A Monograph of Marasmius (Basidiomycota) From Northern Thailand

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A monograph of Marasmius (Basidiomycota) from Northern Thailand based

on morphological and molecular (ITS sequences) data
Article in Fungal diversity · July 2009
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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 present here a monograph of Marasmius from

the region based on morphological and
This paper is part of a series dealing with molecular (ITS sequences) data.
the macrofungi of northern Thailand (Le et al., In this treatment we accept the generic
2007a,b; Wannathes et al., 2009). The genus circumscription of Marasmius as delimited by
Marasmius Fr. (Agaricales, Marasmiaceae) is Wilson and Desjardin (2005). They restricted
worldwide in distribution and is composed of Marasmius to a monophyletic lineage based on
approximately 600 species. Recently nLSU rDNA sequences data including only
Marasmius of Southeast Asia has received sections Globulares, Hygrometrici, Leveilleani,
renewed interest through the research of Marasmius, Neosessiles, Scotyphysini and Sicci
Desjardin et al. (2000) in Indonesia, Tan et al. as defined by Singer (1976, 1986). Other
(2007) in Malaysia, and Desjardin et al. (2004) sections recognized by Singer (1976, 1986) are
and Wannathes et al. (2004, 2007) in Thailand. currently excluded from Marasmius. Wilson
To date, no monographic treatment of and Desjardin (2005) transferred members of
Marasmius has been published from Thailand. Marasmius section Alliacei to the genus
During the years 2003 to 2007, we conducted Mycetinis. Section Androsacei was elevated to
extensive fieldwork in three provinces (Chiang generic rank as Setulipes by Antonín (1987);
Mai, Chiang Rai and Phrae) of northern section Epiphylli belongs in the family
Thailand. As a result of these expeditions, we Physalacriaceae (Wilson and Desjardin 2005);
209
and section Fusicystides is treated as a groups, and 3 species of Crinipellis for
synonym of Setulipes (Desjardin unpubl.). outgroup rooting purposes.
Materials and methods DNA extraction, Polymerase Chain Reaction

and sequencing:
Morphological protocols Genomic DNA was extracted from two
Color terms and notations in parentheses sources: mostly from dried herbarium specimens,
are those of Kornerup and Wanscher (1978). with fifteen extractions from living cultures
All measurements and colors reported for (Table 1). Extraction of dried herbarium tissue
microscopic features were observed from dried (1-10 mg) was done using the E.Z.N.A. fungal
material rehydrated in 100% ethanol followed DNA Miniprep Kit or E.Z.N.A. Forensic DNA
by distilled water, 3% potassium hydroxide Extraction Kit (Omega Bio-tek Inc., Norcross,
(KOH) or Melzer’s reagent. The terms used to GA, USA). Pure culture extractions were
describe lamellae spacing refer to the number performed with a modified SDS extraction
of lamellae that reach from the stipe to the protocol of Roger and Bendich (1994) and
pileus margin and do not include the lamellulae, Johnson (1998).
whose spacing is indicated by the number of ITS regions 1 and 2, and the 5.8S rDNA,
series present. Spore statistics include: xm, the were amplified using primers ITS1-F, ITS5,
arithmetic mean of the spore length by spore ITS 4, ITS4-B (Gardes and Bruns, 1993; White
width (± standard deviation) for n spores et al., 1990) and ITS4-m (TTG AGC TTT TCC
measured in a single specimen; xmr, the range CTC TTC AC) which was designed in this study.
of spore means, and xmm, the mean of spore When the amplification failed the intermediate
means (± SD) when more than one specimen is primers ITS2 and ITS3 (White et al., 1990)
available; Q, the quotient of spore length by were also used. The DNA fragment was
spore width in any one spore, indicated as a amplified on GeneAmp PCR System 9600
range of variation in n spores measured; Qm, (Applied Biosystems®, Perkin-Elmer
the mean of Q-values in a single specimen; Qmr, Corporation, Nortwalk, CT, USA) or PCT-
the range of Qm values and Qmm, the mean of 100™ (MJ research, Inc., Watertown, MA, USA)
Qm values where more than one specimen is automated thermal cycle. PCR products were
available; n, the number of spores measured purified using ExoSAPIT Kit (USB
per specimen; s, the number of specimens Corporation, Cleveland, OH, USA) or GFX™
involved. Specimens are deposited in the PCR DNA or Gel band Purification Kit
mycological herbarium at Chiang Mai (Amersham Biosciences, USA) and run on an
University (CMU), and the H. D. Thiers ABI 3100 Genetic Analyzer System (Applied
Herbarium at San Francisco State University Biosystems®, Foster City, CA, USA). Sequence
(SFSU). editing was performed with Sequencher 4.2
Phylogenetic protocols software (GeneCodes Co., Ann Arbor, MI,
USA). Edited sequences have been deposited
Taxon sampling: in GenBank (Table 1).
One hundred and twenty five Internal
Transcribed Spacer (ITS) nrDNA sequences Phylogenetic analyses:
representing 62 species were used in the Initial sequence alignment was
analyses, of which 116 sequences of performed with Clustal X (Thompson et al.,
Marasmius were generated as part of this study 1997) using the default settings, followed by
(Table 1), 4 sequences of Marasmius and 2 manual alignment with MacClade 4.04
sequences of Crinipellis were obtained from (Maddison and Maddison, 2001). Parsimony
Y.S. Tan (Univ. of Malaya), 2 sequences of analyses were conducted using PAUP* 4.0b10
Marasmius were obtained from GenBank, and 1 (Swofford, 2002). Searches employed a
sequence of Crinipellis were obtained from J. heuristic search method with all characters
Kerekes (San Francisco State Univ.). The weighted equally, gaps treated as missing data,
sequences represent 59 species of Marasmius random stepwise sequence addition, tree
in nine traditionally accepted infrageneric bisection reconnection (TBR), and collapse of
210
Fungal Diversity
Table 1. List of Marasmius specimens sequenced (ITS1-5.8S-ITS2) for this study, itemized by
infrageneric group.
GenBank
Species Sections/ Series Collection No.
accession No.
M. albimyceliosus Globulares NW422 EU935544
M. calvus Globulares NW331 EU935481
M. grandiviridis Globulares NW152 EU643514
M. grandiviridis Globulares NW349** EU643515
M. laticlavatus Globulares NW231 EU643510
M. mokfaensis Globulares DED7726 EU643516
M. mokfaensis Globulares NW020** EU643517
M. pellucidus Globulares NW321 EU935508
M. purpureostriatus Globulares NW158 EU935539
M. purpureostriatus Globulares NW318 EU935538
M. pseudopurpureostriatus Globulares NW286 EU643513
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
M. brevicollus Marasmius/ Sicciformes NW128 EU935558
M. cafeyen Marasmius/ Sicciformes NW058** EU935546
M. cafeyen Marasmius/ Sicciformes NW130*** EU935547
M. cafeyen Marasmius/ Sicciformes NW200* EU935548
M. crinis-equi Marasmius/ Sicciformes NW348 EU935555
M. aff. crinis-equi Marasmius/ Sicciformes NW182 EU935564
M. aff. crinis-equi Marasmius/ Sicciformes NW205 EU935565
M. guyanensis Marasmius/ Sicciformes NW254* EU935552
M. guyanensis Marasmius/ Sicciformes NW280* EU935553
M. guyanensis Marasmius/ Sicciformes TYS314* EU935554
M. nigrobrunneus Marasmius/ Sicciformes NW162 EU935570
M. nigrobrunneus Marasmius/ Sicciformes NW223 EU935572
M. nigrobrunneus Marasmius/ Sicciformes NW416* EU935571
M. nigrobrunneus “f. cinnamomeus” Marasmius/ Sicciformes NW025 EU935576
M. nigrobrunneus “f. cinnamomeus” Marasmius/ Sicciformes NW260* EU935574
M. nigrobrunneus “f. cinnamomeus” Marasmius/ Sicciformes NW327* EU935577
M. nigrobrunneus “f. cinnamomeus” Marasmius/ Sicciformes TYS281 EU935575
M. purpureobrunneolus Marasmius/ Sicciformes NW215* EU935556
M. purpureobrunneolus Marasmius/ Sicciformes NW370 EU935557
M. purspureisetosu Marasmius/ Sicciformes NW155 EU935563
M. ruforotula Marasmius/ Sicciformes NW257* EU935550
M. rufurotula Marasmius/ Sicciformes NW312 EU935551
M. straminiceps Marasmius/ Sicciformes NW256 EU935549
M. subruforotula Marasmius/ Sicciformes NW140 EU935579
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
211
Table 1 (Continued). List of Marasmius specimens sequenced (ITS1-5.8S-ITS2) for this study,
itemized by infrageneric group.
GenBank
accession No.
M. inthanonensis Sicci/ Atrorubentes NW414 EU935515
M. iras Sicci/ Atrorubentes NW276 EU935486
M. iras Sicci/ Atrorubentes NW375 EU935487
M. jasminodorus Sicci/ Atrorubentes NW067** EU935512
M. jasminodorus Sicci/ Atrorubentes NW294 EU935513
M. jasminodorus Sicci/ Atrorubentes NW353 EU935514
M. luteolus Sicci/ Atrorubentes NW138 EU935506
M. luteolus Sicci/ Atrorubentes NW304 EU935507
M. ochroleucus Sicci/ Atrorubentes NW299 EU935503
M. pseudopellucidus Sicci/ Atrorubentes NW186 EU935504
M. pseudopellucidus Sicci/ Atrorubentes NW305 EU935505
M. xestocephalus Sicci/ Atrorubentes JFK69 EU935488
M. xestocephalus Sicci/ Atrorubentes NW344 EU935489
M. bambusiniformis Sicci/ Leonini NW329 EU935521
M. corneri Sicci/ Leonini NW269 EU935482
M. corneri Sicci/ Leonini TYS274* EU935483
M. cremeus Sicci/ Leonini WN366 EU935494
M. cremeus Sicci/ Leonini TYS320 EU935495
M. cupreostipes Sicci/ Leonini NW150 EU935485
M. imitarius Sicci/ Leonini NW297 EU935496
M. makok Sicci/ Leonini NW201 EU935524
M. bondoi Sicci/ Haematocephali NW011 EU935472
M. brunneoolivascens Sicci/ Haematocephali NW112 EU935516
M. brunneoolivascens Sicci/ Haematocephali NW277** EU935519
M. brunneoolivascens Sicci/ Haematocephali NW373 EU935517
M. brunneoolivascens Sicci/ Haematocephali NW397*** EU935518
M. ganyao Sicci/ Haematocephali NW005 EU935499
M. graminipes Sicci/ Haematocephali NW078 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 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
212
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
zero length branches. Analyses were performed Bayesian phylogenetic analyses were
in two steps: first, 1000 random sequence carried out using Metropolis-coupled Markov
addition replicates were performed, saving no chain Monte Carlo (MCMCMC) methods with
more than 5 trees per replicate, with MaxTrees MrBayes v3.1.2 (Huelsenbeck & Ronquist,
set to auto increase. Second, with MaxTrees 2001; Ronquist and Huelsenbeck, 2003), under
reset to 15 K, the most parsimonious trees a GTR+I+G model as determined above. The
resulting from the first step were used as analyses consisted of two parallel searches, run
starting trees and the analysis was allowed to for 5,000,000 generations, with six chains and
swap to completion. random starting trees. Default settings in
The appropriate model of sequence MrBayes were used for the incremental heating
evolution for phylogenetic analyses was scheme of the chains (3 heated and 1 cold
determined using the program MrModeltest chain), unconstrained branch length
v2.3 (Nylander, 2004). Maximum likelihood (unconstrained: exponential (10.0)), and
(ML) analyses were conducted in PAUP*, and uninformative topology (uniform) priors. The
employed the GTR+I+G model of sequence chains were sampled every 1000 generations.
evolution. ML searches followed an iterative Trees sampled prior to searches reaching a split
search strategy. First, a neighbor joining (NJ) deviation frequency of 0.03 were discarded as
tree was constructed using a Jukes-Cantor (JC) the burn-in, and the resulting trees were used to
distance model. Next, with the search criterion calculate Bayesian posterior probabilities.
reset to ML, the model parameters were To further analyze the relationships
estimated from the NJ tree, fixed, and an ML among three Marasmius species complexes,
search was conducted. After this initial search was sub-alignments were constructed by pruning
completed, the model parameters were re- taxa from the larger ITS dataset to represent the
estimated from the resulting ML tree(s), fixed, species complexes and sister taxa. Boostrap
and another ML search was intiated with analyses were performned in PAUP* on
starting trees obtained via NJ. This step was resulting sub-alignments, consisting of 2000
completed a third and final time, for a total of replicates under a branch and bound search
three independent ML searches. Upon algorithm with furthest sequence addition and
completion of the final search, trees resulting MulTrees on.
from all three iterations were compared to
insure that the searches had converged on Results
equally likely topologies. Maximum likelihood
bootstrapping was performed in GARLI v0.951 Molecular Phylogeny
(Zwickl, 2006) with 300 replicates under the Phylogenetic analyses were performed on
GTR+I+G model of sequence evolution, with an ITS dataset of 125 sequences of which 112
all parameter values estimated by the program sequences were Marasmius species and 3
and using the default run termination settings. sequences were Crinipellis species, with the
213
latter used as an outgroup for rooting purposes. Section Marasmius subsection Sicciformes
Two new Thai Marasmius species were not (collariate species with Siccus-type broom cells)
included in the analyses because of the is not monophyletic, with species clustering
presence of multiple different ITS copies in into more than three separate clades: clade B
each species. Cloning was not conducted to with 0.91 PP and 60% BS; clade C with 0.92
address this problem. Sequencing products PP and 85 % BS; clade E with no statistical
ranged from 270 nucleotides (Marasmius support; plus four unsupported taxa at the base
mokfaensis NW020; partial sequence) to 883 of the tree. A single member of section
nucleotides (Marasmius tantulus NW239). All Neosessiles (M. tenuissimus) and the monotypic
sequences were aligned and the ends trimmed section Leveilleani (M. leveilleanus) form a
to create a dataset of 701 nucleotides that clade (F) with relatively strong support (0.97
included 354 parsimony informative characters. PP and 82% BS) although its placement in the
Each of the ML analysis iterations tree is not supported statistically. Within sect.
recovered a single tree, the likelihood values of Sicci, the four infrasectional taxa circumscribed
which did not differ significantly. We have by Singer (1976) based on micromorphological
selected the topology resulting from the third features (viz., series Atrorubentes,
iteration to present here (Fig. 1; -lnL = Haematocephali, Leonini, Spinulosi) are not
10983.475). Parsimony analyses produced 15 monophyletic (Fig. 1). Members of ser.
