3 ± 1.0 × 4.0 ± 0 µm, Q = 4.5-5.5, Qm = 4.8,
n = 25 spores, s = 1 specimen], narrowly
clavate to cylindrical, smooth, hyaline,
inamyloid, thin-walled. Basidia not observed.
Basidioles clavate. Cheilocystidia common, of
Siccus-type broom cells; main body 10-15 × 56 µm, cylindrical to clavate, hyaline, inamyloid,
thin-walled; apical setulae 3-12(-16) × 1-1.5(-2)
µm, cylindrical, often wavy, seldom forked,
subacute, hyaline, thin-walled. Pleurocystidia
scattered to rarely present, 45-47 × 8-10 µm,
cylindrical to fusoid, usually wavy or
constricted at the apex, hyaline, inamyloid, thin285
often forked, conical to subacute, yellow,
slightly thick-walled (up to 1 µm). Clamp
connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on wood or on
dicotyledonous leaves, Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Inthanon National Park, Junction of
Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E
98º 29.6’ alt. 1,703 m., 2 August 2003, N. Wannathes
078 (CMU: holotype; SFSU: isotype).
Fig. 60. Marasmius graminipes (N. Wannathes 078). 1.
Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5. Pleurocystidia 6. Pileipellis 7.
Caulocystidia, Scale bar 1 = 20 mm, 2-7 = 10 µm
walled. Pileipellis hymeniform, mottled,
composed of Siccus-type broom cells; main
body 12-27 × 8-12 µm, clavate to broadly
clavate or turbinate, hyaline, inamyloid, thinto thick-walled; apical setulae 5-18 × 1-2 µm,
crowded, cylindrical, subacute, dark brown,
thick-walled. Pileus trama hyphae 5-10 µm
diam., interwoven, cylindrical to inflated,
smooth, hyaline, dextrinoid, thin-walled.
Lamellar trama hyphae 5-12 µm diam., regular
to interwoven, cylindrical, smooth, hyaline,
weakly dextrinoid, thin-walled, non-gelatinous.
Stipitipellis hyphae 4-9(-11) µm diam., parallel,
cylindrical, yellow to light brown, smooth,
inamyloid, thin-walled, non-gelatinous. Stipe
trama hyphae 6-8 µm diam., parallel,
cylindrical, hyaline, smooth, dextrinoid, thinwalled, non-gelatinous. Caulocystidia abundant,
usually present in cespitose clusters of many
cells, composed of Siccus-type broom cells
with main body 6-28 × 7-8 µm, cylindrical,
hyaline to pale yellow, inamyloid, thin-walled;
apical setulae 5-40 × 1.5-2 µm, cylindrical,
286
Discussion: Distinctive features of
M. graminipes include: a striate, brownish grey
pileus with dark brown disc; close (20-24),
broad, greyish white lamellae; a brown,
pruinose stipe; narrowly clavate basidiospores
with mean 19.3 × 4 µm; simple, strangulate
pleurocystidia; and numerous grass-like tufts of
Siccus-type broom cells on the stipe surface.
The new species is similar to several browncolored Asian species. Marasmius coklatus (No.
31) differs in forming smaller basidiospores
(mean 11.2 × 6.1 µm), and numerous hymenial
setae, pileosetae and caulosetae. Marasmius
nummularioides Desjardin & Y.S. Tan,
described from Malaysia, differs in forming a
non-striate pileus coloured more reddish brown,
fewer lamellae (16-18) with brown edges,
smaller basidiospores (mean 16.7 × 4.1 µm),
and Siccus-type broom cells on the stipe
surface with more well-developed basal cells,
although they do form setulae up to 60 µm long
(Tan et al., 2007). Marasmius bondoi (see
below) differs in forming slightly smaller
basidiospores with mean range 14.2-18.3 × 3.54.0 µm, fewer lamellae (12-18), and a golden
brown stipe that lacks caulocystidia. Although
in the ML phylogenetic tree (Fig. 1) the
presence of M. graminipes makes M. bondoi
paraphyletic, a separate branch and bound
analysis of a subset including only M. bondoi
and M. graminipes isolates with M. plicatulus
Peck as an outgroup (Fig. 74), indicates that M.
graminipes is sister to all M. bondoi isolates
and supports our recognition of M. graminipes
as a distinct taxon.
52. Marasmius brunneoolivascens Wannathes,
Desjardin & Lumyong, sp. nov.
(Figs 59-6, 61)
MycoBank: MB512428
Etymology: ‘brunneo’ (Latin) = brown,
‘olivascens’ (Latin) = olive; referring to the olive brown
color of the pileus and lamellae.
Fungal Diversity
Pileus 6-23 mm diametro, convexus cum umbone
rugoso ubi iuvenis, expansus et plano-convexus et leviter
depressus ubi vetus, striatus usque sulcatus, pruinosus,
hebetatus, primo ubique atrobrunneus demum disco
atrobrunneo et margine brunneo usque cinereo-brunneo.
Contextus cinereo-creameus, tenuis. Lamellae annexae
usque liberae, subdistantes (12-18) cum 2-3 seriebus
lamellularum, latae (1-2 mm), olivaceo-brunneae usque
cinereo-olivaceae vel flavo-olivaceae, saepe brunneomarginatae usque olivaceo-brunneomarginatae, haud
intervenosae. Stipes 15-55 × 0.5-1 mm, centralis,
cylindratus, filo metallico similis, cavus, glaber, haud
insititius, apice flavo-albo, basi brunnea usque rubro
brunnea. Odor saporque non propria. Basidiosporae (11)12-14(-15) × 4-5(-6) µm, fusoideae, inaequilaterales,
laeves, hyalinae, inamyloideae, tenuitunicatae Basidia
20-22 × 6-8 µm,cylindrata usque clavata, 4-spora.
Basidiolae fusoideae usque clavatae. Cheilocystidia
abundantia, typi Sicci; 12-28 × 5-9 µm, cylindrata usque
clavata, interdum catenulatus, hyalina, inamyloidea,
tenuitunicata; setulis apicalibus 2-9(-10) × 1-1.5 µm,
cylindratis,
subacutis,
flavis
usque
brunneis,
crassetunicatis. Pleurocystidia 2 typorum: a) cellulis typi
Sicci, cheilocystidio similibus, dispersis, raris, 12-22 ×
6-8 µm, cylindratis usque clavatis, hyalinis, inamyloideis,
tenuitunicatis; setulis apicalibus 2-7 × 1-1.5 µm,
cylindratis, subacutis, flavis, inamyloideis, crassetunicatis; b) cellulis haud setulosis 30-50 × 5-9 µm,
cylindratis usque clavatis, constrictis et interdum lobatis
ad apicem, hyalinis, inamyloideis, tenuitunicatis.
Pileipellis hymeniformis, maculosus, typi Sicci; 8-23 ×
5-12 µm, cylindratus usque clavatus vel pyriformis,
saepe ramosus, hyalinus usque pallide flavus,
inamyloideus, tenui- usque crassetunicatus; setulis
apicalibus 2-7(-12) × 1-1.5(-2.5) µm, coartatis,
cylindratis, subacutis usque acutis, brunneis usque
atrobrunneis, crassetunicatis. Trama pilei intertexta,
dextrinoidea usque fortiter dextrinoidea. Trama lamellae
intertexta usque regularis, hyphis 3-8(-22) µm diametro,
cylindratis
usque
inflatis,
laevibus,
hyalinis,
dextrinoideis usque fortiter dextrinoideis, tenuitunicatis,
haud gelatinosis. Stipitipellis subparallelus, hyphis 3-9(13) µm diametro, cylindratis, brunneis usque laete
brunneis, laevibus, dextrinoideis, crassetunicatis (usque
ad 3 µm), haud gelatinosis. Trama stipitis parallela,
hyphis (3-)5-13(-20) µm diametro, cylindratis, hyalinis
usque pallide flavis, laevibus, dextrinoideis, tenuitunicatis, haud gelatinosis. Caulocystidia nulla. Fibulae
praesentes in omnibus texturis.
Typus: Chiang Mai Province, Doi Suthep-Pui
National Park, Sangasabhasri Land on the way to Huai
Kok Ma Village, N 18º 48.4’ E 98º 54.6’ alt. 1,146 m.,
dispersus usque gregarius in folis plantae dicotyledoneae
vel ligno, 14 August 2003, N. Wannathes 112 (CMU:
holotypus; SFSU: isotypus).
Pileus 6-23 mm diam., convex with a
rugulose umbo when young, expanding to
plano-convex and slightly depressed in age,
striate to sulcate, pruinose, dull, dark brown
overall when young, in age disc dark brown
Fig. 61. Marasmius brunneoolivascens (N. Wannathes 227).
1. Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5a. Siccus-type pleurocystidia 5b. Nonsetulose pleurocystidia 6. Pileipellis, Scale bar 1 = 20
mm, 2-6 = 10 µm
and margin brown to greyish brown. Context
greyish cream, thin. Lamellae adnexed to free,
subdistant (12-18) with 2-3 series of lamellulae,
broad (1-2 mm), olive brown to greyish olive
or yellowish olive, sometimes with brown to
olive-brown edges, non-intervenose. Stipe 1555 × 0.5-1 mm, central, cylindrical, wiry,
hollow, glabrous, non-insititious, apex
yellowish white, base brown to reddish brown.
Odor and taste not distinctive.
Basidiospores (11-)12-14(-15) × 4-5(-6)
µm [xmr = 12.6-13.8 × 4.0-5.0, xmm = 13.0 ± 0.3
× 4.4 ± 0.4 µm, µm, Qmr = 2.7-3.4, Qmm = 3.0 ±
0.2, n = 25 spores, s = 11 specimens], fusoid,
inequilateral, smooth, hyaline, inamyloid, thinwalled. Basidia20-22 × 6-8 µm, cylindrical to
clavate, 4-spored. Basidioles fusoid to clavate.