K equally parsimonious trees 819 steps, C.I. = Atrorubentes fall into two clades (AA, AC);
0.427, R.I. = 0.701),), which did not differ members of ser. Haematocephali belong to
significantly in topology from those recovered in three clades (AA, AB, AE) and two isolated
the ML analyses. Bayesian analyses reached a lineages; members of ser. Leonini are scattered
standard deviation of split frequencies of 0.03 throughout Clade A; while members of ser.
after 1,592,000 generations, and the intial 4000 Spinulosi fall into two clades (AC, AF) plus an
trees recovered were excluded as the burn-in. isolated lineage sister to AA (M. coarctatus).
Maximum likelihood bootstrap values (BS) and Clade AF, with 1.0 PP support, contains
Bayesian posterior probalilites (PP) support members of sect. Globulares, sect. Sicci series
many of the terminal nodes in the phylogeny, Leonini and Spinulosi.
but fail to recover the deeper nodes with strong In the ML analysis, three species
support. complexes were uncovered, based on M.
All members of sections Globulares plus bondoi (sect. Sicci ser. Haematocephali; clade
Sicci belong to a monophyletic clade with AB) with 1.0 PP and 74% BS support, M.
relatively strong support (1.0 PP and 76% BS; haematocephalus (sect. Sicci ser.
clade A), although the sections themselves are Haematocephali; clade AE) with 1.0 PP and
not monophyletic. Eight species currently 99% BS support, and M. nigrobrunneus (sect.
placed in sect. Globulares fall into three Marasmius subsect. Sicciformes) in an
distinct subclades (AA, AD, AF), although five unresolved position near the base of the tree
of them belong to a single subclade (AD) but (Fig. 1). To help clarify the taxonomic
with low support (.67 PP, 52% BS). Members boundaries of these species complexes, three
of section Marasmius, comprised of species separate bootstrap branch and bound analyses
with lamellae attached to a collarium, do not were performed based on datasets that included
form a monophyletic group (clades B-F and the ingroup taxa with their sister taxa (as
other species at the base of the tree). Section resolved in the ML tree) used as outgroup.
Marasmius subsection Marasmius (collariate These results will be discussed in the
species with Rotalis-type broom cells) is commentaries that follow the species
monophyletic for the four included species descriptions.
with 1.0 PP and 83% BS support (clade D).
214
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
0.77/ 78 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
0.66/ 54 M. auratus NW076 - AA
1.0/ 97 M. ochroleucus NW299 -SA
1.0/ 87
M. pseudopellucidus NW305 (type) -SA
0.98/
M. pseudopellucidus NW186 -SA
0.99/ 92 M. luteolus NW138 -SA
63
1.0/ M. luteolus NW304 -SA
0.95/ 74 M. brunneoolivasscens NW112 (type) -SH
M. brunneoolivascens NW373 -SH
M. suthepensis TYS280 (type) -SH
1.0/ 100 M. coarctatus NW315 -SS
0.93/ 85 M. coarctatus NW385 (type) -SS
1.0/ 100
M. bambusiniformis NW329 -SL
0.64/ 60 M. makok NW201 (type) -SL
0.52 / 53 M. bondoi NW390 -SH
M. graminipes NW078 (type) -SH
M. bondoi NW011 -SH
1.0/ 74
M. bondoi NW386 (type) -SH
M. bondoi NW237 -SH
AB
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 -
A
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
0.77
M. nummularius NW266 -
M. nummularius NW396 -
1.0/ 100 1.0/ 97 M. xestocephalus JFK69 -SA AC
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
AD
0.67/ 52 M. laticlavatus NW412 -G
M. laticlavatus NW293 -G
1.0/ 100 1.0/ 98 M. mokfaensis DED7726 (type) -G
0.9 M. mokfaensis NW020 -G
0.76 M. haematocephalus f. haematocephalus NW296 -SH
M. haematocephalus f. haematocephalus NW428 -SH
1.0/ 100
1.0/ 93 M. haematocephalus f. haematocephalus TYS277 -SH
0.79/ 63 M. haematocephalus f. atrobrunneus NW117 -SH
0.86/ 65 M. haematocephalus f. violaceus NW193 -SH AE
1.0/
M. haematocephalus f. violaceus NW413 -SH
1.0/ 76 1.0/
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 AF
0.97/ 55 M. araneocephalus NW358 (type) -SS
M. cohaerans var. lachnophyllus DED 4071 -SS
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 B
0.91/ 60 M.rufurotula NW312 -MS
1.0/ 93 M. guyanensis NW254 -MS
0.99/ 94 M. guyanensis TYS314 -MS
1.0/ 93 M. guyanensis NW280 -MS
M. crinisequi NW348 -MS
1.0 1.0/ 100 M. purpureobrunneolus NW215 -MS
0.92/ 85 M. purpureobrunneolus NW370 -MS C
M. brevicollus NW128 -MS
0.91/ 64 M. rotula DQ182506 -MM
0.52 M. apatelius NW427 -MM
1.0/ 83
1.0 M. apatelius NW437 -MM D
M. tantulus NW239 (type) -MM
0.99 M. somalomoensis NW232 -MM
100 M. aff. crinisequi NW182 -MS
M. aff. crinisequi NW205 -MS E
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 F
M. leveilleanus NW248 -L
1.0/ 100 M. leveilleanus NW268 -L
M. nigrobrunneus f. cinnamoneus NW119 -MS
M. nigrobrunneus f. cinnamoneus TYS281 (type) -MS
0.84/ 67 M. nigrobrunneus f. cinnamoneus NW327 -MS
M. nigrobrunneus NW162 -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
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 10. Stipe arising directly from rhizomorphs.............. 11

10. Stipe arising directly from the substrate;
rhizomorphs present or absent............................. 16
Key to species of Marasmius from northern
Thailand 11. Basidiomes and rhizomorphs associated with wood;
lamellae remote to distant (5-10) ............................. 12
1. Collarium present; stipe insititious.......................... 11. Basidiomes and rhizomorphs associated with
................................................ (sect. Marasmius) 2 leaves, typically bamboo and other grasses or rarely
1. Collarium absent; stipe insititious or non-insititious dicot leaves; lamellae subdistant (8-17) .............. 14
............................................................................. 21
12. Pileus very pale, cream-colored . 9. M. pallenticeps
2. Pileipellis composed of Rotalis-type broom cells 12. Pileus more deeply pigmented, reddish brown to
...........................................(subsect. Marasmius) 3 orangish brown, orange to greyish orange .......... 13
2. Pileipellis composed of Siccus-type broom cels.......
..........................................(subsect. Sicciformes) 6 13. Pileus reddish brown to orangish brown; lamellae
distant (8-10); basidiospores 10-12 × 4-4.5 µm, xmm
3. Basidiospores (17-) 20-22 × 4 µm; basidiomes very = 10.6 × 4.6 µm, Qm = 2............. 10. M. crinis-equi
small, with pilei up to 1.5 mm diam and stipes up to 13. Pileus orange to greyish orange; lamellae remote
1 mm long.......................................... 1. M. tantulus (5-8); basidiospores 8-10 × 4-6 µm, xmm = 9 × 4.9
3. Basidiospores ≤ 15 µm long; basidiomes larger, µm, Qm = 1.9 ....................... 11. M. aff. crinis-equi
with pilei usually > 1.5 mm diam and stipes > 5 mm
long......................................................................... 4 14. Pileus 2-6 mm diam, reddish brown; stipe 4-10 mm
long ....................................... 12. M. subruforotula
4. Lamellae distant (6-10), non-marginate; pileus 14. Pileus 2-28 mm diam, black, dark grey, greyish
yellowish white to brownish cream with a light brown, brown or brownish orange; stipe 8-215 mm
brown central spot .........................2. M. delicatulus long ..................................................................... 15
4. Lamellae subdistant to close (9-18), brown-
marginate; pileus brown with a hint of olive or 15. Pileus black, dark grey or greyish brown; stipe 50-
brownish cream to yellowish brown....................... 5 215 mm long ...........................................................
.............13A. M. nigrobrunneus f. nigrobrunneus
5. Pileus 2-4 mm diam, brown with a hint of olive, 15 Pileus brown to brownish orange; stipe 8-60 mm
with a reddish brown papilla in the center surround long ... 13B. M. nigrobrunneus “f. cinnamomeus”
by a pale zone; stipe 8-47 mm long ..........................
...............................................3. M. somalomoensis 16. Pileus reddish brown; lamellae subdistant (11-13);
5. Pileus 2-10 mm diam, light brownish cream to stipe 4-10 mm long ............... 12. M. subruforotula
yellowish brown without olive tones, with or 16. Pileus brown, brownish orange, brownish yellow,
without a small brown papilla, lacking a pale central rusty brown, cream-brown, light yellow; lamellae
zone; stipe 8-15 mm long .................4. M. apatelius distant (8-12); stipe 8-35(-60) mm long .............. 17
6. Pileus minutely hispidulous from the presence of 17. Pileus light yellow to yellowish white; rhizomorphs
pileosetae ................................................................ 7 absent ...................................... 14. M. straminiceps
6. Pileus glabrous, lacking pileosetae ......................... 8 17. Pileus more deeply pigmented, brown, brownish
orange, brownish yellow, rusty brown or cream-
7. Pileosetae uncommon, 36-82 µm long; pileipellis brown; rhizomorphs present or absent ................ 18
broom cells and cheilocystidia with numerous (5-20)
apical setulae; lamellae distant (7-8); pileus 0.2-2 18. Stipe arising directly from rhizomorphs or from
mm diam; stipe arising directly from rhizomorphs... bamboo or other grass leaves ..................................
................................................ 5. M. berambutanus .......... 13B. M. nigrobrunneus “f. cinnamomeus”
7. Pileosetae common, 45-350 µm long; pileipellis 18. Stipe arising directly from dicot leaves............... 19
broom cells and cheilocystidia with few (2-6)
apical setulae; lamellae subdistant (13-17); pileus 19. Basidiospores narrow, 10-12 × 3-4 µm, xmm = 11.1
1-4 mm diam; stipe not arising directly from × 3.5 µm, Qmm = 3.2.................. 15. M. guyanensis
rhizomorphs..........................6. M. purpureisetosus
19. Basidiospores broader, 9-12 × 4-7 µm, xmmr = 10-
10.2 × 4.3-5.8, Qmmr = 1.8-2.4............................. 20
8. Basidiospores > 13 µm long.................................. 9
8. Basidiospores ≤ 12 µm long................................ 10
20. Pileus disc brown with a cream-brown margin;
basidiospores 9-11 × 4-5 µm, xmm = 10 × 4.3 µm,
9. Basidiospores 16-23 × 3.5-5.5 µm, xmm = 19.7 ×
Qmm = 2. .......................................... 16. M. cafeyen
4.6 µm; lamellae distant (6-9) ......7. M. brevicollus
20. Pileus disc brownish yellow or rusty brown with a
9. Basidiospores 13-16 × 3-4 µm, xmm = 14.2 × 3.2
greyish orange to light yellow margin;
µm; lamellae distant to subdistant (9-13) ................
basidiospores 10-12 × 4.5-7 µm, xmm = 10.2 × 5.8
......................................8. M. purpureobrunneolus
µm, Qmm = 1 ................................17. M. ruforotula
217
21. Pileipellis composed of a hymeniform layer of 29. Stipe none or rudimentary, lateral to strongly
clavate to pyriform, non-setulose cells.................... eccentric, less than 2 mm long ................................
............................................ (sect. Globulares)... 22 ................(sect. Neosessiles)... 26. M. tenuissimus
21. Pileipellis composed of Siccus-type broom cells 29 29. Stipe central, well-developed .............................. 30
22. Pleurocystidia present, fusoid............ 18. M. calvus 30. Stipe insititious; lamellae free, well-separated from
22. Pleurocystidia absent........................................... 23 the stipe apex; coarse black rhizomorphs may be
present ..... (sect. Leveilleani)... 27. M. leveilleanus
23. Basidiospores < 10 µm long................................ 24 30. Stipe non-insititious; lamellae attached, typically
23. Basidiospores > 18 µm long................................ 25 adnexed to adnate; rhizomorphs absent ..................
........................................................ (sect. Sicci) 31
24. Cheilocystidia indistinct, basidiomorphous; pileus
context thin but opaque, subhygrophanous; 31. Setae present on pileus, lamellae and/or stipe.........
lamellae relatively broad (2-3 mm), strongly .................................................. (ser. Spinulosi) 32
intervenose; pileus with brown to creamy brown 31. Setae absent......................................................... 37
disc and striae; basidiomes usually solitary.............
..............................................19. M. albimyceliosus 32. Pleurocystidia and pleurosetae absent................. 33
24. Cheilocystidia distinct, irregularly cylindrical to 32. Pleurocystidia or pleurosetae present.................. 35
fusoid, ventricose or clavate; pileus context
extremely thin, pellucid, hygrophanous; lamellae 33. Basidiospores 6-7 × 2.5-3.5 µm; lamellae very
narrow (< 2 mm), often forked and intervenose; crowded, > 22 with 4-5 series of lamellulae; pileus
pileus with cream to orange white disc, pure white brown with a hint of olive ...........28. M. coarctatus
elsewhere; basidiomes usually in dense cespitose 33. Basidiospores 12-15 × 3-5 µm; lamellae close to
clusters ........................................20. M. pellucidus subdistant, 12-22 with 2-3 series of lamellulae;
pileus orange, brownish orange or reddish brown...
25. Pileus with bluish grey, purple or violaceous ............................................................................ 34
pigments .............................................................. 26
25. Pileus lacking bluish grey, purple or violaceous 34. Pileus orange to brownish orange; lamellae close
pigments .............................................................. 28 (16-22); pileosetae abundant, golden, up to 300 µm
or more long; caulosetae simple, acicular ...............
26. Basidiomes robust, with pilei 30-90 mm diam, ...................................................... 29. M. trichotus
bluish grey to purplish grey overall when young 34. Pileus reddish brown with brown margin; lamellae
becoming grayish brown with paler ridges in age; subdistant (12-18); pileosetae absent or rare,
basidiospores 27-33 × 5-6 µm (xmm = 30 × 5.3 µm) reddish brown, up to 115 µm long; caulosetae often
................................................... 21. M. mokfaensis with apical setulae................. 30. M. nummularius
26. Basidiomes smaller, with pilei 13-38 mm diam,
striped, with dark violet to grayish violet disc and 35. Basidiospores 10-13 × 5.5-7 µm .... 31. M. coklatus
plicae and yellow to grayish yellow ridges; 35. Basidiospores 6-8 × 2.5-4 µm ............................. 36
basidiospores 20-30 × 4-7 µm (xmr = 21.3-28.2 ×
4.6-6.2 µm).......................................................... 27 36. Pileus smooth to striatulate, not venose-reticulate,
brown with a hint of olive; lamellae very crowded
27. Pileus rather small, 13-20 mm diam, with dark (> 22) with 4-5 series of lamellulae; pileipellis
violet to violet disc and plicae, and narrow pale formed from Siccus-type broom cells, simple
grayish yellow to grayish white ridges; stipe up to pileosetae up to 60 µm long and transitional cells
1.5 mm thick; basidiospores 21-30 × 4-7 µm (xmm = with 2-4 apical setulae up to 37 µm long ................
23.8 × 5.3 µm).................. 22. M. purpureostriatus ....................................................28. M. coarctatus
27. Pileus larger, 14-38 mm diam, with violet to purple 36. Pileus venose-reticulate, dark purplish brown;
disc and narrow plicae, and broad yellow to cream lamellae subdistant (10-12) with 3 series of
ridges; stipe 1.5-3 mm thick; basidiospores 20-24.5 lamellulae; pileipellis formed from Siccus-type
× 5-6 µm (x = 22.7 × 5.6 µm) ................................. broom plus arachnoid cells with 2-4 setulae 30-70
...............................23. M. pseudopurpureostriatus µm long ..............................32. M. araneocephalus
28. Pileus 37-88 mm diam, yellowish green with darker 37. Stipe pruinose overall, pruinosity formed from
olive green plicae; stipe 3-7 mm thick; irregularly cylindrical, obtuse to subacute, non-
cheilocystidia clavate to irregular, sometimes lobed, setulose cells, Siccus type cells typically absent on
5-12 µm diam; basidiospores 26-30 × 4-5 µm (x = stipe surface (if present then they are in
26.7 × 4.4 µm).........................24. M. grandiviridis combination with non-setulose cells) ......................
28. Pileus 15-19 (-33) mm diam, yellowish grey to ............................................ (ser. Atrorubentes) 38
greyish cream with a brown disc; stipe 1-2 mm 37. Stipe glabrous or pruinose, if pruinose then
thick; cheilocystidia broadly clavate, 11-16 µm pruinosity formed only from Siccus-type broom
diam; basidiospores 26-35 × 5-7 µm (xmm = 30.4 × cells ..................................................................... 47
5.6 µm) ......................................25. M. laticlavatus
218
Fungal Diversity
38. Cheilocystidia simple, cylindrical to clavate, 46. Pileus disc smooth, dark olive to olive when young;
Siccus-type broom cells absent from lamellar edge stipe long, up to 100 mm long.................................
............................................................................. 39 ............................................. 41. M. inthanonensis
38. Cheilocystidia of Siccus-type broom cells .......... 40 46. Pileus disc rugulose, dark reddish brown when
young; stipe shorter, up to 45 mm long...................
39. Pileus disc golden yellow, margin orange to ..................... 38A. M. araucariae var. araucariae
yellowish orange; stipe base dark reddish brown;
basidiospore xmm = 12.2 × 4.1 µm, Qmm = 3.0; 47. Pleurocystidia absent....................(ser. Leonini) 48
basidiomes solitary on dicot leaves . 33. M. auratus 47. Pleurocystidia present .(ser. Haematocephali)... 54
39. Pileus disc pale yellowish white, margin white,
fading to white overall in age; stipe base pale 48. Pileus white to cream colored ........ 42. M. cremeus
orange to grayish orange; basidiospore xmm = 11.0 48. Pileus more deeply pigmented, olive, brown,
× 4.1 µm, Qmm = 2.7; basidiomes cespitose on reddish brown, brownish orange, greyish orange or
bamboo debris ................. 34. M. pseudopellucidus orange.................................................................. 49
40. Pileus light yellow to cream overall; lamellae close 49. Basidiospores 9-12 × 5-6 µm, Qm ≤ 2.0 ..................
to crowded (20-24).................. 35. M. ochroleucus ...............................................43. M. hypochroides
40. Pileus more deeply pigmented, olive brown, brown, 49. Basidiospores in the range 12-26 × 3-5 µm, Qm ≥
reddish brown, brownish orange or yellowish 4.0 ....................................................................... 50
brown; lamellae subdistant to close (12-20)........ 41
50. Pileus conical, with a dark greyish brown disc and
41. Caulocystidia of two types: Siccus-type broom cells olive margin ......................................44. M. makok
plus simple cylindrical cells ................................ 42 50. Pileus convex, brown, reddish brown, brownish
41. Caulocystidia of one type: simple or lobed, orange, greyish orange or orange, lacking olive
cylindrical cells, Siccus-type broom cells absent tones .................................................................... 51
from stipe surface ................................................ 44
51. Stipe distinctly copper colored overall, 70-250 mm
42. Pleurocystidia present as scattered Siccus-type long ......................................... 45. M. cupreostipes
broom cells; pileus bright brownish orange to 51. Stipe base brown, dark brown or reddish brown
golden orange with paler creamy orange margin; (never copper colored), 13-90 mm long.............. 52
lamellae distant (8-13), narrow (1-2 mm diam) ......
.........................................................36. M. luteolus 52. Pileus 10-43 mm diam; stipe 1-3 mm thick ............
42. Pleurocystidia absent; pileus dark brown to dark .........................................................46. M. corneri
reddish brown with light brown to brownish orange 52. Pileus 2-15 mm diam; stipe < 0.5 mm thick........ 53
margin; lamellae subdistant (12-20), broad (2-5
mm diam) ............................................................ 43 53. Pileus brown to reddish brown; lamellae distant
(10-12); basidiospore xmm = 18.6 × 4.4 µm;
43. Odor strongly of jasmine tea; lamellae pale basidiomes mostly on woody sticks ........................
yellowish white; pileus disc rugulose, dark reddish ......................................................47. M. imitarius
brown .................................... 37. M. jasminodorus 53. Pileus brownish orange to greyish orange; lamellae
43. Odor absent; lamellae brownish orange to grayish subdistant (10-18); basidiospore xmm = 16.2 × 3.7
brown or brown; pileus disc smooth, dark brown to µm; basidiomes on dicot leaves ..............................
brown or brownish orange....................................... .........................................48. M. bambusiniformis
............................38B. M. araucariae var. siccipes
54. Basidiospores 8-12 × 4-5(-6) µm, xmm = 9.6 × 4.8
44. Caulocystidia apically lobed or forked; pileus with µm, Qmm = 2.1 .............................. 49. M. confertus
distinct yellow tones when young (disc yellowish 54. Basidiospores 11-32 × 3.5-5.5 µm, xmmr = 12.6-
brown, margin grayish yellow to yellowish orange 28.2 × 3.7-4.5 µm, Qmmr = 3.0-6.6....................... 55
...............................................39. M. xestocephalus
44. Caulocystidia all simple, cylindrical cells; pileus 55. Basidiospores 25-32 × 4-4.5 µm, xmm = 28.2 × 4.3
lacking yellow tones when young, with dark brown,
µm, Qmm = 6.6; stipe 100-195 × 0.5 mm; lamellae
reddish brown or olive tones ............................... 45
distant (6-8); basidiomes on bamboo leaves ...........
.........................................................50. M. ganyao
45. Basidiospores 12-14 × 3-4 µm with xmm = 12.8 ×
55. Basidiospores 11-25 × 3.5-5.5 µm, xmmr = 12.6-22
3.7 µm, Qmm = 3.5; pileus dark brown overall when
× 3.7-4.5 µm, Qmmr = 3.0-5.9; stipe < 100 mm long;
young..................................................... 40. M. iras
lamellae distant to close (8-24); basidiomes
45. Basidiospores 8-12 × 3-4 µm with xmr = 10.4-10.5
typically on dicot leaves or wood, rarely on
× 3.9-4.0 µm, Qmr = 2.6-2.7; pileus olive or dark bamboo or other grass debris .............................. 56
reddish brown overall when young ..................... 46
219
56. Lamellae close (20-24); stipe surface with 1965. [Type: Marasmius pararotula Singer].
caulocystidia formed from grass-like clusters of = subsect. Pararotulae (Singer) Singer, Fl.
setulae 6-40 x 1-2 µm lacking a basal cell or arising Neotrop. Monogr. 17: 92. 1976.
from a rudimentary basal cell.... 51. M. graminipes
56. Lamellae distant to subdistant (8-18); stipe surface
glabrous, lacking caulocystidia ........................... 57 1. Marasmius tantulus Wannathes, Desjardin
& Lumyong, sp. nov. (Fig. 2)
57. Basidiospores 11-15 × 3.5-5.5 µm, xmmr = 12.6- MycoBank: MB512412.
13.2 × 4-4.5 µm, Qmmr = 3.0-3.7.......................... 58 Etymology: ‘tantulus’ = very small; referring to
57. Basidiospores 13-25 × 3.5-5 µm, xmmr = 16.1-22 × the very tiny basidiomes.
3.7-4.2 µm, Qmmr = 4.2-5.9.................................. 59 Pileus < 1.5 mm diametro, convexus, breviter
umbilicatus, sine papilla, striatus, glaber, hebetatus,
58. Pileus dark brown overall or with brown margin in cinereo-creameus. Contextus creameus, tenuis. Lamellae
age; lamellae olive brown to grayish olive or adnatae usque collarium infirmus evolutum, remotae (3-
yellowish olive; Siccus-type broom cells often 4), angustae, creameae, haud marginatae, haud
scattered on lamellar sides....................................... intervenosae. Stipes < 1 × < 0.3 mm, excentricus, sursum
.......................................52. M. brunneoolivascens angustatus cum basi subbulbosa, subvelutinus, insititius,
58. Pileus disc bright brown, margin brownish orange brunneus, sine rhizomorpha. Odor saporque non propria.
to brownish yellow; lamellae yellowish white; Basidiosporae (17-)20-22 × 4 µm, fusoideae usque
Siccus type broom cells absent on lamellar sides .... anguste clavatae, laeves, hyalinae, inamyloideae,
...................................................53. M. suthepensis tenuitunicatae. Basidia non observata. Basidiolae
fusoideae usque clavatae. Cheilocystidia vulgaria, typi
59. Pileus deep reddish brown, maroon, dark violet, Rotalis; 18-23 × 10-18 µm, subglobosa usque late
purple, purplish brown or yellowish orange with clavata, hyalina, inamyloidea, tenuitunicata; setulis
white margin; basidiospores 19-25 × 3.5-5 µm, xmm divergentibus 1-2 × 1 µm, gongylodibus usque conicis,
= 20.7 × 4.0 mm, Qmm = 5.2 forms of ..................... obtusis, confertis ad dimidium superiorem, hyalinis,
.........................................54. M. haematocephalus tenui- usque crassetunicatis. Pleurocystidia nulla.
59. Pileus dark brown to brown, ferruginous, brownish Pileipellis hymeniformis, haud maculosus, typi Rotalis;
orange, brownish yellow or grayish brown (lacking 27-35 × 16-22 µm, late clavatus, subglobosus, pyriformis
deep red, violet, purple or light yellow tones); vel adumbratim inaequabilis, hyalinus, inamyloideus,
basidiospores 13-19 × 3.5-5 µm, xmmr = 16.1-18.1 × tenuitunicatus; setulis divergentibus 1-2 × 1 µm,
4.0-4.2 µm, Qmmr = 4.2-4.3.................................. 60 gongylodibus usque conicis, obtusis, saepe confertis ad
dimidium superiorem, flavis, crassetunicatis. Trama pilei
60. Basidiomes on monocot leaves; pileus typically intertexta, inamyloidea. Trama lamellae intertexta usque
small, 1-5(-8) mm diam....................................... 61 regularis, hyphis 3-6 µm diametro, cylindratis, laevibus,
60. Basidiomes on dicot leaves and wood; pileus hyalinis, inamyloideis, tenuitunicatis, haud gelatinosis.
typically larger, 4-30(-48) mm diam ................... 62 Stipitipellis parallelus, hyphis 5-10 µm diametro,
cylindratis cum eminentiis digitiformibus dispersis, laete
61. Pileus dark brown overall; lamellae distant (8-11); flavis usque brunneo-flavis, laevibus, leniter
stipe 5-15 mm long.................................................. dextrinoideis usque dextrinoideis, tenuitunicatis, haud
........54B. M. haematocephalus “f. atrobrunneus” gelatinosis. Trama stipitis subparallela, hyphis 4-6 µm
61. Pileus disc brown to ferruginous, margin diametro, cylindratis, hyalinis, laevibus, inamyloideis,
ferruginous to grayish orange; lamellae subdistant tenuitunicatis, haud gelatinosis. Caulocystidia nulla.
(11-13); stipe 22-35 mm long.................................. Fibulae praesentes in omnibus texturis.
..................................................55. M. hypophaeus Typus: Chiang Mai Province, Doi Inthanon
National Park Junction of Highway 1,009 and road to
62. Pileus 4-15 mm diam, pallid, light brown to grayish Mae Chaem, N 18º 31.6’ E 98º 29.6’ alt. 1,703 m.,
brown with paler margin; cheilocystidia setulae 3-6 dispersus usque gregarius in ligno, 28 July 2004, N.
µm long ................................ 56. M. aff. pallescens Wannathes 239 (CMU: holotypus).