Cheilocystidia abundant, of Siccus-type broom
cells; main body 12-28 × 5-9 µm, cylindrical to
287
clavate, sometimes catenulate, hyaline,
inamyloid, thin-walled; apical setulae 2-9(-10)
× 1-1.5 µm, cylindrical, subacute, yellow to
brown, thick-walled. Pleurocystidia composed
of 2 type of cells: a) Siccus-type broom cells
like the cheilocystidia, scattered and rare; main
body 12-22 × 6-8 µm, cylindrical to clavate,
hyaline, inamyloid, thin-walled; apical setulae
2-7 × 1-1.5 µm, cylindrical, subacute, yellow,
inamyloid, thick-walled; b) non-setulose cells,
30-50 × 5-9 µm, cylindrical to clavate,
constricted and sometimes lobed at the apex,
hyaline, inamyloid, thin-walled. Pileipellis
hymeniform, mottled, composed of Siccus-type
broom cells; main body 8-23 × 5-12 µm,
cylindrical to clavate or pyriform, often
branched, hyaline to pale yellow, inamyloid,
thin-to thick-walled; apical setulae 2-7(-12) ×
1-1.5(-2.5) µm, crowded, cylindrical, subacute
to acute, brown to dark brown, thick-walled.
Pileus trama interwoven, dextrinoid to strongly
dextrinoid. Lamellar trama hyphae 3-8(-22)
µm diam., interwoven to regular, cylindrical to
inflated, smooth, hyaline, dextrinoid to strongly
dextrinoid,
thin-walled,
non-gelatinous.
Stipitipellis hyphae 3-9(-13) µm diam.,
subparallel, cylindrical, brown to light brown,
smooth, dextrinoid, thick-walled (up to 3 µm),
non-gelatinous. Stipe trama hyphae (3-)5-13(20) µm diam., parallel, cylindrical, hyaline to
pale yellow, smooth, dextrinoid, thin-walled,
non-gelatinous. Caulocystidia absent. Clamp
connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves or on wood, Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Sangasabhasri
Land on the way to Huai Kok Ma Village, N 18º 48.4’ E
98º 54.6’ alt. 1,146 m., 29 July 2003, N. Wannathes 055
(CMU, SFSU); same location, 14 August 2003, N.
Wannathes 112 (CMU: holotype; SFSU: isotype) and
N.Wannathes 113 (CMU); same location, 16 July 2004,
N. Wannathes 228 (CMU); same location, 24 June
2005,N. Wannathes 322 (CMU); Chiang Mai Province,
Mae Taeng District, Tung Joaw Village, N 19º 8.07’ E
98º 38.9’ alt. 1423 m., 31 July 2003, N. Wannathes 063
(CMU, SFSU); same location, 22 August 2004, N.
Wannathes 277 (CMU, SFSU); Chiang Mai Province,
Huai Nam Dang National Park, North 19º 18.3’ E 98º
35.8’ alt. 1,538 m., 29 June 2005, N. Wannathes 373
(CMU, SFSU); Chiang Mai Province, Doi Suthep-Pui
National Park, Medicinal Plant Garden, 3 August 2005,
N. Wannathes 397 (CMU, SFSU).
288
Discussion: Marasmius brunneoolivascens is characterized by a striate to sulcate
pileus coloured brown to greyish brown with a
dark brown disc, subdistant (12-18) broad
lamellae colored olive brown to greyish olive
or yellowish olive with or without brown to
olive brown edges, a glabrous stipe, fusoid
basidiospores with mean 13.0 × 4.4 µm and
mean Q = 3, two types of pleurocystidia
(simple strangulate cells and scattered Siccustype broom cells), and an absence of
caulocystidia. This new species is similar to
two African species. Marasmius grandisetulosus Singer differs in forming white to creamcoloured lamellae, longer basidiospores (17-23
µm long) and lacks olive tones (Singer, 1964;
Antonín, 2007). Marasmius elaeocephalus
Singer sensu Antonín (2007) differs in forming
an olive brown pileus with yellow tones on the
margin, yellowish white lamellae with
concolorous edges, and lacks two types of
pleurocystidia.
53. Marasmius
suthepensis
Desjardin & Lumyong, sp. nov.
Wannathes,
(Figs 59-8, 62)
MycoBank: MB512429
Etymology: ‘suthep’ = Doi Suthep-Pui National
Park; referring to the place where the holotype specimen
was collected.
Pileus 10-27 mm diametro, obtuse conicus usque
convexus, cum vel sine umbone, laevis usque striatus,
glaber, hebetatus, disco clare brunneo, margine brunneoflavo usque brunneo-aurantiaco. Contextus flavo-albus,
tenuis. Lamellae annexae usque liberae, subdistantes
(12-18) cum 3-4 seriebus lamellularum, latae (1-2 mm),
pallide flavo-albae, haud marginatae, haud intervenosae.
Stipes 18-55 × 1 mm, centralis, cylindratus, filo
metallico similis, cavus, glaber, haud insititius, apice
flavo-albo, basi brunnea. Odor saporque non propria.
Basidiosporae (10-)11-14(-15) × (3.5-)4-5 µm, anguste
ellipsoideae usque fusoideae, laeves, hyalinae,
inamyloideae, tenuitunicatae Basidia 13-15 × 5-6 µm,
cylindrata usque clavata, 4-spora. Basidiolae cylindratae
usque clavatae. Cheilocystidia abundantia, typi Sicci; (6)12-18 × (3-)7-11 µm, cylindrata usque clavata, hyalina,
inamyloidea, tenuitunicata; setulis apicalibus 2-6 × 1 µm,
cylindratis usque conicis, obtusis usque subacutis, flavis
usque flavo-brunneis, tenuitunicatis. Pleurocystidia
vulgaria, 27-43 × 5-7 µm, cylindrata usque fusoidea,
undulata usque constricta et interdum lobata ad apicem,
hyalina, inamyloidea, tenuitunicata. Pileipellis hymeniformis, maculosus, typi Sicci; (7-)10-17(-22) × 4-10 µm,
cylindratus usque clavatus vel pyriformis, hyalinus
usque pallide flavus, inamyloideus, tenuitunicatus;
setulis apicalibus 3-8 × 1(-1.5) µm, coartatis, cylindratis,
subacutis, brunneis usque laete brunneis, crassetunicatis.
Fungal Diversity
Trama pilei intertexta, hyphis (4-)6-13(-17) µm diametro,
cylindratis usque inflatis, laevibus, hyalinis, fortiter
dextrinoideis, tenuitunicatis. Trama lamellae intertexta
usque regularis, hyphis 3-7(-12) µm diametro,
cylindratis
usque
inflatis,
laevibus,
hyalinis,
dextrinoideis,
tenuitunicatis,
haud
gelatinosis.
Stipitipellis subparallelus, hyphis 3-7(-9) µm diametro,
cylindratis, brunneis usque atrobrunneis, laevibus,
inamyloideis usque leniter dextrinoideis, tenuitunicatis,
haud gelatinosis. Trama stipitis parallela, hyphis (3-)510(-13) µm diametro, cylindratis, hyalinis, laevibus,
dextrinoideis,
tenuitunicatis,
haud
gelatinosis.
Caulocystidia nulla. Fibulae praesentes in omnibus
texturis.
Typus: Chiang Mai Province, Doi Suthep-Pui
National Park, Mokfa Waterfall, on Hwy 1095, N19º
6.5’.E 98º 46.3’ alt. 1014 m., dispersus usque gregarius
in folis plantae dicotyledoneae vel ligno, 29 June 2004,
Y.S. Tan 280 (CMU: holotypus; SFSU: isotypus).
Pileus 10-27 mm diam., obtusely conical
to convex, with or without an umbo, smooth to
striate, glabrous, dull, disc bright brown,
margin brownish yellow to brownish orange.
Context thin, yellowish white. Lamellae
adnexed to free, subdistant (12-18) with 3-4
series of lamellulae, broad (1-2 mm), pale
yellowish
white,
non-marginate,
nonintervenose. Stipe 18-55 × 1 mm, central,
cylindrical, wiry, hollow, glabrous, noninsititious, apex yellowish white, base brown.
Odor and taste not distinctive.
Basidiospores (10-)11-14(-15) × (3.5-)45 µm [xmr = 11.7-13.6 × 3.9-4.1 µm, xmm =
12.6 ± 1.0 × 4.0 ± 0.1 µm, Qmr = 2.9-3.8, Qmm
= 3.7 ± 1.0, n = 25 spores, s = 3 specimens],
narrowly ellipsoid to fusoid, smooth, hyaline,
inamyloid, thin-walled. Basidia 13-15 × 5-6
µm, cylindrical to clavate, 4-spored. Basidioles
cylindrical to clavate. Cheilocystidia abundant,
of Siccus-type broom cells; main body (6-)1218 × (3-)7-11 µm, cylindrical to clavate,
hyaline, inamyloid, thin-walled; apical setulae
2-6 × 1 µm, cylindrical to conical, obtuse to
subacute, yellow to yellowish brown, thinwalled. Pleurocystidia common, 27-43 × 5-7
µm, cylindrical to fusoid, wavy to constricted
and sometimes lobed at the apex, hyaline,
inamyloid, thin-walled. Pileipellis hymeniform,
mottled, composed of Siccus-type broom cells;
main body (7-)10-17(-22) × 4-10 µm, cylindrical
to clavate or pyriform, hyaline to pale yellow,
inamyloid, thin-walled; apical setulae 3-8 × 1(1.5) µm, crowded, cylindrical, subacute, brown
to light brown, thick-walled. Pileus trama
Fig. 62. Marasmius suthepensis (Y.S. Tan 280). 1.
Basidiomes 2. Basidia 3. Basidiospores 4. Cheilocystidia
5. .Pleurocystidia 6. Pileipellis, Scale bar 1 = 20 mm, 2-6
= 10 µm
hyphae (4-)6-13(-17) µm diam., interwoven,
cylindrical to inflated, smooth, hyaline,
strongly dextrinoid, thin-walled. Lamellar
hyphae 3-7(-12) µm diam., trama interwoven
to regular, cylindrical to inflated, smooth,
hyaline, dextrinoid, thin-walled, non-gelatinous.