62. Pileus more robust, 8-30(-48) mm diam, more Pileus < 1.5 mm diam., convex,
deeply pigmented, brown to brownish orange, shallowly umbilicate, without papilla, striate,
yellowish brown with brownish cream margin;
cheilocystidia setulae 3-15(-21) µm long ...............
glabrous, dull, greyish cream. Context cream,
.......................................................... 57. M. bondoi thin. Lamellae adnate to a poorly developed
collarium, remote (3-4), narrow, cream, non-
Enumeration of taxa marginate, non-intervenose. Stipe < 1 × < 0.3
mm, eccentric, tapering upward with subbulbous
Section Marasmius, subsect. Marasmius at base, subvelutinous, insititious, brown;
Type: Marasmius rotula (Scop.: Fr.) Fr. rhizomorphs absent. Odor and taste not
= sect. Pararotulae Singer, Sydowia 18: 339.
distinctive.
220
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, thin-
walled, 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).
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
Fig. 2. Marasmius tantulus (N. Wannathes 239). 1. were previously known with basidiospores 20-
Basidiomes 2. Basidiospores 3. Basidioles 4. 22 µm long. The most similar species are M.
Cheilocystidia 5. Pileipellis 6. Stipitipellis, Scale bar 1 =
2 mm, 2-6 = 10 µm arimanus Dennis from Trinidad, and M.
rosulatus Desjardin & R.H. Petersen from New
Zealand. Marasmius arimanus differs in
Basidiospores (17-)20-22 × 4 µm [xm =
forming mummy brown pilei 2-3 mm diam.
20.1 ± 1.3 × 4.0 ± 0 µm, Q = 4.3-5.5, Qm = 5.0,
and basidiospores 11-14 × 3.5-4 µm (fide
n = 9 spores, s = 1 specimen], fusoid to
Singer, 1976). Marasmius rosulatus differs in
narrowly clavate, smooth, hyaline, inamyloid,
forming pure white basidiomes with
thin-walled. Basidia not observed. Basidioles
basidiospores 7.8-9.6 × 4.0-4.8 µm and grows
fusoid to clavate. Cheilocystidia common, of
on Cyperales leaves (Desjardin and Petersen,
Rotalis-type broom cells; main body 18-23 ×
1989).
10-18 µm, subglobose to broadly clavate,
hyaline, inamyloid, thin-walled; divergent 2. Marasmius delicatulus Wannathes,
setulae 1-2 × 1 µm, knob-like to conical, obtuse, Desjardin & Lumyong, sp. nov.
dense over upper half of cell, hyaline, thin- to (Figs 3, 17-1)
thick-walled. Pleurocystidia absent. Pileipellis MycoBank: MB512413
hymeniform, not mottled, composed of Rotalis- Etymology: ‘delicatulus’ = delicate; referring to
type broom cells; main body 27-35 × 16-22 µm, the tiny, delicate basidiomes.
broadly clavate, subglobose, pyriform, or Pileus 1-3 mm diametro, convexus usque late
convexus, subumbilicatus, sine papilla, striatus glaber,
irregular in outline, hyaline, inamyloid, thin- hebetatus, flavo-albus usque brunneo-creameus cum
walled; divergent setulae 1-2 × 1 µm, knob-like macula laete brunnea ad centrum. Contextus flavo-albus,
to conical, obtuse, often dense over upper half tenuis. Lamellae adnatae usque collarium parvum,
of cell, yellow, thick-walled. Pileus trama distantes (6-10), angustae, flavo-albae usque creameae,
interwoven, inamyloid. Lamellar trama hyphae haud marginatae, haud intervenosae. Stipes 12 × 0.1-0.2
mm, centralis, cylindratus, filo metallico similis, nodis
3-6 µm diam., interwoven to regular, dispersis, glaber, insititius, brunneus usque rubro-
cylindrical, smooth, hyaline, inamyloid, thin- brunneus, plerumque rhizomorphis brunneis. Odor
walled, non-gelatinous. Stipitipellis hyphae 5- saporque non propria. Basidiosporae 8-10 × 3.5-5 µm,
10 µm diam., parallel, cylindrical with ellipsoideae, laeves, hyalinae, inamyloideae,
scattered finger-like projections, light yellow to tenuitunicatae. Basidia non observata. Basidiolae
221
fusoideae usque clavatae. Cheilocystidia vulgaria, typi Basidiospores 8-10 × 3.5-5 µm [xmr = 8.6
Rotalis; 13-16 × 9-13 µm, clavata usque late clavata,
× 3.9-4.2 µm, xmm = 8.6 ± 0 × 2.7 ± 0.3 µm,
subglobosa, hyalina, inamyloidea, tenuitunicata; setulis
divergentibus 1-2 × 1 µm, gongylodibus usque conicis, Qmr = 2.1-2.2, Qmm = 2.2 ± 0.1, n = 25 spores, s
obtusis, confertis ad dimidium superiorem, hyalinis = 2 specimens], ellipsoid, smooth, hyaline,
usque pallide flavis, crassetunicatis. Pleurocystidia nulla. inamyloid, thin-walled. Basidia not observed.
Pileipellis hymeniformis, haud maculosus, typi Rotalis; Basidioles fusoid to clavate. Cheilocystidia
18-23 × 14-18 µm, late clavatus usque subglobosus, common, of Rotalis-type broom cells; main
pyriformis, hyalinus, inamyloideus, tenuitunicatus;
setulis divergentibus 1-2 × 1 µm, gongylodibus usque body 13-16 × 9-13 µm, clavate to broadly
conicis, obtusis, confertis ad dimidium superiorem, clavate, subglobose, hyaline, inamyloid, thin-
brunneo-flavis usque laete brunneis, crassetunicatis. walled; divergent setulae 1-2 × 1 µm, knob-like
Trama pilei intertexta, inamyloidea. Trama lamellae to conical, obtuse, dense over upper half of cell,
intertexta, hyphis 4-8 µm diametro, cylindratis usque
inflatis, laevibus, hyalinis, inamyloideis, tenuitunicatis,
hyaline to pale-yellow, thick-walled. Pleuro-
haud gelatinosis. Stipitipellis parallelus, hyphis 3-5 µm cystidia absent. Pileipellis hymeniform, not
diametro, cylindratis, brunneis, laevibus, leniter mottled, composed of Rotalis-type broom cells;
dextrinoideis usque inamyloideis, crassetunicatis (usque main body 18-23 × 14-18 µm, broadly clavate
ad 1 µm), haud gelatinosis. Trama stipitis parallela, to subglobose, pyriform, hyaline, inamyloid,
hyphis 2-4 µm diametro, cylindratis, hyalinis, laevibus,
inamyloideis, tenuitunicatis, haud gelatinosis. thin-walled; divergent setulae 1-2 × 1 µm,
Caulocystidia nulla. Fibulae praesentes in omnibus knob-like to conical, obtuse, dense over upper
texturis. half of cell, brownish yellow to light brown,
Typus: Phrae Province, Maung District, Pa Dang, thick-walled. Pileus trama interwoven, inamy-
Nong Kam Village, Nong Kam reservoir, dispersus
loid. Lamellar trama hyphae 4-8 µm diam.,
usque gregarius in folis plantae dicotyledoneae, 18
August2005, N. Wannathes 426 (CMU: holotypus; interwoven, cylindrical to inflated, smooth,
SFSU: isotypus). hyaline, inamyloid, thin-walled, non-gelatinous.
Pileus 1-3 mm diam., convex to broadly Stipitipellis hyphae 3-5 µm diam., parallel,
convex, subumbilicate, without papilla, striate, cylindrical, brown, smooth, weakly dextrinoid
glabrous, dull, yellowish white to brownish to inamyloid, thick-walled (up to 1 µm), non-
cream with a light brown central spot. Context gelatinous. Stipe trama hyphae 2-4 µm diam.,
yellowish white, thin. Lamellae adnate to a parallel, cylindrical, hyaline, smooth, inamyloid,
small collarium, distant (6-10), narrow, thin-walled, non- gelatinous. Caulocystidia
yellowish white to cream, non-marginate, non- absent. Clamp connections present in all tissues.
intervenose. Stipe 5-12 × 0.1-0.2 mm, central, Habit, habitat and known distribution:
Scattered to gregarious on dicotyledonous
leaves, Northern Thailand.
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).
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
Fig. 3. Marasmius delicatulus (N. Wannathes 426). 1. central, insititious stipe not arising from
Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
Pileipellis Scale bar 1 = 4.5 mm, 2-4 = 10 µm
rhizomorphs, and moderately-sized basidio-
spores (xmm = 8.6 × 4.1 µm; Qmm = 2.2). The
cylindrical, wiry, with scattered nodes, new Thai species is morphologically similar to
glabrous, insititious, brown to reddish brown; several other species. Marasmius diminutivus
brown rhizomorphs usually present. Odor and Y.S. Tan, Desjardin & Vikineswary, from
taste not distinctive. Malaysia, differs in forming smaller (0.5-1 mm
222
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, Fig. 4. Marasmius somalomoensis (N. Wannathes 232).
differs in forming white pilei 2-5 mm diam and 1. Basidiomes 2. Basidioles 3. Cheilocystidia 4.
a more robust stipe up to 0.5 mm thick (Singer, Basidiospores 5. Pileipellis, Scale bar 1 = 10 mm, 2-5 =
10 µm
1976).
brown, thick-walled. Pileus trama interwoven,
3. Marasmius somalomoensis Antonín, inamyloid. Lamellar trama hyphae 2-4(-8) µm
Mycotaxon 88: 66. 2003. (Figs 4, 17-2) diam., interwoven, cylindrical to inflated,
Pileus 2-4 mm diam., convex, umbilicate, smooth, hyaline, inamyloid, thin-walled, non-
with a tiny papilla when young, papilla absent gelatinous. Stipitipellis hyphae 3-5 µm diam.,
in age, sulcate, glabrous, dull, with a reddish subparallel, cylindrical, dark brown, smooth,
brown spot in centre, surrounded by a pale weakly dextrinoid to inamyloid, thick-walled
zone, reddish brown with a hint of olive (up to 1 µm), non-gelatinous. Stipe trama
elsewhere. Context cream, thin. Lamellae hyphae 3-6 µm diam., subparallel, cylindrical,
adnate to a small collarium, subdistant (12-18), hyaline, smooth, inamyloid, thin- to thick-
broad (0.5-1 mm), cream with brown edge, walled (up to 1 µm), non-gelatinous.
non-intervenose. Stipe 8-47 × 0.3 mm, central, Caulocystidia absent. Clamp connections
cylindrical, wiry, with scattered nodes, present in all tissues.
glabrous, insititious, apex cream, base dark Habit, habitat and known distribution:
brown; rhizomorphs absent. Odor and taste not Scattered to gregarious on dicotyledonous
distinctive. leaves, Northern Thailand.
Basidiospores 8-10 × 4-5 µm [xm = 8.8 Material examined: Thailand. Chiang Mai
± 0.6 × 4.3 ± 0.3 µm, Q = 1.8 - 2.2, Qm = 2.1, n Province, Doi Suthep-Pui National Park, Sangasabhasri
Land N 18º 48.4’ E 98º 54.6’ alt. 1,146 m., 27 July 2004,
= 25 spores, s = 1 specimen], ellipsoid, smooth,
N. Wannathes 232 (CMU, SFSU).
hyaline, inamyloid, thin-walled. Basidia not Discussion: Maramius somalomoensis,
observed. Basidioles fusoid to clavate. originally described from material collected in
Cheilocystidia common, of Rotalis-type broom Cameroon, is characterized by having a light
cells; main body 11-19 × 7-11 µm, clavate to brown to brownish orange pileus with darker
broadly clavate, hyaline, inamyloid, thin- papilla surrounded by a pale zone, moderately
walled; divergent setulae 2-3 × 1 µm, knob-like distant lamellae, basidiospores in the range 7-
to conical, obtuse, dense over upper half of cell, 10 × 3.7-5.5 µm, and Rotalis-type pileipellis
brownish yellow to yellow, thick-walled. cells (Antonín, 2003). The Thai material differs
Pleurocystidia absent. Pileipellis hymeniform, from the protologue only in forming pilei with
not mottled, composed of Rotalis-type broom more reddish brown tones, more lamellae (12-
cells; main body 18-23 × 14-18 µm, broadly 18 vs 11-13) with brown edges, and slightly
clavate, subglobose, turbinate to pyriform, longer stipes (up to 47 mm vs 10-17 mm).
hyaline, inamyloid, thin-walled; divergent
setulae 1-3 × 1 µm, knob-like to conical, obtuse, 4. Marasmius apatelius Singer, Bull, Jard.
dense over upper half of cell, brown to light 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 = 1.7-
2.3, n = 25 spores, s = 7 specimens], ellipsoid,
curved in profile, smooth, hyaline, inamyloid,
thin-walled. Basidia 16-24 × 7-9 µm, clavate 4- Fig. 5. Marasmius apatelius (N. Wannathes 251). 1.
spored. Basidioles clavate to fusoid. Basidiomes 2. Basidioles and basidia 3. Basidiospores 4.
Cheilocystidia common, of Rotalis-type broom Cheilocystidia 5. Pileipellis, Scale bar 1 = 10 mm, 2-5 =
cells; main body (13)17-32(-34) × (7-)12-20 10 µm
µm, clavate to broadly clavate, pyriform, leaves, bamboo debris or wood, Africa and
hyaline, inamyloid, thin-walled; divergent northern Thailand.
setulae 0.5-4 × 0.8-1.5 µm, knob-like to conical, Material examined: Thailand. Chiang Mai
obtuse, pale-yellow to light brown, thick- Province, Doi Suthep-Pui National Park, Mokfa
Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
walled. Pleurocystidia absent or of scattered m. 3 July 2003, N. Wannathes 017 (CMU, SFSU); same
Rotalis-type broom cells like the cheilocystidia. location, 21 August 2004, N. Wannathes 265 (CMU);
Pileipellis hymeniform, not mottled, composed Chiang Mai Province, Mushroom Research Centre, 27
of Rotalis-type broom cells; main body 16-35 × km marker on Hwy 1095, 27 July 2003, N. Wannathes
035 (CMU); same location, 27 July 2003, N. Wannathes
10-20 µm, clavate to broadly clavate, pyriform
043 (CMU); Chiang Mai province, Chiang Mai
or turbinate, hyaline, inamyloid, thin- to thick- University, 11 August 2004, N. Wannathes 251 (CMU,
walled; divergent setulae 1-3 × 0.5-1 µm, SFSU); Phrae Province, Maung District, Pa Dang, Nong
knob-like to conical, obtuse, dense over upper Kam Village, near Nong Kam reservoir, 18 August 2005,
half of cell, yellow to light brown, thick-walled. N. Wannathes 427 (CMU, SFSU); Chiang Mai Province,
Chiang Mai University, close to Ang Kaew reservoir, 22
Pileus trama interwoven, inamyloid. Lamellar August 2005, N. Wannathes 437 (CMU, SFSU)
trama hyphae 3-6(-13) µm diam., interwoven Discussion: The specimens from
to regular, cylindrical to inflated, smooth, Thailand matched quite closely the descriptions
hyaline, inamyloid, thin-walled, non-gelatinous. of populations from Africa (Singer 1965;
Stipitipellis hyphae 3-6 µm diam., parallel to Pegler 1977), except for forming a few more
subparallel, cylindrical, light brown to brown, lamellae per basidiome (13-14 vs 9-12).
smooth, weakly dextrinoid to inamyloid, thin- Distinctive features of this species include an
to thick-walled (up to 1 µm), non-gelatinous. entirely brown, umbilicate pileus lacking a
Stipe trama hyphae (2-)3-7(-10) µm diam., dark central spot, collariate, subdistant lamellae
parallel, cylindrical, hyaline, smooth, inamy- with or without a pale brown edge, an absence
loid, thin-walled, non-gelatinous. Caulocystidia of well-developed rhizomorphs, and small
absent. Clamp connections present in all tissues. basidiospores with mean 7.2 × 3.9 µm. This is
Habit, habitat and known distribution: the first report of this species outside Africa.
Scattered to gregarious on dicotyledonous Marasmius tubulatus Petch, is similar, but
224
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, Fig. 6. Marasmius berambutanus (N. Wannathes 044). 1.
distant (8), broad, cream, non-marginate, non- Basidiomes 2. Basidioles 3. Basidiospores 4.
intervenose. Stipe 3-12 × 0.1-0.2 mm, central, Cheilocystidia 5a. Siccus-type pileipellis 5b. Cells
transitional between broom cells and setae 5c. Pileosetae,
cylindrical, wiry, with scattered nodes, Scale bar 1 = 2 mm, 2-5 = 10 µm
glabrous, insititious, apex white, base black,
arising directly from black rhizomorphs. Odor µm, cylindrical, conical to subacute, thick-
and taste not distinctive. walled; c) pileosetae 36-82 × 6-8 µm, common,
Basidiospores 7-10 × (3.5-)4-5.5 µm [xm lanceolate to fusoid, hyaline to yellow,
= 8.4 ± 0.6 × 4.7 ± 0.6 µm, Q = 1.4 - 2.3, Qm = inamyloid, thick-walled. Pileus trama interwo-
2.1, n = 25 spores, s = 1 specimen], ellipsoid, ven, inamyloid. Lamellar trama hyphae 3-6 µm
smooth, hyaline, inamyloid, thin-walled. diam., interwoven, cylindrical to inflated,
Basidia not observed. Basidioles cylindrical to smooth, hyaline, inamyloid, thin-walled, non-
clavate. Cheilocystidia abundant, of Siccus- gelatinous. Stipitipellis hyphae (2-)3-5 µm
type broom cells; main body 13-20 × 7-11 µm, diam., subparallel, cylindrical, dark brown,
cylindrical to clavate or broadly clavate, smooth, dextrinoid, thick-walled (up to 1 µm),
hyaline, inamyloid, thin-walled; apical setulae non-gelatinous. Stipe trama hyphae 2-4 µm
3-19 × 1.5-2.5 µm, cylindrical, often wavy and diam., subparallel, cylindrical, hyaline, smooth,
forked, obtuse to subacute, hyaline, thin-walled. inamyloid, thin-walled, non-gelatinous.
Pleurocystidia absent. Pileipellis hymeniform, Caulocystidia absent. Clamp connections
weakly mottled, composed of 3 types of cells: a) present in all tissues.
Siccus-type broom cells with main body 9-20 ×
Scattered to gregarious on undetermined
8-12 µm, clavate to broadly clavate or pyriform,
dicotyledonous leaves or on wood, Java,
hyaline, inamyloid, thin-walled; apical setulae
Malaysia and northern Thailand.
3-15 × 1-2.5 µm, conical to cylindrical, obtuse Material examined: Thailand. Chiang Mai
to subacute, yellow, thin-walled; b) transitional Province, Mushroom Research Centre, 27 km marker on
cells, similar to Siccus-type broom cells with Hwy 1095, 27 June 2003, N. Wannathes 004 (CMU).
main body 5-11 × 6-9 µm, cylindrical to Discussion: Marasmius berambutanus is
clavate, with 3-5 large apical setulae 9-28 × 2-3 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.
by Tan et al. (2007) based on Malaysian
material.
6. Marasmius purpureisetosus Corner, Nova

Hedwigia 111: 90. 1996. (Figs 7, 17-5)
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, Fig. 7. Marasmius purpureisetosus (N. Wannathes 155).
wiry, with scattered nodes, glabrous, insititious, 1. Basidiomes 2. Basidiospores 3a. Cheilocystidia 3b.
Cheilosetae 4a. Siccus-type pileipellis 4b. Cells transitional
apex creamish brown, base light brown to between broom cells and setae 4c. Peleosetae, Scale bar 1 =
brown; rhizomorphs present. Odor and taste 6.5 mm, 2-4 = 10 µm
not distinctive.
Basidiospores 8-10 × 3-5 µm [xm = 8.6 ± cells with main body 17-36 × 9-12 µm, clavate
0.6 × 3.9 ± 0.4 µm, Q = 1.6 - 2.9, Qm = 2.3, n = to broadly clavate, with only 2-3 large apical
25 spores, s = 1 specimen], ellipsoid, smooth, setulae 15-27 × 3-5 µm, cylindrical, conical to
hyaline, inamyloid, thin-walled. Basidia not subacute, thick-walled; c) pileosetae 80-350 ×
observed. Basidioles fusoid to clavate. 8-12 µm, common, lanceolate to fusoid, brown,
Cheilocystidia abundant, composed of 2 types inamyloid, thick-walled (up to 3 µm). Pileus
of cells: a) Siccus-type broom cells with main trama interwoven, inamyloid. Lamellar trama
body 13-16 × 7-10 µm, clavate to broadly hyphae 3-8(-10) µm diam., interwoven, cylin-
clavate, hyaline to yellow, inamyloid, thick- drical to inflated, smooth, hyaline, inamyloid,
walled; apical setulae 5-10 × 2-3 µm, thin-walled, non-gelatinous. Stipitipellis hy-
cylindrical to conical, obtuse, ranging from 2-4 phae 3-5 µm diam., subparallel, cylindrical,
per cell, yellow, thick-walled; b) non-setulose dark brown, smooth, inamyloid to weakly
cells, 23-38 × 6-8 µm, fusoid, hyaline, dextrinoid, thick-walled (up to 1 µm), non-
inamyloid, thin- to thick-walled. Pleurocystidia gelatinous. Stipe trama hyphae 3-6 µm diam.,
absent. Pileipellis hymeniform, not mottled, subparallel, cylindrical, hyaline, smooth,
composed of 3 types of cells: a) Siccus-type weakly dextrinoid, thin-walled, non-gelatinous.
broom cells with main body 14-20 × 8-15 µm, Caulocystidia absent. Clamp connections
clavate to broadly clavate or pyriform, hyaline present in all tissues.
to pale yellow, inamyloid, thick-walled; apical Habit, habitat and known distribution.
setulae 4-15 × 2-3 µm, conical to cylindrical, Scattered to gregarious on undetermined
obtuse, brown to dark brown, thick-walled; b) dicotyledonous leaves, Java, Malaysia,
transitional cells, similar to Siccus-type broom northern Thailand and Singapore.
226
Fungal Diversity

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 Fig. 8. Marasmius brevicollus (N. Wannathes 128). 1.
papilla, glabrous to minutely velutinous, dull, Basidiomes 2. Basidia 3. Basidiospores 4. Cheilocystidia
brownish red to brownish purple. Context 5. Pileipellis, Scale bar 1 = 6.5 mm, 2-5 = 10 µm
collarium, distant to subdistant (6-9), broad, thin- to thick-walled; apical setulae 1-3(-6) × 1-
cream with brown edge, non-intervenose. Stipe 1.5 µm, conical to cylindrical, subacute to
10-25 × 0.1-0.2 mm, central, cylindrical, wiry, obtuse, yellowish brown to dark brown, thick-
with or without nodes, glabrous, insititious, walled. Pileus trama interwoven, inamyloid.
brownish yellow, rhizomorph absent. Odor and Lamellar trama hyphae 3-5(-7) µm diam.,
taste not distinctive. interwoven, cylindrical to inflated, smooth,
Basidiospores (16-)18-22(-23) × (3.5-)4- hyaline, inamyloid, thin-walled, non-gelatinous.
5.5 µm [xmr = 19.0-21.1 × 4.0-5.1 µm, xmm = Stipitipellis hyphae 2-5 µm diam., subparallel,
cylindrical, yellowish brown, smooth, dextri-
19.7 ± 0.6 × 4.6 ± 0.5 µm, Qmr = 3.8-4.9, Qmm
noid, thin-walled, non-gelatinous. Stipe trama
= 4.6 ± 0.5, n = 25 spores, s = 5 specimens],
hyphae 2-5 µm diam., parallel, cylindrical,
clavate to fusoid, smooth, hyaline, inamyloid,
hyaline, smooth, inamyloid, thin-walled, non-
thin-walled. Basidia 30-40 × 10-12 µm, gelatinous. Caulocystidia absent. Clamp
cylindrical to clavate, 4-spored. Basidioles connections present in all tissues.
fusoid to clavate. Cheilocystidia abundant, of Habit, habitat and known distribution:
Siccus-type broom cells; main body 12-26 × 6- Scattered to gregarious on undetermined
15(-19) µm, cylindrical to clavate, broadly monocotyledonous leaves and stems, Malaysia,
clavate or pyriform, often branched, hyaline, northern Thailand and Singapore.
inamyloid, thin-walled; apical setulae 1-6 × 1- Material examined: Thailand. Chiang Mai
1.5 µm, conical to cylindrical, obtuse to Province, Mae Taeng District, Pa Deng Village,
subacute, yellowish brown to dark brown, Pathammikaram temple on Hwy 1095, 7 August 2003, N.
Wannathes 094(CMU) and 095 (CMU, SFSU); same
thick-walled. Pleurocystidia absent. Pileipellis
location, 23 August 2003, N. Wannathes 123 and 126
hymeniform, mottled, composed of Siccus-type (CMU); same location, 25 August 2003, N. Wannathes
broom cells; main body 7-27(-30) × 9-15 µm, 128 (CMU, SFSU).
clavate to broadly clavate, pyriform or irregular Discussion: Diagnostic features of
in outline, usually branched, hyaline, inamyloid, 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 Fig. 9. Marasmius purpureobrunneolus (N. Wannathes
Singaporean material (Corner, 1996) in 370). 1. Basidiomes 2. Cheilocystidia 3. Basidiospores 4.
Pileipellis, Scale bar 1 = 13.5 mm, 2-4 = 10 µm
forming a shorter stipe (10-25 mm vs 20-60
mm) and paler stipe color (brownish yellow vs Siccus-type broom cells; main body 12-17 × 8-
dark brown). The Singaporean material has a 13 µm, clavate to broadly clavate or irregular
broad range of the lamellae number (6-12) in outline, hyaline, inamyloid, thin- to thick-
which encompasses the range of lamellae walled; apical setulae 1-3 × 1 µm, cylindrical
number for Thai and Malaysian populations. to conical, obtuse to subacute, pale yellow,
Corner (1996) suggested that M. brevicollus thick-walled. Pleurocystidia absent. Pileipellis
was allied with M. nigrobrunneus (Pat.) Sacc., hymeniform, weakly mottled, composed of
but our molecular data (Fig. 1) indicate that
Siccus-type broom cells; main body 15-25 × 7-
M. brevicollus is allied with M. purpureo-
15 µm, clavate to broadly clavate, pyriform or
brunneolus and together they are placed in a
irregular in outline, hyaline, inamyloid, thin- to
clade distantly related to M. nigrobrunneus.
thick-walled; apical setulae 1-3 × 1 µm, conical
to cylindrical, subacute to obtuse, yellowish
for M. brevicollus by Tan et al. (2007) based on
Malaysian material.
inamyloid. Lamellar trama hyphae 4-8(-13)
µm diam., regular to interwoven, cylindrical to
8. Marasmius purpureobrunneolus Henn.,
inflated, smooth, hyaline, inamyloid, thin-
Monsunia 1: 151. 1900. (Fig. 9)
= Marasmius acierufus Corner, Beih. Nova
walled, non-gelatinous. Stipitipellis hyphae 3-5
Hedwigia 111: 25. 1996 µm diam., subparallel, cylindrical, yellowish
Pileus 2-5 mm diam., hemispherical to brown to brown, smooth, dextrinoid, thick-
truncately conical, umbilicate, with or without walled (up to 1 µm), non-gelatinous. Stipe
a papilla, striate, pruinose to pubescent, dull, trama hyphae (2-)3-6 µm diam., subparallel,
dark reddish brown (9F8) to violet brown cylindrical, hyaline, smooth, inamyloid, thin-
(11F4) with a reddish brown spot in centre. walled, non-gelatinous. Caulocystidia absent.
Context cream, thin. Lamellae adnate to a small Clamp connections present in all tissues.
collarium, subdistant (9-13), narrow, yellowish Habit, habitat and known distributions.
white with reddish brown edge, non- Scattered to gregarious on undetermined
intervenose. Stipe 10-32 × 0.1-0.2 mm, central, dicotyledonous leaves or on wood, Java,
cylindrical, wiry, with or without nodes, Malaysia and northern Thailand.
glabrous, insititious, brown (6E5) to dark
Province, Mae Taeng District, “Water Conservation taste
brown (7F7) overall; rhizomorphs absent. Odor not distinctive. Area” Highway 1,095 at 22 km marker,
and taste not distinctive. N. 19º 7.5’ E 98º 45.7’ alt. 724 m. 5 July 2004, N.
Basidiospores (12-)13-16 × 3-4 µm [xmr Wannathes 215 (CMU, SFSU); Chiang Mai Province,
= 14.0-14.4 × 3.2-3.3 µm, xmm = 14.2 ± 0.2 × 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
3.2 ± 0.1 µm, Qmr = 4.4-4.5, Qmm = 4.4 ± 0.1, n June 2005, N. Wannathes 370 (CMU, SFSU).