Stipitipellis hyphae 3-7(-9) µm diam.,
subparallel, cylindrical, brown to dark brown,
smooth, inamyloid to weakly dextrinoid, thinwalled, non-gelatinous. Stipe trama hyphae (3)5-10(-13) µm diam., parallel, cylindrical,
hyaline, smooth, dextrinoid, thin-walled, nongelatinous. Caulocystidia absent. Clamp
connections present in all tissues.
Habit, habitat and known distribution.
Scattered to gregarious on dicotyledonous
leaves or on wood, Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Sangasabhasri
Land on the way to Huai Kok Ma Village, N 18º 48.4’ E
98º 54.6’ alt. 1,146 m., 30 June 2003, N. Wannathes 012
(CMU, SFSU); same location, 29 July 2003, N.
Wannathes 057 (CMU, SFSU); Chiang Mai Province,
Doi Suthep-Pui National Park, Mokfa Waterfall, on Hwy
1095, N19º 6.5’.E 98º 46.3’ alt. 1014 m., 29 June 2004,
Y.S. Tan 280 (CMU: holotype; SFSU: isotype).
Discussion: Diagnostic features of
Marasmius suthepensis include: a smooth to
striate pileus coloured brownish yellow to
brownish orange with bright brown disc;
subdistant (12-18), broad lamellae that are pale
yellowish white and non-marginate; a glabrous,
brown stipe; basidiospores with mean 12.6 ×
289
4.0 µm and mean Q = 3.7; simple cylindrical
pleurocystidia, and an absence of caulocystidia.
The Thai new species is similar to two browncoloured neotropical species. Marasmius
hinnuleus Berk. & M.A. Curtis, described from
Cuba, differs in forming smaller (4-12 mm vs
10-27 mm), sulcate to plicate pilei coloured
pale reddish brown to cinnamon-rufous, and
slightly longer spores with mean 14 × 3.8 µm
(Singer, 1976; Pegler, 1983). Marasmius
helvolus Berk., described from Brazil, differs
in forming distinctly sulcate to plicate pilei,
fewer lamellae (9-11) and usually lacks
pleurocystidia (rare in some basidiomes)
(Singer, 1976; Pegler, 1983; Desjardin and
Ovrebo, 2006). Marasmius suthepensis is also
similar to M. brunneoolivascens (see above),
which may occur at the same site, but the latter
species differs in forming olive toned lamellae
and Siccus-type broom cells are often present
as pleurocystidia on the lamellar sides.
54. Marasmius haematocephalus (Mont.) Fr.,
Epicr. Syst. Mycol: 382. 1838.
≡ Agaricus haematocephalus Mont. Ann. Sci.
Nat., Bot., sér. 2, 8: 369. 1837.
≡ Androsaceus haematocephalus (Mont.) Pat., J.
Bot. (Morot) 3: 336. 1889.
= Marasmius rhodocephalus Fr., Nova Acta
Regiae Soc. Sci. Upsal 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.
There occur in Thailand six distinct
forms of M. haematocephalus, that share
basidiospores in the range 19-25 × 3.5-5 µm
with xmm = 20.7 × 4.0 µm and Qmm = 5.2, welldeveloped, simple and strangulate pleurocystidia, two types of cheilocystidia, lack
caulocystidia, and fall into a well-supported
clade based on ITS sequences (Fig. 1). They
differ in pileus size and pigmentation, ranging
from yellowish orange to deep reddish brown,
maroon, dark violet, purple and purplish brown,
and in substrate preference. The ITS sequence
data, however, are of limited use in
distinguishing the forms. A branch and bound
analysis that includes sequences of all M.
290
haematocephalus forms with M. siccus as an
outgroup (Fig. 70) indicates that several forms
are well-supported by BS values 92-100%
(forms
haematocephalus,
“atrobrunneus”,
“violaceus”), whereas other forms show
invariability (forms “variabilis”, “luteocephalus”) or form a polytomy (“f. robustus”).
Marasmius haematocephalus as it occurs in
Southeast Asia will remain a complex of
morphologically and genetically distinct
entities without formal taxonomic recognition
until further specimens are analyzed in the
hopes of clarifying their differences and
relationships. The forms recognized below will
not be formally proposed and are named herein
only to aid in documenting their existence.
Key to forms of Marasmius haematocephalus
1. Pileus yellowish orange or with a pale brownish
yellow disc and nearly white margin .........................
......... 54A. M. haematocephalus “f. luteocephalus”
1. Pileus with red, violet, olive, greyish blue or brown
pigments................................................................... 2
2. Pileus 1-5 mm diam, dark brown overall, lacking red,
violet or purple...........................................................
..........54B. M. haematocephalus “f. atrobrunneus”
2. Pileus > 5 mm diam, with red, violet, purple, olive, or
greyish blue tones ................................................... 3
3. Pileus olive to greyish blue overall when young, disc
often with red, orange or reddish purple tones in age
.................54C. M. haematocephalus “f. variabilis”
3. Pileus lacking olive and greyish blue tones ............. 4
4. Pileus typically 10-20(-30) mm diam., dark reddish
brown to dark violet brown (10-11F6-8) when young,
fading to reddish-greyish brown in age......................
..................54D. M. haematocephalus “f. robustus”
4. Pileus typically <10 mm diam, deep reddish brown to
maroon or deep violet-purple................................... 5
5. Pileus magenta, deep violet-purple or dark purple
(13-14E-F6-8); basidiospores normally with mean in
the range 20.5-22 × 3.8-4.2 µm; basidiomes often
associated with bamboo leaves .................................
.................54E. M. haematocephalus “f. violaceus”
5. Pileus deep reddish brown to maroon (9-11E-F7-8);
basidiospores normally with mean in the range 17.518 × 4.0-4.3 µm; basidiomes usually associated with
dicot leaves ................................................................
....... 54F. M. haematocephalus f. haematocephalus
54A Marasmius
haematocephalus
“f.
luteocephalus” Wannathes, Desjardin &
Lumyong, nom. prov.
(Figs 63, 69-1)
Fungal Diversity
Pileus 4-21 mm diam., convex to planoconvex, sulcate, pruinose, dull, yellowish
orange overall or with disc pale brownish
yellow and margin yellowish orange to
yellowish white or nearly white. Context cream,
thin. Lamellae free to adnexed, distant (11-13),
broad (1-3 mm), cream with yellowish orange
edges, non-intervenose. Stipe 20-42 × 0.3-0.8
mm, central, cylindrical, wiry, hollow, glabrous,
non-insititious, apex yellowish white, base dark
brown. Odor and taste not distinctive.
Basidiospores (20-)21-23(-24) × 3.5-4
µm [x = 22 ± 0.9 × 3.7 ± 0.3 µm, Q = 5.0-6.6,
Qm = 5.9, n = 25 spores, s = 1 specimen],
clavate to fusoid, often curved in profile,
smooth, hyaline, inamyloid, thin-walled.
Basidia not observed. Basidioles fusoid to
clavate. Cheilocystidia composed of 2 types of
cells: a) Siccus-type broom cells with main
body 12-19 × 7-9 µm, cylindrical to clavate,
hyaline, inamyloid, thin-walled; apical setulae
2-5 × 1 µm, cylindrical, often forked, subacute
to obtuse, hyaline to yellow, thick-walled; b)
non-setulose cells like the pleurocystidia, 36-41
× 11-13 µm, fusoid, usually attenuated and
strangulate at the apex, hyaline, inamyloid,
thin-walled. Pleurocystidia common, 52-58(62) × 10-12 µm, fusoid to clavate, usually
attenuated and strangulate at the apex, hyaline,
inamyloid, thin-walled. Pileipellis hymeniform,
mottled, composed of Siccus-type broom cells;
main body 12-15× 8-14 µm, clavate to broadly
clavate or pyriform, sometime branched,
yellow to hyaline, inamyloid, thin-walled;
apical setulae 2-4 × 1 µm, cylindrical, subacute,
yellow to pale yellow, thick-walled. Pileus
trama interwoven, dextrinoid. Lamellar trama
hyphae 5-10(-20) µm diam., interwoven,
cylindrical to inflated, smooth, hyaline,
dextrinoid,
thin-walled,
non-gelatinous.
Stipitipellis hyphae 3-5 µm diam., subparallel,
cylindrical, greenish brown, smooth, dextrinoid,
thin- to thick-walled, non-gelatinous. Stipe
trama hyphae (4-)7-10 µm diam., parallel,
cylindrical, hyaline, smooth, dextrinoid, thinwalled, non-gelatinous. Caulocystidia absent.
Clamp connections present.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, Northern Thailand.
Fig. 63. Marasmius haematocephalus “f. luteocephalus” (N.
Wannathes 310). 1. Basidiomes 2. Basidiospores 3a.
Siccus-type
cheilocystidia
3b.
Non-setulose
cheilocystidia 4. Pleurocystidia 5. Pileipellis, Scale bar 1
= 10 mm, 2-5 = 10 µm
Material examined: Thailand. Chiang Rai
Province, Muang District, Khun Con Waterfall, 12 June
2005, N. Wannathes 310 (CMU, SFSU).
Discussion: Marasmius haematocephalus
“f. luteocephalus” differs from all other forms
in having a yellowish orange to yellowish
white pileus with pale brownish yellow disc
and cream-coloured lamellae with yellowish
orange edges. This provisional taxon looks like
a paler form of M. haematocephalus var.
leucophyllus Singer, described from Bolivia,
but the latter differs in forming a darker pileus
(fawn to pinkish cinnamon), and smaller
basidiospores in the range 15-20 × 3.5 µm
(Singer, 1976).
54B. Marasmius
haematocephalus
“f.
atrobrunneus” Wannathes, Desjardin &
Lumyong, nom. prov.