= 25 spores, s = 2 specimens], narrowly Discussion: Marasmius purpureobrun-
ellipsoid to fusoid, smooth, hyaline, inamyloid, neolus is characterized by striate, dark reddish
thin-walled. Basidia not observed. Basidioles brown to violet brown pilei, subdistant (9-13)
fusoid to clavate. Cheilocystidia abundant, of lamellae with reddish brown edges, fusoid
228
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. brevi-
collus look very similar in size and color of the Fig. 10. Marasmius pallenticeps (N. Wannathes 346). 1.
pilei. The two species differ in lamellae Basidiomes 2. Cheilocystidia 3. Basidiospores 4.
spacing (9-13 lamellae vs 6-9, respectively) Pileipellis, Scale bar 1 = 3.5 mm, 2-4 = 10 µm
and in basidiospore size (xmm = 14.2 × 3.2 µm 3-5 µm diam., regular to interwoven,
vs 19.7 × 4.6 µm, respectively). cylindrical, smooth, hyaline, inamyloid, thin-
9. Marasmius pallenticeps Singer, Fl. Neotrop. µm diam., subparallel, cylindrical, brown to
Monogr. 17: 127. 1976 (Fig. 10) dark brown, smooth, inamyloid to weakly
Pileus 0.5-1.5 mm diam., convex, dextrinoid, thick-walled (up to 1 µm), non-
subumbilicate with a papilla, striate, pruinose gelatinous. Stipe trama hyphae 1.5-3 µm diam.,
to glabrous, dull, cream to pale yellowish white subparallel, cylindrical, hyaline, smooth,
(2A2) with a dark brown papilla. Context inamyloid, thin-walled, non-gelatinous.
cream, thin. Lamellae adnate to a small Caulocystidia absent. Clamp connections
collarium, distant (7-9), narrow, cream, non- present in all tissues.
marginate, non-intervenose. Stipe 3-6 × 0.1 mm, Habit, habitat and known distribution:
central, cylindrical, wiry, glabrous, insititious, Scattered to gregarious on wood (dicotyledo-
dark brown to black overall, arising directly nous tree), Argentina, New Zealand, Indonesia
from black rhizomorphs. Odor and taste not and northern Thailand.
distinctive. Material examined: Thailand. Chiang Mai
Basidiospores (7-)8-10(-11) × 4.5-5.5 µm Province, Mae Taeng District, Mae Sae Village, Pang Sa
Ded Water Conservation Area, N 19º 14.6', E 98º 38.5'
[xm = 9.0 ± 0.8 × 4.9 ± 0.3 µm, Q = 1.4 - 2.2, alt. 962 m., 26 June 2005, N. Wannathes 346 (CMU,
Qm = 1.7, n = 25 spores, s = 1 specimen], SFSU).
ellipsoid, smooth, hyaline, inamyloid, thin- Discussion: Distinctive features of
walled. Basidia not observed. Basidioles fusoid Marasmius pallenticeps include a tiny pale
to clavate. Cheilocystidia abundant, of Siccus- cream-coloured pileus with dark brown papilla,
type broom cells; main body 11-18 × 5-10 µm, distant (7-9), collariate lamellae, and a stipe
cylindrical to clavate or broadly clavate, arising directly from rhizomorphs. Our
hyaline, inamyloid, thin-walled; apical setulae specimen is indistinguishable from the
2-4 × 1(-1.5) µm, cylindrical to conical, obtuse specimen reported from Java by Desjardin et al.
to subacute, pale yellow to hyaline, thick- (2000).
walled. Pleurocystidia absent. Pileipellis
hymeniform, weakly mottled, composed of 10. Marasmius crinis-equi F. Muell. ex
Siccus-type broom cells; main body 9-13 × 4- Kalchbr., Grevillea 8: 153. 1880. (Fig. 11)
≡ Marasmius equicrinis F. Muell. ex Berk., J.
11 µm, cylindrical to clavate or broadly clavate, Linn. Soc., Bot. 18: 383. 1881.
hyaline, inamyloid, thin-walled; apical setulae ≡ Androsaceus crinis-equi (F. Muell. ex Kalchbr.)
1-4 × 1 µm, cylindrical to conical, subacute, Overeem, Hoofd van Het Mus. Econ. Bot. Buitenzorg 1:
pale yellow, thick-walled. Pileus trama 69. 1927.
interwoven, inamyloid. Lamellar trama hyphae = 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
collarium, distant (8-10), narrow, cream to
orangish yellow, non-marginate, non-
intervenose. Stipe 4-6 × 0.1-0.2 mm, central,
cylindrical, wiry, glabrous, insititious, dark Fig. 11. Marasmius crinis-equi (N. Wannathes 182). 1.
brown to black overall, arising directly from Basidiomes 2. Cheilocystidia 3. Basidiospores 4.
copious, branched, black rhizomorphs. Odor Pileipellis, Scale bar 1 = 6.5 mm, 2-4 = 10 µm
and taste not distinctive.
Basidiospores (8-)10-12 × 4-4.5 µm [xmr politan in tropical habitats.
= 10.5-10.6 × 4.1-5.1 µm, xmm = 10.6 ± 0.1 × Province, Doi Suthep-Pui National Park, Mokfa
4.6 ± 0.7 µm, Qmr = 2.1-2.6, Qmm = 2.3 ± 0.4, n Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
= 25 spores per 2 specimens], ellipsoid, smooth, m., 25 July 2003, N. Wannathes 029 (CMU, SFSU);
hyaline, inamyloid, thin-walled. Basidia 22-23 Chiang Mai Province, Mae Taeng District, Mae Sae
Village, Pang Sa Ded Water Conservation Area, N 19º
× 6-7 µm, clavate, 4-spored. Basidioles fusoid
14.6', E 98º 38.5' alt. 962 m., 26 June 2005, N.
to clavate. Cheilocystidia abundant, of Siccus- Wannathes 348 (CMU, SFSU).
type broom cells; main body 11-22 × 7-12 µm, Discussion: Marasmius crinis-equi is one
clavate to broadly clavate, subglobose or of the more widely distributed species in the
irregular in outline, hyaline, inamyloid, thin- genus, reported thoughout tropical areas of the
walled; apical setulae 2-5 × 1-2 µm, conical to world. It is the causal agent of 'Horse-hair
cylindrical, obtuse to subacute, pale yellow, Blight' that is known to be pathogenic on tea,
thick-walled. Pleurocystidia absent. Pileipellis rubber, cocoa and nutmeg crops (Pegler, 1983).
hymeniform, weakly mottled, composed of This species grows on leaves and petioles of
Siccus-type broom cells; main body9-22 ×8-21 living or dead trees, and on monocotyledonous
µm, clavate to broadly clavate, pyriform, stems and clums (Singer, 1976) and its
subglobose or irregular in outline, often rhizomorphs are often found covering aerial
branched, hyaline, inamyloid, thin-walled; branches of living trees. Marasmius crinis-equi
apical setulae 1-5 × 1-1.5 µm, conical to is easy to recognize by a small, brown to
cylindrical, subacute to obtuse, yellow to brownish orange pileus with a dark brown
yellowish brown, thick-walled. Pileus trama central papilla, distant collariate lamellae,
interwoven, inamyloid. Lamellar trama hyphae stipes that arise directly from black rhizo-
3-8(-12) µm diam., regular to interwoven, morphs, and Siccus-type pileipellis elements
cylindrical to inflated, smooth, hyaline, inamy- and cheilocystidia.
loid, thin-walled, non- gelatinous. Stipitipellis
hyphae (2-)3-6 µm diam., subparallel, cylin- 11. Marasmius aff. crinis-equi F. Muell. ex
drical, brown to dark brown, smooth, Kalchbr., Grevillea 8: 153. 1880 (Fig. 12)
dextrinoid, thick-walled (up to 3 µm), non- Pileus 1-2 mm diam., hemispherical to
gelatinous. Stipe trama hyphae 2-5 µm diam., convex, shallowly umbilicate, papillate, striate
subparallel, cylindrical, hyaline, smooth, to sulcate, glabrous, dull, light orange (6A4) to
inamyloid, thin-walled, non-gelatinous. Caulo- greyish orange with a brown papilla. Context
cystidia absent. Clamp connections present in greyish cream, thin. Lamellae adnate to a small
all tissues. collarium, distant (5-8), broad, cream, non-
Habit, habitat and known distribution: marginate or with orangish cream edges, non-
Scattered to gregarious on wood, cosmo- intervenose. Stipe 3-8 × 0.1-0.2 mm, central,
230
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 = 8.9-
9.2 × 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. Cheilo-
cystidia abundant, of Siccus-type broom cells;
main body 10-22 × 8-13 µm, clavate to broadly
Fig. 12. Marasmius aff. crinis-equi (N. Wannathes 348).
clavate or pyriform, hyaline, inamyloid, thin- 1. Basidiomes 2. Basidioles 3. Basidiospores 4.
walled; apical setulae 1-4 × 1 µm, conical to Cheilocystidia 5. Pileipellis, Scale bar 1 = 4.5 mm, 2-5 =
cylindrical, obtuse to subacute, yellow, thick- 10 µm
hymeniform, weakly mottled, composed of of this taxon is collected, we tentatively
Siccus-type broom cells; main body 11-19 × 7- identify it as M. aff. crinis-equi.
16 µm, clavate to broadly clavate, pyriform or
irregular in outline, hyaline, inamyloid, thin- to 12. Marasmius subruforotula Singer, Bull.
Jard. Bot. Etat Brux. 34: 339. 1964 (Fig. 13)
thick-walled; apical setulae 2-7 × 1 µm,
cylindrical to conical, subacute to obtuse,
convex, umbilicate, with or without a papilla,
yellowish brown to brown, thick-walled. Pileus
striate to sulcate, minutely velutinous, dull,
trama interwoven, inamyloid. Lamellar trama
reddish brown to brown with dark brown
hyphae 3-8 µm diam., interwoven, cylindrical,
papilla or spot at the centre. Context cream,
smooth, hyaline, inamyloid, thin-walled, non-
thin. Lamellae adnate to a small collarium,
gelatinous. Stipitipellis hyphae 4-8 µm diam.,
subdistant (11-13), narrow, cream-grey, non-
subparallel, cylindrical, black to dark brown,
smooth, weakly dextrinoid to inamyloid, thick- marginate, non-intervenose. Stipe 4-10 × 0.1-
walled (up to 1 µm), non-gelatinous. Stipe 0.2 mm, central, cylindrical, wiry, glabrous,
trama hyphae 2-3(-5) µm diam., subparallel, shiny, insititious, black overall, arising from
cylindrical, hyaline, smooth, inamyloid, thin- leafy substrate or seldom directly from black
walled, non-gelatinous. Caulocystidia absent. rhizomorphs. Odor and taste not distinctive.
Scattered to gregarious on wood, northern
Thailand.
Province, Doi Inthanon National Park, Junction of
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 Fig. 13. Marasmius subruforotula (N. Wannathes 140).
crinis-equi in all taxonomically significant 1. Basidiomes 2. Cheilocystidia 3. Basidiospores 4.
details except for having broader basidiospores Pileipellis, Scale bar 1 = 10 mm, 2-4 = 10 µm
with Qmm = 1.9 (vs. 2.3) and paler pilei. The
molecular data (Fig. 1), however, indicate that Basidiospores 8-10 × 4-5 µm [xm = 9.5 ±
these specimens are not closely allied to M. 1.0 × 4.5 ± 0.5 µm, Q = 1.8 - 2.5, Qm = 2.1, n =
crinis-equi sensu stricto. Until further material 25 spores, s = 1 specimen], ellipsoid, smooth,
231
hyaline, inamyloid, thin-walled. Basidia not 13A. Marasmius nigrobrunneus (Pat.) Sacc. f.
observed. Basidioles fusoid to clavate. Cheilo- nigrobrunneus Syll. Fung. 11:37. 1895.
cystidia abundant, of Siccus-type broom cells; (Figs 14, 17-4)
main body 14-16 × 6-12 µm, cylindrical to Pileus 3-28 mm diam, hemispherical to
clavate or broadly clavate, hyaline, inamyloid, convex when young, becoming plano-convex
thin-walled; apical setulae 2-5 × 1 µm, and shallowly umbilicate in age, with or
cylindrical, often forked, subacute, pale yellow, without a small black papilla in the umbilicus,
thick-walled. Pleurocystidia absent. Pileipellis striate to plicate; surface dull, dry, glabrous to
hymeniform, weakly mottled, composed of pruinose; disc dark grey to black, margin
Siccus-type broom cells; main body 8-20 × 10- greyish brown (6F3) to brownish grey (9F2).