(Fig. 64)
Pileus 1-5 mm diam., obtusely conical to
convex with or without a rugulose disc, striate,
291
minutely velutinous, dull, dark brown overall.
Context brownish white, thin. Lamellae
adnexed, distant (8-11), narrow, brownish
cream, non-marginate, non-intervenose. Stipe
5-15 × 0.2 mm, central, cylindrical, wiry,
hollow, glabrous, insititious, apex yellowish
white, base brown to dark brown. Odor and
taste not distinctive.
Basidiospores 17-19 × 4-5 µm [x = 18.1
± 0.7 × 4.2 ± 0.3 µm, Q = 3.8-4.8, Qm = 4.3, n
= 25 spores, s = 1 specimen], clavate to fusoid,
often curved in profile, smooth, hyaline,
inamyloid, thin-walled. Basidia not observed.
Basidioles fusoid to clavate. Cheilocystidia
composed of 2 types of cells: a) Siccus-type
broom cells with main body 6-14 × 5-12 µm,
cylindrical to clavate or pyriform, hyaline,
inamyloid, thin-walled; apical setulae 2-4 × 1
µm, cylindrical, often forked, subacute, hyaline
to yellow, thick-walled; b) non-setulose cells
like the pleurocystidia, 35-40 × 7-9 µm, fusoid
to clavate, usually attenuated and strangulate at
the apex, often with 2-4 successive apical
constrictions, hyaline inamyloid, thin-walled.
Pleurocystidia common, 34-48 × 7-9 µm,
clavate to fusoid, usually attenuated and
strangulate at the apex, sometimes with 2
successive apical constrictions, hyaline,
inamyloid, thin-walled. Pileipellis hymeniform,
mottled, composed of Siccus-type broom cells;
main body 12-16 × 7-10 µm, clavate to broadly
clavate or pyriform, sometime branched,
yellow to hyaline, inamyloid, thin- to thickwalled; apical setulae 2-5 × 1 µm, cylindrical,
subacute, brown to dark brown, thick-walled.
Pileus trama interwoven, inamyloid. Lamellar
trama hyphae 4-6 µm diam., regular to
interwoven, cylindrical, smooth, hyaline,
inamyloid,
thin-walled,
non-gelatinous.
Stipitipellis hyphae 3-5 µm diam., parallel,
cylindrical, brown to greenish brown, smooth,
inamyloid, thin- to thick-walled (up to 0.5 µm),
non-gelatinous. Stipe trama hyphae 2-5 µm
diam., parallel, cylindrical, hyaline, smooth,
weakly dextrinoid, thin-walled, non-gelatinous.
Caulocystidia absent. Clamp connections
present.
Habit, habitat and known distribution:
Scattered to gregarious on grass leaves,
Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Mae Taeng District, Street to Pathummigaram
292
Fig. 64. Marasmius haematocephalus “f. atrobrunneus” (N.
Wannathes 117). 1. Basidiomes 2. Basidiospores 3a.
Siccus-type
cheilocystidia
3b.
Non-setulose
cheilocystidia 4. Pleurocystidia 5. Pileipellis, Scale bar 1
= 10 mm, 2-5 = 10 µm
temple in Padeng Village, 22 August 2003, N.
Wannathes 117 (CMU, SFSU).
Discussion: Marasmius haematocephalus “f.
atrobrunneus” differs from all other forms of
this species in forming a smaller (1- 5 mm
diam.) dark brown pilei without red, violet,
purple or yellowish orange tones.
54C. Marasmius
haematocephalus
“f.
variabilis” Wannathes, Desjardin & Lumyong,
nom. prov.
(Figs 65, 69-3)
Pileus 5-12 mm diam., convex, sulcate,
pruinose, dull, when young olive to greyish
blue overall or disc dark purple and margin
brownish purple with dark brownish grey to
dark olive sulcae, when mature disc dark
purple and margin fading to reddish brown or
purplish brown with greyish blue sulcae.
Context brownish grey, thin. Lamellae adnexed,
subdistant (10-13), narrow to broad (1-2 mm),
pinkish cream with or without pinkish brown
edges, non-intervenose. Stipe 20-32 × 0.5-1
mm, central, cylindrical, wiry, hollow, glabrous,
non-insititious, apex purple, base dark
brownish purple. Odor and taste not distinctive.
Basidiospores (17-)19-22(-24) × 3.5-5(5.5) µm [xmr = 20.3-21.6 × 3.9-4.2 µm, xmm =
21.0 ± 0.9 × 4.1 ± 0.3 µm, Qmr = 5.1-5.3, Qmm
= 5.2 ± 0.1, n = 25 spores, s = 2 specimens],
clavate to fusoid, often curved in profile,
smooth, hyaline, inamyloid, thin-walled.
Basidia not observed. Basidioles fusoid to
Fungal Diversity
clavate. Cheilocystidia composed of 2 types of
cells: a) Siccus-type broom cells with main
body 15-20 × 5-7 µm, cylindrical to clavate,
hyaline, inamyloid, thin-walled; apical setulae
2-10 × 1(-1.5) µm, cylindrical, subacute to
acute, greenish brown, thick-walled; b) nonsetulose cells like the pleurocystidia, 40-52× 78 µm, fusoid to clavate, usually attenuated and
strangulate at the apex, hyaline, inamyloid,
thin-walled. Pleurocystidia common, 54-67 ×
10-12 µm, clavate to fusoid, usually attenuated
and strangulate at the apex, hyaline, inamyloid,
thin-walled. Pileipellis hymeniform, not
mottled, composed of Siccus-type broom cells;
main body 10-16 × 6-8 µm, cylindrical to
clavate or broadly clavate, hyaline, inamyloid,
thin-walled; apical setulae 4-7 × 1 µm,
cylindrical, subacute to acute, dark greenish
brown, thick-walled. Pileus trama interwoven,
weakly dextrinoid. Lamella trama hyphae 3-9
µm diam., interwoven, cylindrical, smooth,
hyaline, weakly dextriomoid, thin-walled, nongelatinous. Stipitipellis hyphae 3-5 µm diam.,
subparallel, cylindrical, brown, smooth,
inamyloid to weakly dextrinoid, thin- to thickwalled (up to 1 µm), non-gelatinous. Stipe
trama hyphae 3-6 µm diam., parallel,
cylindrical, hyaline, smooth, inamyloid to
weakly dextrinoid, thin-walled, non-gelatinous.
Caulocystidia absent. Clamp connections
present.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous or
bamboo leaves, Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mokfa Waterfall,
on Hwy 1095, N19º 6.5’ E 98º 46.3’ alt. 1014 m., 25
june 2005, N. Wannathes 336 (CMU, SFSU); Phrae
Province, Muang District, Cherng Thong Waterfall, 16
August 2005, N. Wannathes 415 (CMU); Phrae Province,
Muang District, Pa Dang, Nong Kam Village, near Nong
Kam reservoir, 18 August 2005, N. Wannathes 430
(CMU, SFSU).
Discussion: Marasmius haematocephalus “f.
variabilis” differs from the other forms of M.
haematocephalus in forming pilei that display
the full rainbow of colours found in other
forms throughout development, including blue,
olive, red, violet, purple and grey. However, “f.
variabilis” still forms pinkish cream lamellae
with or without pinkish brown edges, and a
stipe that has a purple apex and dark brownish
purple base, colors characteristic of f.
Fig. 65. Marasmius haematocephalus “f. variabilis” (N.
Wannathes 433). 1. Basidiomes 2. Basidiospores 3a.
Siccus-type
cheilocystidia
3b.
Non-setulose
cheilocystidia 4. Pleurocystidia 5. Pileipellis, Scale bar 1
= 10 mm, 2-5 = 10 µm
haematocephalus.
54D. Marasmius haematocephalus “f.
robustus” Wannathes, Desjardin & Lumyong,
nom. prov.
(Figs 66, 69-2)
Pileus 10-20(-30) mm diam., obtusely
conical when young, convex to plano-convex
in age, disc rugulose, sulcate to plicate,
glabrous, dull, dark reddish brown to dark
violet brown (10-11F6-8) when young, fading
to reddish-greyish brown to greyish purple in
age. Context greyish cream, thin. Lamellae
adnexed, distant to subdistant (9-15), broad (24 mm), yellowish grey (2B2) to greyish yellow
(4B3), non-marginate or brown-marginate,
non-intervenose. Stipe 13-43(-62) × 0.5-1 mm,
central, cylindrical, wiry, hollow, glabrous,
non-insititious, apex greyish white, base dark
brown. Odor and taste not distinctive.
Basidiospores (16-)18-23(-24) × (3.5-)45(-6) µm [xmr = 18.5-21.6 × 3.9-4.5 µm, xmm =
20.0 ± 1.3 × 4.1 ± 0.1 µm, Qmr = 4.2-5.5, Qmm
= 4.9 ± 0.4, n = 25 spores, s = 9 specimens],
293
clavate to fusoid, often curved in profile,
smooth, hyaline, inamyloid, thin-walled.
Basidia not observed. Basidioles fusoid to
clavate. Cheilocystidia composed of 2 types of
cells: a) Siccus-type broom cells with main
body (7-)11-19(-23) × 6-11 µm, cylindrical to
clavate, hyaline, inamyloid, thin-walled; apical
setulae 2-6(-10) × 1µm, cylindrical, subacute to
acute, yellow, thin- to thick-walled; b) nonsetulose cells like the pleurocystidia, 35-53× 812 µm, fusoid to clavate, usually attenuated
and strangulate at the apex, hyaline, inamyloid,
thin-walled. Pleurocystidia common, 38-62 ×
6-15 µm, clavate to fusoid, usually attenuated
and strangulate at the apex, hyaline, inamyloid,
thin-walled. Pileipellis hymeniform, not
mottled, composed of Siccus-type broom cells;
main body 10-28 × 6-13 µm, cylindrical to
clavate or broadly clavate, hyaline, inamyloid,
thin- to thick-walled; apical setulae 2-5(-7) × 1
µm, cylindrical, subacute to acute, brown to
dark brown, thick-walled. Pileus trama
interwoven, dextrinoid. Lamellar trama hyphae
(2-)4-6(-14) µm diam., interwoven, cylindrical
to inflated, smooth, hyaline, weakly dextrinoid,
thin-walled,
non-gelatinous.
Stipitipellis
hyphae 3-9(-12) µm diam., subparallel,
cylindrical, brown to dark brown, smooth,
weakly dextrinoid, thin- to thick-walled (up to
1 µm), non-gelatinous. Stipe trama hyphae 312 µm diam., parallel, cylindrical, hyaline,
smooth, dextrinoid, thin-walled, non-gelatinous.
Caulocystidia absent. Clamp connections
present.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves or wood, Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Sangasabhasri
Land on the way to Huai Kok Ma Village, N 18º 48.4’ E
98º 54.6’ alt. 1,146 m., 29 July 2003, N. Wannathes 053
(CMU, SFSU); same location, 14 August 2003, N.
Wannathes 110 and 111, (CMU, SFSU); Chiang Mai
Province, Doi Inthanon National Park, Junction of
Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E
98º 29.6’ alt. 1,703 m., 3 July 2004, N. Wannathes 203
(CMU, SFSU); Chiang Rai Province, Muang District,
Khun Con Waterfall, 12 June 2005, N. Wannathes 301
(CMU, SFSU); Chiang Mai Province, Doi Suthep-Pui
National Park, Medicinal Plat Garden, 16 June 2005, N.
Wannathes 314 (CMU, SFSU); Chiang Mai Province,
Doi Suthep-Pui National Park, Mokfa Waterfall, on Hwy
1095, N19º 6.5’.E 98º 46.3’ alt. 1014 m., 25 June 2005,
N. Wannathes 330 (CMU, SFSU); Phrae Province,
294
Fig. 66. Marasmius haematocephalus “f. robustus” (N.
Wannathes 433). 1. Basidiomes 2. Basidiospores 3a.
Siccus-type
cheilocystidia
3b.
Non-setulose
cheilocystidia 4. Pleurocystidia 5. Pileipellis, Scale bar 1
= 10 mm, 2-5 = 10 µm
Muang District, Cherng Thong Waterfall, 16 August
2005, N. Wannathes 417 (CMU, SFSU); same location,
19 August 2005, N. Wannathes 433 (CMU, SFSU).
Discussion: Marasmius haematocephalus “f.
robustus” is unique because of its robust pilei
(10-30 mm diam.) and range of pileus colours
that are similar to both forms haematocephalus
and “violaceus”. This provisional Thai form is
phenetically similar to M. haematocephalus var.
macrocephalus Singer, described from Bolivia,
except that the latter grows on monocot leaves
(Singer, 1976).
54E. Marasmius
haematocephalus
“f.
violaceus” Wannathes, Desjardin & Lumyoung,
nom. prov.
(Figs 67, 69-4)
Pileus 4-18 mm diam., obtusely conical
to convex, umbonate, sometimes with a
rugulose disc, striate to sulcate, pruinose to
finely velutinous, dull, dark purple (14F7) to
dark magenta (13F7) overall when young, in
age disc dark violet to greyish violet (14C4)
Fungal Diversity
and margin pinkish purple to violet. Context
purplish white (14A3), thin. Lamellae adnexed,
distant (7-12), narrow to broad (1-2 mm),
purplish pink (14A3), non-marginate, nonintervenose. Stipe 13-30 × 0.2-0.5 mm, central,
cylindrical, wiry, hollow, glabrous, noninsititious, apex purplish white, base dark
brown. Odor and taste not distinctive.
Fig. 67. Marasmius haematocephalus “f. violaceus” (N.
Wannathes 413). 1. Basidiomes 2. Basidiospores 3a.
Siccus-type
cheilocystidia
3b.
Non-setulose
cheilocystidia 4. Pleurocystidia 5. Pileipellis, Scale bar 1
= 10 mm, 2-5 = 10 µm
Basidiospores (19-)20-24(-25) × 3.5-5
µm [xmr = 20.6-21.8 × 3.8-4.2 µm, xmm = 21.4
± 0.6 × 4.0 ± 0.1 µm, Qmr = 5.3-5.5, Qmm = 5.4
± 0.1, n = 25 spores, s = 4 specimens], clavate
to fusoid, often curved in profile, smooth,
hyaline, inamyloid, thin-walled. Basidia not
observed. Basidioles fusoid to clavate.
Cheilocystidia composed of 2 types of cells: a)
Siccus-type broom cells with main body 10-21
× 5-11 µm, cylindrical to clavate or irregular in
outline, hyaline, inamyloid, thin-walled; apical
setulae 2-5(-14) × 1(-2) µm, cylindrical,
subacute to acute, yellow, thin- to thick-walled;
b) non-setulose cells like the pleurocystidia,
38-74× 8-10 µm, fusoid to clavate, usually
attenuated and strangulate at the apex, hyaline,
inamyloid,
thin-walled.
Pleurocystidia
common, 38-73 × 7-13 µm, clavate to fusoid,
usually attenuated and strangulate at the apex,
hyaline, inamyloid, thin-walled. Pileipellis
hymeniform, not mottled, composed of Siccustype broom cells; main body 10-17 × 4-10 µm,
cylindrical to clavate or broadly clavate,
hyaline, inamyloid, thin- to thick-walled; apical
setulae 2-5(-8) × 1 µm, cylindrical, subacute to
acute, brown, thick-walled. Pileus trama
interwoven, dextrinoid. Lamellar trama hyphae
3-8(-12) µm diam., interwoven, cylindrical,
smooth, hyaline, dextrinoid, thin-walled, nongelatinous. Stipitipellis hyphae 2-6(-8) µm
diam., subparallel, cylindrical, brown, smooth,
dextrinoid, thin- to thick-walled (up to 1 µm),
non-gelatinous. Stipe trama hyphae 3-8(-10)
µm diam., parallel, cylindrical, hyaline, smooth,
dextrinoid,
thin-walled,
non-gelatinous.
Caulocystidia absent. Clamp connections
present.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves or on bamboo debris, Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Mae Taeng District, New Waterfall, on 36 km.
marker of Hwy1095, 2 July 2004, N. Wannathes 193 and
195 (CMU, SFSU); Chiang Mai Province, Doi SuthepPui National Park, Mokfa Waterfall, on Hwy 1095, N19º
6.5’.E 98º 46.3’ alt. 1014 m., 25 June 2005, N.
Wannathes 339 (CMU, SFSU); Phrae Province, Muang
District, Cherng Thong Waterfall, 16 August 2005, N.
Wannathes 413 (CMU, SFSU).
Discussion: This taxon represents the
dark violet-purple form commonly encountered
in Southeast Asia and reported only as M.
haematocephalus from Papua New Guinea by
Desjardin and Horak (1997). It differs from f.
haematocephalus, as reported by Desjardin et
al. (2000) from Indonesia, in that the latter
form has deep reddish brown to maroon pilei
(lacking obvious deep purple and violet tones).
The provisional Thai form may be identical to
M. haematocephalus var. violaceus Singer,
described from Bolivia (Singer, 1976),
although we are hesitant to accept Southeast
Asian populations as representing a New
World taxon without data to support vicariance
or recent dispersal.
A bipolar mating system was reported for
this form of M. haematocephalus by Tan et al.
(2007) based on material from Malaysia.
295
54F. Marasmius haematocephalus (Mont.) Fr.
f. haematocephalus
(Figs 68, 69-5)
Pileus 3-10(-14) mm diam., obtusely
conical when young, convex with or without a
rugulose umbo in age, striate to sulcate,
pruinose, dull, disc deep reddish brown to
maroon, margin greyish red (11C6) to violet
brown (11D7). Context pinkish cream, thin.
Lamellae adnexed, distant (9-14) with 0-1
series of lamellulae, narrow, purplish cream,
non-marginate, non-intervenose. Stipe 10-40 ×
< 0.5 mm, central, cylindrical, wiry, hollow,
glabrous, non-insititious, apex light violet to
light purple, base brown to purplish brown.
Odor and taste not distinctive.
Basidiospores (15-)17-20(-25) × (3-)4-5
µm [xmr = 16.8-21.9 × 4.0-4.3 µm, xmm = 18.4
± 1.9 × 4.0 ± 0.1 µm, Qmr = 3.9-4.7, Qmm = 4.6
± 0.5, n = 25 spores, s = 6 specimens], clavate
to narrowly fusoid, often curved in profile,
smooth, hyaline, inamyloid, thin-walled.
Basidia 23-25 × 5-6 µm, cylindrical to clavate,
4-spored. Basidioles fusoid to clavate.
Cheilocystidia composed of 2 types of cells: a)
Siccus-type broom cells with main body 8-17 ×
5-11 µm, cylindrical to clavate, or pyriform,
often branched, hyaline, inamyloid, thin-walled;
apical setulae 1-4(-5) × 1 µm, cylindrical,
subacute, yellow to brownish yellow, thin- to
thick-walled; b) non-setulose cells like the
pleurocystidia, 25-41 × 7-12 µm, fusoid to
clavate, usually attenuated and strangulate at
the apex, often with 2-4 successive apical
constrictions, hyaline inamyloid, thin-walled.
Pleurocystidia common, 32-54 × 6-12(-15) µm,
clavate to fusoid, usually attenuated and
strangulate at the apex, sometimes with 2
successives apical constrictions, hyaline,
inamyloid, thin-walled. Pileipellis hymeniform,
mottled, composed of Siccus-type broom cells;
main body (6-)8-20 × 5-12 µm, clavate to
broadly clavate or pyriform, sometimes
branched, yellow to hyaline, inamyloid, thin- to
thick-walled; apical setulae 2-5 × 1(-1.5) µm,
cylindrical, subacute, brown to dark brown,
thick-walled. Pileus trama interwoven, weakly
dextrinoid to dextrinoid. Lamellar trama
hyphae 3-7(-12) µm diam., interwoven,
cylindrical to inflated, smooth, hyaline, weakly
dextrinoid to dextrinoid, thin-walled, non-
296
gelatinous. Stipitipellis hyphae 3-7 µm diam.,
parallel, cylindrical, brown to greenish brown,
smooth, dextrinoid, thin- to thick-walled (up to
1 µm), non-gelatinous. Stipe trama hyphae 35(-7) µm diam., parallel, cylindrical, hyaline,
smooth, dextrinoid, thin-walled, non-gelatinous.