15 µm, clavate to broadly clavate or pyriform, Context thin, greyish white to cream. Lamellae
hyaline, inamyloid, thin-walled; apical setulae adnate to a collarium, subdistant (9-17) with no
3-5 × 1 µm, cylindrical to conical, subacute, lamellula, non-intervenose, broad (2-4 mm),
yellowish brown to brown, thick-walled. Pileus yellowish white when young, becoming pale-
trama interwoven, inamyloid. Lamellar trama yellow in age, with brown to dark brown edges.
hyphae 5-12(-18) µm diam., interwoven, Stipe 52-214 × 0.3-0.5 mm, central, cylindrical,
cylindrical to inflated, smooth, hyaline, wiry, shiny, solid, rarely with a few nodes,
inamyloid, thin-walled, non-gelatinous. Stipiti- glabrous, insititious, apex olive brown grading
pellis hyphae 3-6 µm diam., subparallel, to dark brown at the base when young, dark
cylindrical, dark brown to black, smooth, brown overall in age, arising directly from
inamyloid, thick-walled (up to 1 µm), non- black rhizomorphs. Odor and taste not
gelatinous. Stipe trama hyphae 3-6 µm diam., distinctive.
subparallel, cylindrical, hyaline to yellow, Basidiospores (6-)8-10(-13) × (3-)4-5(-7)
smooth, inamyloid, thin-walled, non-gelatinous. μm [xmr = 7.4-10.6 × 4.1-6.3 μm, xmm = 9.0 ±
Caulocystidia absent. Clamp connections 0.9 × 4.7 ± 0.6 μm, Q = 1.4-2.9, Qmr = 1.7-2.2,
present in all tissues. Qmm = 1.9 ± 0.1, n = 25 spores, s = 11
Habit, habitat and known distribution: specimens], ellipsoid, smooth, hyaline,
Scattered to gregarious on undetermined inamyloid, thin-walled. Basidia 25-27 × 5.0-7.5
dicotyledonous leaves or on wood, Africa and μm, clavate to cylindrical, 4-spored. Basidioles
Northern Thailand. fusoid to cylindrical. Cheilocystidia common,
Material examined: Thailand. Chiang Mai of Siccus-type broom cells; main body (11-)14-
Province, Doi Suthep-Pui National Park, Sangasabhasri
Land on the way to Huai Kok Ma Village, N 18º 48.4’ E 20(-30) × (4-)7-14(-19) μm, cylindrical to
98º 54.6’ alt. 1,146 m., 29 August 2003, N. Wannathes clavate, pyriform or sometimes irregular in
140 (CMU, SFSU). outline, hyaline, thin- to thick-walled; apical
Discussion: Marasmius subruforotula is setulae (2-)3-9(-15) × 1-2(-2.5) μm, crowded,
characterzied by a small (2-6 mm), reddish cylindrical or irregular in outline, obtuse to
brown pileus, subdistant (11-13) collariate subacute, dark grey to black, inamyloid, thick-
lamellae, a short stipe 4-10 mm long, and walled. Pleurocystidia absent. Pileipellis
ellipsoid basidiospores with mean 9.5 × 4.5 µm. hymeniform, mottled to weakly mottled,
The Thai specimens differ from M. composed of 3 types of cells: a) Siccus-type
nigrobrunneus “f. cinnamomeus” in forming broom cells with main body (11-)15-27 ×
reddish brown pilei and usually arising directly (7-)10-15(-18) μm, clavate to broadly clavate,
from leaves, although some basidiome arise pyriform or sometimes irregular in outline,
from rhizomorphs. Our molecular data (Fig. 1) yellowish brown, inamyloid, thin- to thick-
suggest that this species is distinct from M. walled; apical setulae 2-9 × 1-2(-2.5) μm,
nigrobrunneus and its forms, and is closer to M. cylindrical to conical or irregular in outline,
curreyi. Until further material is available for obtuse to subacute, dark grey to black,
comparison with the African, M. subruforotula, inamyloid, thin- to thick-walled; b)
we are only tentatively accepting the Thai Globulares-type cells, 14-28 × 11-12 μm,
species as M. subruforotula. subglobose to clavate, sometimes irregular in
232
Fungal Diversity
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,
Fig. 14. Marasmius nigrobrunneus (N. Wannathes 294). SFSU).
1. Basidiomes 2. Basidioles 3. Basidiospores 4. Discussion: The specimens of Marasmius
Cheilocystidia 5a. Siccus-type pileipellis 5b. Cells nigrobrunneus from northern Thailand match
transitional between broom cells and non-setulose cells quite closely those reported by Singer (1958,
5c. Globulares-type pileipellis, Scale bar 1 = 20 mm, 2-5 1976) and Pegler (1983, 1986) from Sri Lanka,
= 10 µm
India, Sierra Leone, Trinidad, Bolivia and
Argentina, including forming lamellae with
outline, dark grey to black, inamyloid, thin-
greyish brown edges and stipes that often arise
walled; c) cells transitional between broom
from black rhizomorphs. They differ only
cells and non-setulose cells with main body 20-
slightly from specimens reported from Vietnam
22 × 15-17 μm, broadly clavate to pyriform
(type; Patouillard, 1891) and Papua New
with a few knob- like apical setulae 6-13 × 4-8 Guinea (Desjardin and Horak, 1997) that lack a
μm, dark grey to black, thin- to thick-walled. colored lamellar edge, and were not reported to
Pileus trama interwoven; lamellar trama arise from black rhizomorphs. The species is
regular; hyphae 3-7 μm diam, cylindrical to commonly found on bamboo debris and the
inflated, smooth, hyaline, inamyloid, thin- combination of dark grey to greyish brown
walled, non-gelatinous. Stipitipellis hyphae 4-8 pileus, distant collariate lamellae with or
μm diam, parallel, cylindrical, smooth, dark without colored edge, long, black, wiry stipes
brown, dextrinoid, thick-walled (up to 2 μm), often associated with rhizomorphs, and
non-gelatinous. Stipe trama hyphae 3-8 μm versiform pileipellis broom cells are distinctive
diam, parallel, cylindrical, smooth, hyaline, features for the species. A recent type study
inamyloid, thick-walled (up to 2 μm). was published by Yang (2000).
Caulocystidia absent. Clamp connections A tetrapolar mating system was reported
present in all tissues. by Wannathes et al. (2007) for M.
Habit, habitat, and known distribution: nigrobrunneus based on specimens from
Scattered to gregarious on bamboo leaves and northern Thailand.
stems, or rarely on undetermined dicotyle-
donous leaves. Africa, Caribbean region, South 13B. Marasmius nigrobrunneus “f. cinna-
America, India, Sri Lanka, Papua New Guinea, momeus” Wannathes, Desjardin & Lumyong
Vietnam, Thailand. nom. prov. (Fig. 15)
Material examined: Thailand. Chiang Mai Etymology: ‘cinnamomeus’ = cinnamon; referring
Province, Doi Suthep-Pui National Park, Sangra to the color of the pileus.
Sabhasri Land to Huai Kok Ma Village, N18°48.402’, Pileus 2-7(-15) mm diam., obtusely
E98°54.617’, 1146 meters, 30 Jun. 2003, N. Wannathes conical, umbilicate, with or without a papilla,
010 (CMU, SFSU); Chiang Mai Province, Doi Suthep-
striate to sulcate, glabrous to pruinous, dull,
Pui National Park, Mok Fa waterfall, N19°6.581’,
E98°46.353’, 1014 meters, 13 Aug. 2003, N. Wannathes
brown to orangish brown (6C5-C8) with a
101 (CMU, SFSU); same location, 29 Jun 2004, Y. S. brown papilla or spot in the centre. Context
233
cream, thin. Lamellae adnate to a small 2-7(-12) µm diam., interwoven, cylindrical to
collarium, distant to subdistant (8-11(-15)), inflated, smooth, hyaline, inamyloid, thin-
broad, cream, non-marginate or with orange to walled, non-gelatinous. Stipitipellis hyphae 3-
brown edges, non-intervenose. Stipe 8-60 × 4(-8) µm diam., subparallel, cylindrical, brown,
0.1-0.3 mm, central, cylindrical, wiry, glabrous, smooth, dextrinoid, thin- to thick-walled, non-
insititious, brown to dark brown overall, gelatinous. Stipe trama hyphae 3-6(-10) µm
usually arising directly from rhizomorphs. diam., parallel, cylindrical, hyaline, smooth,
Odor and taste not distinctive. inamyloid, thin-walled, non-gelatinous. Caulo-
Basidiospores 8-11 × 4-5(-6) µm [xmr = cystidia absent. Clamp connections present in all
8.9-10.1 × 4.3-5.4 µm, xmm = 9.5 ± 0.5 × 4.7 ± tissues.
0.4 µm, Qmr = 1.9-2.1, Qmm =2.0 ± 0.1, n = 25 Habit, habitat and known distribution:
spores, s = 7 specimens], ellipsoid, smooth, Scattered to gregarious on undetermined
hyaline, inamyloid, thin-walled. Basidia 31-37 monocotyledonous leaves and stems, Northern
× 6-10 µm, cylindrical to clavate, 4-spored. Thailand.
Basidioles cylindrical to clavate. Cheilocystidia Province, Mae Taeng District, Pong Dued Hot Spring,
abundant, of Siccus-type broom cells; main on Hwy1095, 25 July 2003, N. Wannathes 025 (CMU,
body 10-21(-32) × 6-12 µm, cylindrical to SFSU); Chiang Mai Province, Doi Suthep-Pui National
clavate or broadly clavate, hyaline, inamyloid, Park, Mokfa Waterfall, on Hwy 1095, N19º 6.5’.E 98º
46.3’ alt. 1014 m., 25 July 2003, N. Wannathes 030
thin-walled; apical setulae 2-4(-7) × 1-1.5 µm, (CMU); same location, 29 June 2004, Y. S. Tan 281
cylindrical to conical, obtuse to subacute, pale (CMU, SFSU); same location, 25 June 2005, N.
yellow, thick-walled. Pleurocystidia absent. Wannathes 327 (CMU, SFSU); Chiang Mai Province,
Pileipellis hymeniform, weakly mottled, Mae Taeng District, Pa Deng Village, street to
composed of 2 types of cells: a) Siccus-type Pathammikaram temple on Hwy 1095, 22 August, 2003,
N. Wannathes 119 and 120 (CMU, SFSU); Chiang Mai
broom cells with main body 8-18(-24) × 7-16 Province, Mae Taeng District, New Waterfall, on 36 km.
µm, clavate to broadly clavate, pyriform or marker of Hwy1095, 20 August 2004, N. Wannathes 260
irregular in outline, hyaline, inamyloid, thin- to (CMU, SFSU).
thick-walled; apical setulae 2-6 × 1-1.5 µm, Discussion: Marasmius nigrobrunneus “f.
cylindrical to conical, subacute to obtuse, cinnamomeus” is distinct because of the
yellow to yellowish brown, thick-walled; b) following features: a brown to brownish orange
non-setulose cells, scattered, 12-17 × 8-15 µm, (cinnamon) pileus 2-7(-15) mm diam., a stipe
clavate to broadly clavate, often branched, in the range 8-60 × 0.1-0.3 mm that arises
hyaline, inamyloid, thick-walled. Pileus trama directly from rhizomorphs or from the substrate,
interwoven, inamyloid. Lamellar trama hyphae and is usually associated with non-bamboo
grasses. Marasmius nigrobrunneus f. nigro-
brunneus 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.
Fig. 15. Marasmius nigrobrunneus “f. cinnamomeus” (N. curreyi as outgroup, indicates that f.
Wannathes 119). 1. Basidiomes 2. Cheilocystidia 3. nigrobrunneus forms a clade distinct from “f.
Basidiospores 4. Pileipellis, Scale bar 1 = 20 mm, 2-4 = cinnamomeus”, providing support for
10 µm
recognition of the latter as a distinct form (Fig. 16)
234
Fungal Diversity
M. nigrobrunneus NW162
88
M. nigrobrunneus “f. cinnamomeus” NW119
74
M. nigrobrunneus “f. cinnamomeus” TYS281
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, broadly clavate or sometimes irregular in

Desjardin, & Lumyong, Mycol. Res. 111: 991. outline, rarely lobed, hyaline, thick-walled
2007. (Fig. 18) apically, thin-walled elsewhere; apical setulae
Pileus 2-4 mm diam, hemispherical to (1-)3-6(-7) × 1-2 μm, cylindrical to conical,
convex, subumbilicate to depressed, with or usually wavy in outline, obtuse to subacute,
without a papilla, striate, dull, dry, glabrous, pale yellow, inamyloid, thin- to thick-walled.
light yellow (4A4) to yellowish white (3A2) Pileus trama and lamellar trama interwoven;
with a reddish brown central spot. Context thin, hyphae (2-)3-6 μm diam, cylindrical, smooth,
yellowish white (3A2). Lamellae adnate to a hyaline, inamyloid, thin-walled, non-gelatinous.
collarium, distant (9-10) with no lamellula, Stipitipellis hyphae 3-7 μm diam, parallel,
non-intervenose, narrow, cream, non-marginate. cylindrical, smooth, brown, weakly dextrinoid
Stipe 11-35 × 0.1-0.3 mm, central, cylindrical, to dextrinoid, thin-walled, non-gelatinous;
tough, terete, glabrous, insititious, yellowish Stipe trama hyphae (2-)4-8 μm diam, parallel,
white at apex, brown to dark brown at base; cylindrical, smooth, hyaline, weakly dextrinoid
rhizomorphs absent. Odor and taste not to dextrinoid, thin- to thick-walled ( up to 1
distinctive. μm). Caulocystidia absent. Clamp connections
Basidiospores (7-)8-10(-12) × 4-5(-6.5) present in all tissues.
μm [xmr = 8.4-9.4 × 4.4-5.1 μm, xmm = 8.9 ± 0.7 Habit, habitat, and known distribution.
× 4.8 ± 0.5 μm, Q = 1.4-2.2, Qmr = 1.8-1.9, Qmm Scattered to gregarious on undetermined
= 1.8 ± 0.1, n = 25 spores, s = 2 specimens], dicotyledonous leaves or wood. Northern
ellipsoid, smooth, hyaline, inamyloid, thin- Thailand.
walled. Basidia 25-30 × 6-8 μm, clavate, 4- Material examined. Thailand. Chiang Mai
Province, Doi Suthep-Pui National Park, Mok Fa
spored. Basidioles clavate to fusoid. Cheilo-
waterfall on Hwy 1095, N19º 6.581’, E98º 46.353’, 1014
cystidia common, of Siccus type broom cells; meters, 13 Aug. 2003, N. Wannathes106 (CMU, SFSU);
main body (7-)10-18 × (4-)7-12 μm, clavate to Chiang Mai Province, Doi Inthanon National Park,
broadly clavate, pyriform, cylindrical or junction of Hwy1009 and road to Mae Chem, N18º
irregular in outline, rarely lobed, hyaline, thin- 31.58’, E98º 29.64’, 1703 meters, 28 Jul. 2004, N.
Wannathes 244 (CMU: holotype; SFSU: isotype);
walled; apical setulae (2-)4-6(-7) × 0.8-1.5 μm, Chiang Mai Province, Mae Taeng District, New
cylindrical to conical or irregular in outline, Waterfall, on 36 km. marker of Hwy1095, N. Wannathes
obtuse to subacute, hyaline to pale-yellow, 256 (CMU, SFSU).
inamyloid, thin- to thick walled. Pleurocystidia Discussion: Distinctive features of this
absent. Pileipellis hymeniform, not mottled, new species include the following: a pale
composed of Siccus-type broom cells; main yellow (stramineus), umbilicate pileus with a
body 11-20 × 6-10 μm, cylindrical to clavate, reddish brown spot; collariate, 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 rhizomorphs, has thick-walled cheilocystidia,

obvious rhizomorphs; basidiospores with mean and narrower basidiospores with mean 10.3 ×
of 8.9 × 4.8 µm and Qmm = 1.8; Siccus-type 4.3 µm and Qmm = 2.4 (Desjardin et al. 2000).
broom cells on pileipellis and lamellar edges; The new species is macromorphologically
and growth on dicotyledonous leaves. These similar to M. platyspermus Singer, a species
features indicate placement in sect. Marasmius described from Argentina, but the latter forms
subsect. Sicciformis. Marasmius straminiceps Rotalis-type broom cells and belongs in subsect.
is phenetically similar to M. conicopapillatus Marasmius (Singer 1976).