Caulocystidia absent. Clamp connections
present.
Fig. 68. Marasmius haematocephalus f. haematocephalus
(N. Wannathes 428). 1. Basidiomes 2. Basidiospores 3a.
Siccus-type
cheilocystidia
3b.
Non-setulose
cheilocystidia 4. Pleurocystidia 5. Pileipellis, Scale bar 1
= 10 mm, 2-5 = 10 µm
Habit, habitat and known distribution.
Scattered to gregarious on dicotyledonous
leaves, Africa, neotropics, Java, Malaysia,
Papua New Guinea, Sri Lanka and Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mokfa
Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
m., 29 June 2004, Y.S. Tan 277 (CMU, SFSU); Chiang
Rai Province, Muang District, Horticulture Research
Center, 12 June 2005, N. Wannathes 296 (CMU, SFSU);
Chiang Mai Province, Doi Suthep-Pui National Park,
Medicinal Plant Garden, 16 June 2005, N. Wannathes
316 (CMU, SFSU); Phrae Province, Muang District,
Suen Keun, Na Koo Haa Waterfall, 15 August 2005, N.
Wannathes 409 (CMU, SFSU); Phrae Province, Muang
District, Cherng Thong Waterfall, 16 August 2005, N.
Wannathes 418 (CMU, SFSU), same location, 19
August2005, N. Wannathes 434 (CMU, SFSU); Phrae
Province, Muang District, Pa Dang, Nong Kam Village,
near Nong Kam reservoir, 18 August 2005, N.
Wannathes 428 (CMU, SFSU).
Fungal Diversity
Fig. 69. Basidiomata of forms of Marasmius haematocephalus. 1. M. haematocephalus “f. luteocephalus”
(N.Wannathes 310) 2. M. haematocephalus “f. robustus” (N.Wannathes 433) 3. M. haematocephalus “f. variabilis”
(N.Wannathes 430) 4. M. haematocephalus “f. violaceus” (N.Wannathes 413) 5. M. haematocephalus f.
haematocephalus (N.Wannathes 428), Scale bars = 20 mm.
Discussion: Marasmius haematocephalus, a
commonly reported pantropical species, is
characterized by a small (3-14 mm diam.),
obtusely conical to convex, striate to sulcate
pileus coloured deep reddish brown to maroon,
distant (9-14), narrow, pink or purplish cream
lamellae, a wiry, non-insititious, glabrous stipe
with light violet to purple apex and brown
base, clavate basidiospores with mean range
16.8-21.9 × 4.0-4.3 µm, numerous strangulate
pleurocystidia, and an absence of caulocystidia.
Singer (1976) recognized nine distinct varieties
of this species occurring in the New World
tropics distinguished mainly by pileus
pigmentation. As indicated above, we can
distinguish six distinct forms in Thailand, some
of which are nearly indistinguishable from the
varieties reported by Singer (1976) from the
neotropics. Until molecular data from
neotropical populations are compared with
297
M. haematocephalus f. haematocephalus NW409
M. haematocephalus f. haematocephalus NW296
99
M. haematocephalus f. haematocephalusTYS277
M. haematocephalus f. haematocephalus NW434
93
M. haematocephalus f. haematocephalus NW428
M. haematocephalus “f. atrobrunneus” NW117
M. haematocephalus “f. violaceus” NW339
98
M. haematocephalus “f. violaceus” W193
M. haematocephalus “f. violaceus” NW413
83
52
M. haematocephalus “f.variabilis” W430
M. haematocephalus “f. luteocephalus” NW310
M. haematocephalus “f. robustus” NW330
M. haematocephalus “f. robustus” NW433
5 changes
M. siccus DED 255
Fig. 70. Branch and bound bootstrap tree of Marasmius haematocephalus species complex. Numbers above branches
represent support from 2000 bootstrap replicates.
those of the Thai populations, we hesitate to
designate the Thai material with Singer's
epithets.
In the ML tree, 13 sequences of the
macromorphologically distinct yet micromorphologically indistinguishable M. haematocephalus formed a clade (AE) with strong
support (1.0 PP, 99% BS) and some internal
phylogenetic structure. A bootstrap branch and
bound analysis of 2000 replicates of a dataset
including all 13 M. haematocephalus
sequences with M. siccus as outgroup (Fig. 70),
recognized most of the macromorphological
entities as distinct clades. Representative
sequences conforming to forma haematocephalus formed a clade with 99% BS support;
those of “f. violaceus” formed a clade with
98% BS support; “f. variabilis” and “f.
luteocephalus” showed invariable sequences
298
and were sister taxa with 83% BS support;
whereas “f. robustus” was unresolved.
55. Marasmius hypophaeus Berk & M.A.
Curtis, J. Linn. Soc., Bot. 10: 298. 1868.
(Figs 71, 75-1)
Pileus 2-8 mm diam., paraboloid to
convex, disc rugulose, striate, pruinose, dull,
brown (7D8) to ferruginous when young,
greyish orange (6B4) in age. Context cream,
thin. Lamellae free to adnexed, distant (11-13),
narrow, cream with brownish orange edges,
non-intervenose. Stipe 23-35 × < 0.5 mm,
central, cylindrical, wiry, glabrous, insititious
to non-insititious, apex yellowish white, base
dark brown. Odor and taste not distinctive.
Basidiospores 16-18 × 4 µm [x = 17.2 ±
0.7 × 4.0 ± 0 µm, Q = 4.0-4.5, Qm = 4.2, n = 25
spores, s = 1 specimen], narrowly ellipsoid to
Fungal Diversity
hymeniform, weakly mottled, composed of
Siccus-type broom cells; main body 11-19 × 79 µm, cylindrical to clavate, pale yellow,
inamyloid, thin-walled; apical setulae 2-5 × 11.5 µm, crowded, cylindrical, often wavy,
subacute, brownish yellow to light brown, thinto thick-walled. Pileus trama interwoven,
dextrinoid. Lamellar trama hyphae (3-)4-7(-10)
µm diam., interwoven, cylindrical to inflated,
smooth, hyaline, dextrinoid, thin-walled, nongelatinous. Stipitipellis hyphae 3-4 µm diam.,
subparallel, cylindrical, yellowish brown to
light brown, smooth, weakly dextrinoid, thinwalled, non-gelatinous. Stipe trama hyphae 3-7
µm diam., parallel, cylindrical, hyaline, smooth,
weak dextrinoid, thin-walled, non-gelatinous.
Caulocystidia absent. Clamp connections
present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on monocotyledonous
or dicotyledonus leaves, Cuba, Java and
Northern Thailand.
Material examined: Thailand. Chiang Rai
Province, Khun Chae National Park, N 19º 4.4' E 99º
23.5' alt. 963 m., 10 June 2005, N. Wannathes 285
(CMU, SFSU)
Fig. 71. Marasmius hypophaeus (N. Wannathes 285). 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4a. Nonsetulose cheilocystidia 4b. Siccus-type pleurocystidia 5.
Pileipellis, Scale bar 1 = 20 mm, 2-5 = 10 µm
fusoid, smooth, hyaline, inamyloid, thin-walled.
Basidia not observed. Basidioles cylindrical to
clavate. Cheilocystidia common, of Siccus-type
broom cells; main body 10-15 × 6-8 µm,
cylindrical to clavate, hyaline, inamyloid, thinwalled; apical setulae 5-11 × 1(-3) µm,
cylindrical, wavy, often forked, conical to
subacute, yellow to brownish yellow, thin- to
thick-walled. Pleurocystidia composed of 2
type of cells: a) rarely present, Siccus broom
cells with main body 13-15 × 4-6 µm,
cylindrical, hyaline, inamyloid, thin-walled;
apical setulae 3-5 × 1 µm, cylindrical, conical
to subacute, hyaline, thick-walled; b) nonsetulose cells, common, 41-48 × 7-9 µm,
cylindrical to clavate, strangulate at the apex,
hyaline, inamyloid, thin-walled. Pileipellis
Discussion: Marasmius hypophaeus is
characterized by a small, striate pileus coloured
ferruginous to greyish orange, distant lamellae,
colored cream lamellae with brownish orange
edges, basidiospores with mean 17.2 × 4.0 µm,
and growth on both moncotyledonous and
dicotyledonous leaves. Described originally
from Cuba, this is the second report of the
species from the Old World tropics. Desjardin
et al. (2000) reported the species from Java.
56. Marasmius aff. pallescens Murrill, N.
Amer. Fl. 9(4): 261. 1915.
(Figs 72, 75-2)
Pileus 4-15 mm diam., obtusely conical
to convex, sometimes with a small papilla
when young, broadly conical with rugulose
disc in age, sulcate, pruinose, dull, light brown
(7D5-6) to pale greyish brown (7D3-4) when
young, grey to greyish orange (6B-C3) in age.
Context light brownish cream, thin. Lamellae
adnexed, subdistant (12-16), broad (1-3 mm),
light brownish cream, non-marginate, nonintervenose. Stipe 20-50 × 0.5-0.8 mm, central,
cylindrical, wiry, glabrous, non-insititious,
apex yellowish white, base dark brown. Odor
and taste not distinctive.
299
µm diam., interwoven, cylindrical to inflated,
smooth, hyaline, dextrinoid, thin-walled, nongelatinous. Stipitipellis hyphae 3-5 µm diam.,
parallel, cylindrical, brown, smooth, dextrinoid,
thick-walled (up to 1µm), non-gelatinous. Stipe
trama hyphae 3-5 µm diam., parallel,
cylindrical, hyaline, smooth, dextrinoid, thinwalled, non-gelatinous. Caulocystidia absent.
Clamp connections present in all tissues.
Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves or on wood, Northern Thailand.
Material examined: Thailand. Phrae Province,
Muang District, Pa Dang, Nong Kam Village, near Nong
Kam reservoir, 18 August 2005, N. Wannathes 424
(CMU, SFSU)
Discussion: The Thai specimen is similar
to M. pallescens Murr. as reported by Pegler
(1983) from the Lesser Antilles in all
taxonomically significant details except for
Pegler's report of slightly smaller basidiospores
(12-17.5 × 3-5 µm, x = 14.5 × 3.8 µm).
Because M. pallescens is currently known only
from the neotropics, until further material is
collected in Thailand, sequenced, and
compared with neotropical specimens, our
identification must remain tentative.
Fig. 72. Marasmius aff. pallescens (N. Wannathes 424). 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pleurocystidia 5. Pileipellis, Scale bar 1 = 20 mm, 2-5 =
10 µm
57. Marasmius bondoi Wannathes, Desjardin
& Lumyong, sp. nov.
(Figs 73, 75-3)
MycoBank: MB512430
Basidiospores (16-)17-18(-19) × 4-5 µm
[x = 17.5 ± 0.7 × 4.1 ± 0.3 µm, Q = 3.6-4.8, Qm
= 4.3, n = 25 spores per 1 specimen], narrowly
ellipsoid to fusoid, smooth, hyaline, inamyloid,
thin-walled. Basidia not observed. Basidioles
cylindrical to clavate. Cheilocystidia common, of
Siccus-type broom cells; main body 6-18 × 510 µm, cylindrical to clavate, hyaline,
inamyloid, thin- to thick-walled; apical setulae
3-6 × 1 µm, cylindrical, conical to subacute,
yellow to pale yellow, thick-walled. Pleurocystidia common, 46-53 × 10-15 µm,
cylindrical to fusoid, constricted at the apex,
hyaline, inamyloid, thin-walled. Pileipellis
hymeniform, mottled, composed of Siccus-type
broom cells; main body 9-17 × 6-8 µm,
cylindrical to clavate, hyaline, inamyloid, thinwalled; apical setulae 3-5 × 1 µm, crowded,
cylindrical, subacute, brownish yellow to light
brown, thick-walled. Pileus trama interwoven,
dextrinoid. Lamellar trama hyphae 3-8(-10)
Etymology: ‘bondoi’ (Thai) = on the mountain;
referring to the habitat where the holotype specimen was
found.
Pileus 12-32 mm diametro, convexus usque
plano-convexus, umbonatus vel subumbonatus, striatus
usque sulcatus, pruinosus, hebetatus, disco atrobrunneo
usque brunneo, margine flavo-brunneo, hygrophanus,
ubique pallide brunneo-flavo post arescendum.
Contextus flavo-brunneus, tenuis. Lamellae annexae
usque liberae, distantes usque subdistantes (10-20) cum
2-4 seriebus lamellularum, latae (2-4 mm), cinereoaurantiacae, haud marginatae, haud intervenosae. Stipes
32-92 × 1-2 mm, centralis, cylindratus, filo metallico
similis, cavus, glaber, haud insititius, apice cinereo-flavo,
basi rubro-brunnea usque violaceo-brunnea. Odor suco
cocois similis. Sapor non proprius. Basidiosporae 1318(-20) × 3-5 µm, anguste ellipsoideae usque fusoideae,
oblique curvae, laeves, hyalinae, inamyloideae,
tenuitunicatae Basidia 25-29 × 7-9 µm, cylindrata usque
clavata, 4-spora. Basidiolae fusoideae usque clavatae.
Cheilocystidia abundantia, typi Sicci; (7-)12-21 × (3-)510 µm, cylindrata usque clavata, hyalina, inamyloidea,
tenuitunicata; setulis apicalibus 3-15(-21) × 1-2 µm,
cylindratis usque conicis, interdum lobatis, subacutis,
flavis usque pallide flavis, tenuitunicatis. Pleurocystidia
vulgaria, 40-58 × 7-15 µm, cylindrata usque fusoidea,
300
Fungal Diversity
constricta et interdum lobata ad apicem, hyalina,
inamyloidea, tenuitunicata. Pileipellis hymeniformis,
maculosus, typi Sicci; (8-)13-20(-25) × 6-10 µm,
cylindratus usque clavatus vel pyriformis, hyalinus
usque pallide flavus, inamyloideus, tenui- usque
crassetunicatus; setulis apicalibus 4-10(-15) × 1-1.5 µm,
coartatis, cylindratis, subacutis, flavo-brunneis usque
brunneis, crassetunicatis. Trama pilei intertexta, hyphis
3-8 µm diametro, cylindratis usque inflatis, laevibus,
hyalinis, fortiter dextrinoideis, tenuitunicatis. Trama
lamellae intertexta usque regularis, hyphis 3-8(-13) µm
diametro, cylindratis usque inflatis, laevibus, hyalinis,
dextrinoideis,
tenuitunicatis,
haud
gelatinosis.
Stipitipellis subparallelus, hyphis 3-6(-9) µm diametro,
cylindratis, flavo- brunneis usque brunneis, laevibus,
leniter dextrinoideis, crassetunicatis (usque ad 2 µm),
haud gelatinosis. Trama stipitis parallela, hyphis 4-13(19) µm diametro, cylindratis, hyalinis, laevibus,
dextrinoideis,
tenuitunicatis,
haud
gelatinosis.
Caulocystidia nulla. Fibulae praesentes in omnibus
texturis.
Typus: Chiang Mai Province, Doi Suthep-Pui
National Park, trial opposite with the way to Huai Kok
Ma Village, gregarius usque confertim gregarius, typice
caespitosus in ligno vel folis plantae dicotyledoneae, 2
July 2005, N. Wannathes 386 (CMU: holotypus; SFSU:
isotypus).
Pileus 12-32 mm diam., convex to planoconvex, umbonate or subumbonate, striate to
sulcate, pruinose, dull, disc dark brown to
brown, margin yellowish brown, hygrophanous, fading to pale brownish yellow overall
with moisture loss. Context yellowish brown
(5D5), thin. Lamellae adnexed to free, distant
to subdistant (10-20) with 2-4 series of
lamellulae, broad (2-4 mm), greyish orange,
non-marginate, non-intervenose. Stipe 32-92 ×
1-2 mm, central, cylindrical, wiry, hollow,
glabrous, non-insititious, apex greyish yellow
(4B4), base reddish brown (8E8) to violet
brown (10F8). Odor of coconut juice. Taste not
distinctive.
Basidiospores 13-18(-20) × 3-5 µm [xmr
= 14.2-18.3 × 3.5-4.6 µm, xmm = 16.4 ± 1.3 ×
3.9 ± 0.3 µm, Qmr = 3.5-4.9, Qmm = 4.2 ± 0.5, n =
25 spores, s = 10 specimens], narrowly ellipsoid
to fusoid, curved in profile, smooth, hyaline,
inamyloid, thin-walled. Basidia 25-29 × 7-9 µm,
cylindrical to clavate, 4-spored. Basidioles fusoid
to clavate. Cheilocystidia abundant, of Siccustype broom cells; main body (7-)12-21 × (3-)510 µm, cylindrical to clavate, hyaline,
inamyloid, thin-walled; apical setulae 3-15(-21)
× 1-2 µm, cylindrical to conical, sometimes
lobed, subacute, yellow to pale yellow, thinwalled. Pleurocystidia common, 40-58 × 7-15
Fig. 73. Marasmius bondoi (N. Wannathes 237). 1.
Basidiomes 2. Basidia 3. Basidiospores 4. Cheilocystidia 5.
Pleurocystidia 6. Pileipellis, Scale bar 1 = 20 mm, 2-6 =
10 µm
µm, cylindrical to fusoid, constricted and
sometimes lobed at the apex, hyaline,
inamyloid, thin-walled. Pileipellis hymeniform,
mottled, composed of Siccus-type broom cells;
main body (8-) 13-20(-25) × 6-10 µm,
cylindrical to clavate or pyriform, hyaline to
pale yellow, inamyloid, thin- to thick-walled;
apical setulae 4-10(-15) × 1-1.5 µm, crowded,
cylindrical, subacute, yellowish brown to
brown, thick-walled. Pileus trama hyphae 3-8
µm diam., interwoven, cylindrical to inflated,
smooth, hyaline, strongly dextrinoid, thinwalled. Lamellar trama hyphae 3-8(-13) µm
diam., interwoven to regular, cylindrical to
inflated, smooth, hyaline, dextrinoid, thinwalled, non-gelatinous. Stipitipellis hyphae 36(-9) µm diam., subparallel, cylindrical,
yellowish brown to brown, smooth, weakly
dextrinoid, thick-walled (up to 2 µm), nongelatinous. Stipe trama hyphae 4-13(-19) µm
diam., parallel, cylindrical, hyaline, smooth,
dextrinoid,
thin-walled,
non-gelatinous.
Caulocystidia absent. Clamp connections
present in all tissues.
301
M. bondoi NW237
M. bondoi NW390
98
M. bondoi NW386
63
M. bondoi NW011
55
M. bondoi NW320
74
M. bondoi NW384
M. bondoi NW399
M. graminipes NW078
M. plicatulus NW439
5 changes
Fig. 74. Branch and bound bootstrap tree of the Marasmius bondoi species complex. Numbers above branches represent
support from 2000 bootstrap replicates.
Habit, habitat and known distribution.
Gregarious to densely gregarious, typically in
cespitose clusters on wood or on
dicotyledonous leaves, Northern Thailand.
Material examined: Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Sangasabhasri
Land on the way to Huai Kok Ma Village, N 18º 48.4’ E
98º 54.6’ alt. 1,146 m., 30 June 2003, N. Wannathes 011
(CMU, SFSU); same location, 6 June 2004, Y.S. Tan
318 (CMU, SFSU); same location, 24 July 2004, N.
Wannathes 237 (CMU, SFSU); same location, 24 June
2005,N. Wannathes 320 (CMU, SFSU); Chiang Mai
Province, Doi Suthep-Pui National Park, trial opposite
with the way to Huai Kok Ma Village, 2 July 2005, N.
Wannathes 384 (CMU, SFSU); same location, 2 July
2005, N. Wannathes 386 (CMU: holotype; SFSU:
isotype), N. Wannathes 388, 389 and 390(CMU, SFSU);
Chiang Mai Province, Doi Suthep-Pui National Park,
Medicinal Plant Garden, 3 August 2005, N. Wannathes
399 (CMU, SFSU).
Discussion: Marasmius bondoi is
characterized by a convex, subumbonate,
sulcate pileus with brown and yellowish brown
tones, distant to subdistant (10-20) lamellae
with or without brown edges, basidiospores
with means in the range 14.2-18.3 × 3.5-4.6 µm,
Siccus-type broom cells with setulae 2-15 × 1-2
µm, fusoid-constricted pleurocystidia, and
growth on wood or dicot leaf substrates. In a
separate branch and bound analysis involving
only M. bondoi and M. graminipes with M.
plicatulus as outgroup (Fig. 74), several clades
302
were recognized that differed in lamellar
spacing, albeit with low BS support, viz.,
specimens with distant (10-15) lamellae
[NW011, 237, 386 and 390] and specimens
with subdistant (13-20) lamellae [NW320, 384
and 399]. All other taxonomically significant
micro- and macromorphological features of all
M. bondoi specimens are overlapping and we
recognize them as representing a single species
with variable lamellar spacing.
Marasmius bondoi is most phenetically
similar to two African species, M. grandisetulosus
Singer and M. tenuisetulosus (Singer) Singer.
Both of the latter species are reported to have
distinctly striped pilei with darker sulcae
(brown or rusty-tawny) and paler ridges
(yellowish grey to orange or tawny). The Thai
M. bondoi has an evenly pigmented pileus that
is never striped.
In the ML tree, clade AB contained seven
sequences of M. bondoi with M. graminipes on
a relatively long branch embedded within the
clade as sister to one sample (NW 390) but
with low support (0.52 PP, 53% BS). In a
branch and bound analysis with 2000 replicates
and M. plicatulus as outgroup (Fig. 74), the
seven M. bondoi sequences cluster with low BS
support (55%) and the morphologically distinct
M. graminipes is sister to the M. bondoi clade.
Fungal Diversity
Fig. 75. Basidiomata of Marasmius section Sicci ser. Haematocephali 1. M. hypopheus (N. Wannathes 285) 2. M. aff.
pallescens (N. Wannathes 424) 3. M. bondoi (N. Wannathes 237), Scale bars = 20 mm.
Discussion
In most cases, the ITS sequence data
supported recognition of morphospecies with
high statistical support. For example, three
specimens of the new species M. jasminodorus
(in clade AA, Fig. 1) that were collected in
diverse geographical sites (different mountains at
different elevations and forest types in different
provinces) formed a clade with 0.98 PP and
100% BS support and showed almost no
genetic variability.
Singer (1976) recognized Marasmius as
representing 11 sections. Based on nLSU rDNA
sequences data, Wilson and Desjardin (2005)
restricted the genus Marasmius to only seven
sections (viz. Globulares, Hygrometrici, Leveilleani, Marasmius, Neosessiles, Scotophysini,
Sicci) and recognized the other four Singerian
sections (viz., Alliacei, Androsacei, Epiphylli,
Fusicystides) as belonging to other genera.
Only five of the sections in Marasmius sensu
stricto were encountered in northern Thailand
(Hygrometrici and Scotophysini excluded) and
of these, only sect. Marasmius subsect.
Marasmius was monophyletic based on the ITS
dataset (Fig. 1; clade D). In addition, Singer
(1976, 1986) and Antonín and Noordeloos
(1993) subdivided sect. Sicci into four series
(viz., Atrorubentes, Haematocephali, Leonini,
Spinulosi) based on the presence or absence of
pleurocystidia, setae or simple cylindrical
caulocystidia. None of these series are
monophyletic in our analyses. There was very
limited statistical support for the deeper nodes
in the ML or MP analyses (Fig. 1) suggesting
that the ITS region may be of limited value in
clarifying the infrageneric delimitation of
Marasmius. Before any formal reorganization
of taxa within Marasmius is proposed, further
phylogenetic analyses based on a global sample
set and additional genes (LSU, rpb2, ef1-α)
must be conducted.
Although the ITS data alone are of
limited use in elucidating infrageneric
phylogenetic structure, they are very useful in
aiding taxonomic delimitations at the species
rank. In most cases, species delimitations based
on morphological characters are concordant
with those suggested by ITS sequence
similarity. And this pattern holds true for
geographically distant and ecologically distinct
populations [cf., M. jasminodorus (.98 PP,
100% BS), M. brunneoolivascens (.99 PP, 92%
303
BS), M. bambusiniformis (1.0 PP, 100% BS),
M. laticlavatus (1.0 PP, 100% BS), M. cafeyen
(1.0 PP, 100% BS)].
Historically, morphological characters
have been used in combination to delimit
sections (collariate lamellae, insititious stipe,
pileipellis anatomy), series (pleurocystidia,
setae or simple caulocystidia), and species
(basidiospore size, broom cell setulae, cystidia
shape). The ITS data support the use of these
morphological characters to aid in delimiting
taxa at various taxonomic ranks, albeit not
necessarily as circumscribed by Singer (1976,
1986), Antonín and Noordeloos (1993),
Desjardin and Horak (1997), and Desjardin et
al. (2000). For example, all taxa that lack
collariate lamellae belong in clades A and F
(Fig. 1) and are phylogenetically distinct from
taxa with collariate lamellae (all other clades).
However, all collariate species (sect.
Marasmius) do not form a monophyletic group.
In addition, all species with non-insititious
stipes belong to the well-supported clade A
(sects. Sicci and Globulares), whereas those
with insititious stipes belong in the other clades.
Moreover, all tested species with a unifactorial
(bipolar) mating system belong in clade A,
whereas those with a bifactorial (tetrapolar)
mating system belong to the clades with
collariate lamellae and insititious stipes. Based
on these data, clade A, representing all
included species of sects. Sicci and Globulares
can be distinguished from all other Marasmius
by the presence of a non-insititious stipe,
bipolar mating system and absence of collariate
lamellae. A pileipellis formed from Siccus-type
broom cells versus Globulares-type cells (nonsetulose) that currently differentiates sect. Sicci
from sect. Globulares is not a phylogenetically
significant feature according to the ITS data.
Of the eight species with Globulares-type
pileipellis elements included in the analyses,
five belong to clade AD, one in clade AA, and
two are in clade AF. Desjardin and Horak
(1997) suggested that the separation of species
into sections Sicci and Globulares based on
pileipellis anatomy was untenable when they
described M. nexus Desjardin & Horak, a
species from Papua New Guinea with a
pileipellis formed from equal numbers of
Siccus-type cells and Globulares-type cells.
304
The presence or absence of pleurocystidia, setae or simple caulocystidia have
been used to distinguish series with sect. Sicci,
although these infrageneric groups are not
monophyletic based on the ITS data
Likewise, basidiospore size is a
taxonomically significant feature to aid in
delimiting species, but is of limited use at
higher taxonomic levels. For example, M.
pellucidus, with xmm = 7.2 × 3.3 µm, belongs to
a subclade within clade AA (with .77 PP, 78%
BS) and is distinct from the other members (M.
araucariae, M. jasminodorus, M. inthanonensis)
who have basidiospore means in the range
10.1-10.4 × 3.6-3.9 µm. Alternatively, shared
basidiospore size may be a clade-defining
feature. Clade AC, with two setoid (SS) species
(M. nummularius, M. trichotus) and two nonsetoid species with simple caulocystidia (SA;
M. iras, M. xestocephalus) all share narrowly
fusoid basidiospores with means in the range
12.1-13.2 × 3.7-4.1 µm.
Combinations of these micro-characters
and ITS data can be used to define clades. For
example, four species within clade AD with 1.0
PP and 100% BS are circumscribed by
Globulares-type pileipellis cells, long-clavate
basidiospores with means in the range 22.830.0 × 4.4-5.6 µm, and an absence of
pleurocystidia and caulocystidia. The ITS data
are a tremendous help in delimiting the
pleurocystidiate species M. haematocephalus
(clade AE with 1.0 PP, 99% BS) from other
taxa, and in distinguishing its various forms
whose pilei range in color from yellowish
orange to brown, olive, greyish blue, red and
violet. Micromorphologically all isolates of this
clade are indistinguishable; however, the
branch and bound analysis (Fig. 70) clearly
distinguishes five of the six forms into wellsupported lineages.
Conclusions
The ITS sequence data generally support
the recognition of species circumscribed by
macro- and micromorphological characters, and
in many cases can be used to delimit clades of
taxa with shared morphological features.
However, ITS sequences are of limited use in
recognizing currently delimited infrageneric
Fungal Diversity
taxa at the series rank and above. Additional
species and genes need to be sequenced and
analyzed with multiple algorithms in order to
clarify the infrageneric delimitation of the
genus Marasmius
Acknowledgments
This research was funded in part by National
Science Foundation (USA) PEET grant (DED-0118776)
to D.E.D. Pibulsongkram Rajabhat University and
Commission on Higher Educations provided partial
financial support to N.W. for her doctoral research. We
thank Ruben Walleyn for providing the colour photo of
M. mokfaensis. Thanks are extended to Huyen Thanh Le
and Yee Shin Tan for help with specimen collection and
Rajesh Jeewon for help with molecular sequencing. The
first author thanks Sonchai Wannathes for help in
writing the Latin diagnoses.
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