Henn., but the latter species differs in forming A tetrapolar mating system was reported
a white to cream-buff pileus with a distinctive, for M. straminiceps by Wannathes et al. (2007).
rather large conical papilla, forms black
236
Fungal Diversity
Fig. 18. Marasmius straminiceps (N. Wannathes 244). 1. Fig. 19. Marasmius guyanensis (N. Wannathes 254). 1.
Basidiomes 2. Basidioles 3. Basidiospores 4. Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5. Pileipellis, Scale bar 1 = 10 mm, 2-5 = Cheilocystidia 5. Pileipellis, Scale bar 1 = 5 mm, 2-5 =
10 µm 10 µm
15. Marasmius guyanensis Mont., Ann. Sci. pellis hyphae 3-6 μm diam, parallel, cylindrical,
Nat., Bot., sér. 4, 1:114. 1854. (Fig. 19) smooth, light brown to brown, inamyloid,
Pileus 0.5-4 mm diam, hemispherical to thick-walled (up to 2 μm), non-gelatinous. Stip
convex, umbilicate, with or without small dark trama hyphae 2-5 μm diam, parallel,
reddish brown papilla in umbilicus, striate, dull, cylindrical, smooth, hyaline to pale-yellow,
dry, glabrous; disc brown (7E6-7), margin weakly dextrinoid to inamyloid, thin-walled.
brownish yellow to orange (6B7-8). Context Caulocystidia absent. Clamp connections
thin, yellowish orange (4A6). Lamellae adnate present in all tissues.
to a collarium, distant (6-11) with no lamellula, Habit, habitat, and known distribution:
non-intervenose, narrow, yellowish orange Scattered to gregarious on undetermined
(4A6), non-marginate. Stipe 15-26 × 0.2-0.3 dicotyledonous leaves. Caribbean region,
mm, central to eccentric, cylindrical, terete, Indonesia, South America, Singapore, Thailand.
tough, glabrous, insititious, dark reddish brown Material examined: Thailand. Chiang Mai
overall; rhizomorphs usually present. Odor and 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., 2 July 2003, N. Wannathes 015 (CMU,
Basidiospores 10-12 (-15) × 3-4(-5) μm SFSU); same location, 5 July 2004, Y. S. Tan 314 (CMU,
[xmm = 11.6 ± 0.7 × 3.8 ± 0.3 μm, xmr = 11.0- SFSU); Chiang Mai Province, Mushroom Research
12.4 × 3.4-4.2 μm, Qmm = 3.1 ± 0.1, Qmr = 3.0- Centre, 27 km marker on Hwy 1095, 28 July 2003, N.
3.3, n = 25 spores, s = 5 specimens], ellipsoid, Wannathes 046 (CMU); Chiang Mai Province, Chiang
Mai University, 11 Aug. 2004, N. Wannathes 254 (CMU,
siccus-type broom cells; main body 10-21 × 6- SFSU); Chiang Mai Province, Mae Taeng District, Doi
11(-20) μm, cylindrical to pyriform, turbinate, Huai Kom Bang in Papae Village, 23 August 2004, N.
subglobose or sometimes irregular in outline, Wannathes 280 (CMU, SFSU).
rarely lobed, hyaline, inamyloid, thin-walled; Discussion: The material from Thailand
apical setulae 1-5 × 1 μm, cylindrical to conical differs slightly from that described by
or irregular in outline, obtuse to subacute, Desjardin et al. (2000) from Indonesia in
brownish yellow, inamyloid, thin- to thick- forming pilei with more brown tones and less
walled. Pileus trama interwoven, inamyloid. orange tones, and in forming basidiospores that
Lamellar trama hyphae 3-7(-9) μm diam, are slightly shorter (mean of 11.1 × 3.5 µm vs.
interwoven, cylindrical, smooth, hyaline, 12.1 × 3.6 µm). In other details, the two
inamyloid, thin-walled, non-gelatinous. Stipiti- populations are similar.
237
A tetrapolar mating system was reported non-intervenose. Stipe 6-31 × < 0.2 mm,
for M. guyanensis by Wannathes et al. (2007) central, cylindrical, wiry, glabrous, insititious,
based on Thai specimens. apex yellowish white, base dark brown;
rhizomorphs sometimes present. Odor and
16. Marasmius cafeyen Wannathes, Desjardin taste not distinctive.
& Lumyong, sp. nov. (Fig. 20) Basidiospores 9-11 × 4-5 µm [xmr = 9.4-
MycoBank: MB512414 10 × 4.2-4.4 μm, xmm = 9.8 ± 0.3 × 4.3 ± 0.1
Etymology: ‘cafeyen’ (Thai) = coffee with milk;
referring to the color of the pileus.
μm, Qmr = 2.2-2.4, Qmm = 2.3 ± 0.1, n = 25
Pileus 2-8 mm diametro, hemisphaericus usque spores, s = 3 specimens], ellipsoid, smooth,
convexus, umbilicatus, cum vel sine papilla atrobrunnea, hyaline, inamyloid, thin-walled. Basidia not
striatus usque sulcatus, minute velutinus, hebetatus,, observed. Basidioles fusoid to clavate.
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, Fig. 20. Marasmius cafeyen (N. Wannathes 130). 1.
cylindratis usque conicis, saepe furcatis, obtusis usque Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
subacutis, pallide flavis, crassetunicatis. Pleurocystidia Pileipellis, Scale bar 1 = 13 mm, 2-4 = 10 µm
nulla. Pileipellis hymeniformis, leniter maculosus, typi
Sicci; 12-18(-25) × 7-14 µm, clavatus usque late clavatus, Cheilocystidia abundant, of Siccus-type broom
pyriformis vel adumbratim inaequabilis, raro ramosus, cells; main body 9-17 × 8-13 µm, clavate to
hyalinus usque pallide flavus, inamyloideus, tenui- usque
crassetunicatus; setulis apicalibus 2-3 × 1 µm, conicis
broadly clavate or pyriform, hyaline, inamyloid,
usque cylindratis, subacutis usque obtusis, flavo- thin-walled; apical setulae 2-6 × 1 µm,
brunneis usque brunneis, crassetunicatis. Trama pilei cylindrical to conical, often forked, obtuse to
intertexta, inamyloidea. Trama lamellae regularis usque subacute, pale yellow, thick-walled.
intertexta, hyphis 4-7(-10) µm diametro, cylindratis Pleurocystidia absent. Pileipellis hymeniform,
usque inflatis, laevibus, hyalinis, inamyloideis,
tenuitunicatis, haud gelatinosis. Stipitipellis subparallelus,
weakly mottled, composed of Siccus-type
hyphis 3-5 µm diametro, cylindratis, brunneis usque broom cells; main body 12-18(-25) × 7-14 µm,
atrobrunneis, laevibus, dextrinoideis, crassetunicatis clavate to broadly clavate, pyriform or irregular
(usque ad 1 µm), haud gelatinosis. Trama stipitis in outline, seldom branched, hyaline to pale
subparallela, hyphis 2-5 µm diametro, cylindratis,
yellow, inamyloid, thin- to thick-walled; apical
hyalinis, laevibus, inamyloideis, tenuitunicatis, haud
gelatinosis. Caulocystidia nulla. Fibulae praesentes in setulae 2-3 × 1 µm, conical to cylindrical,
omnibus texturis. subacute to obtuse, yellowish brown to brown,
Typus: Chiang Mai Province, Mae Taeng District, thick-walled. Pileus trama interwoven,
Pathummikaram temple in Pa Deng Village, dispersus inamyloid. Lamellar trama hyphae 4-7(-10)
usque gregarius in folis plantae dicotyledoneae, 25
August 2003, N. Wannathes 130 (CMU: holotypus;
µm diam., regular to interwoven, cylindrical to
SFSU: isotypus). inflated, smooth, hyaline, inamyloid, thin-
Pileus 2-8 mm diam., hemispherical to walled, non-gelatinous. Stipitipellis hyphae 3-5
convex, umbilicate, with or without dark brown µm diam., subparallel, cylindrical, brown to
papilla, striate to sulcate, minutely velutinous, dark brown, smooth, dextrinoid, thick-walled
dull, margin creamish brown to creamish (up to 1 µm), non-gelatinous. Stipe trama
yellow, disc brown with a dark brown papilla hyphae 2-5 µm diam., subparallel, cylindrical,
or spot at the centre. Context cream, thin. hyaline, smooth, inamyloid, thin-walled, non-
Lamellae adnate to a small collarium, distant gelatinous. Caulocystidia absent. Clamp
(8-12), broad, yellowish white, non-marginate, connections present in all tissues.
238
Fungal Diversity
Habit, habitat and known distribution. non-intervenose, broad (0.8-1 mm), cream
Scattered to gregarious on undetermined (3A2) with orange edges. Stipe 27-34 × 0.1-0.2
dicotyledonous leaves, northern Thailand. mm, central to eccentric, cylindrical, terete,
Material examined: Thailand. Chiang Mai wiry, tough, glabrous, insititious, reddish
brown (7E6) overall; rhizomorphs absent. Odor
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 and taste not distinctive.
(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). Fig. 21. Marasmius ruforotula (N. Wannathes 257). 1.
Marasmius pallipes Singer, described from Basidiomes 2. Basidioles and basidium 3. Basidiospores
South America, differs in forming more 4. Cheilocystidia 5. Pileipellis, Scale bar 1 = 20 mm, 2-5
= 10 µm
cinnamon buff to ochraceus buff pileus, a pale
stipe (stramineus to light chestnut), and slightly Basidiospores (8-)10-12 × (4.5-)5-7 μm
broader basidiospores measuring 4.5-6.2 µm [xmr = 9.3-11.1 × 5.1-6.5 μm, xmm = 10.2 ± 0.9
wide with mean diameter near 5 µm (Singer, × 5.8 ± 0.7 μm, Q = 1.5-2.0, Qmr = 1.6-2.0, Qmm
1976). Marasmius ruforotula Singer, described = 1.8 ± 0.2, n = 25 spores, s = 1 specimen],
from North America and reported here from ellipsoid, smooth, hyaline, inamyloid, thin-
Thailand, differs in forming pilei with reddish walled. Basidia and basidioles clavate to
brown to rusty brown or brownish yellow disc cylindrical. Cheilocystidia common, of Siccus-
and greyish orange to light yellow margin, type broom cells; main body 12-15 × 6-8 μm,
orange-marginate lamellae, and broader cylindrical to pyriform, clavate or broadly
basidiospores with mean width 5.8 µm. clavate, rarely lobed, hyaline, thin-walled;
apical setulae 3-5 × 0.8-1.0 μm, crowded,
17. Marasmius ruforotula Singer, Sydowia 2: cylindrical to conical or wavy, obtuse to
34. 1948. (Fig. 21) subacute, yellow, inamyloid, thin-walled.
Pileus 3-7 mm diam, paraboloid to Pleurocystidia absent. Pileipellis hymeniform,
convex, umbilicate, with a small reddish brown weakly mottled, composed of Siccus-type
papilla in the umbilicus, striate, dull, dry, broom cells; main body 14-20 × 8-14 μm,
glabrous to minutely subvelutinous; disc pyriform to clavate or broadly clavate,
brownish yellow to pale rusty brown, margin sometimes irregular in outline, seldom lobed,
grayish orange (6B4-5) to light yellow (4A4). pale yellow, inamyloid, thin- to thick-walled;
Context thin, cream (3A2). Lamellae adnate to apical setulae (2-)4-8 × 1-1.5(-2) μm,
a collarium, distant (9-10) with no lamellula, 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, thin-
walled. Caulocystidia absent. Clamp
connections present in all tissues.
Habit, habitat, and known distribution:
Fig. 22. Marasmius calvus (N. Wannathes 331). 1.
Scattered to gregarious on undetermined Basidiomes 2. Basidia 3. Basidiospores 4. Cheilocystidia
dicotyledonous leaves. Florida, Guadeloupe, 5. Pleurocystidia 6. Pileipellis, Scale bar 1 = 10 mm, 2-6
Mexico, Ecuador, Papua New Guinea, Java, = 10 µm
Thailand.
Material examined: Thailand. Chiang Mai to close (14-19) with 2-3 series of lamellulae,
Province, New waterfall near 36 km marker on Hwy broad (6-12 mm), brownish orange (5C5), non-
1095, 20 Aug. 2004, N. Wannathes 257 (CMU, SFSU). marginate, non-intervenose. Stipe 82-125 × 2-5
Discussion: The material from northern mm, central, cylindrical with subbulbous base,
Thailand fits within the concept of M. hollow, glabrous to silky, non-insititious, light
ruforotula as it occurs in the old world tropics brown to buff overall. Odor and taste not
as reported by Desjardin and Horak (1997) and distinctive.
Desjardin et al. (2000) although there are Basidiospores 8-10 × 3.5-4 µm [xm = 8.9
subtle differences. The Thai material has larger ± 0.6 × 3.9 ± 0.2 µm, Q = 2.0-2.7, Qm = 2.3, n
basidiospores with mean 10.2 × 5.8 µm versus = 25 spores, s = 1 specimen], narrowly
8.9 × 4.3 mm from Indonesia. Spore size and ellipsoid, slightly curved in profile, smooth,
other basidiome features match quite closely hyaline, inamyloid, thin-walled. Basidia 25-30
those reported for M. gordipes Sacc. & Paol., a × 5 µm, cylindrical to clavate, 4-spored.
species described from Melaka State, Malaysia. Basidioles fusoid to clavate. Cheilocystidia
The latter species, however, was characterized abundant, 8-14 × 7-10 µm, broadly clavate to
as having a reddish brown pileus and an pyriform or irregularly cylindrical, hyaline,
exceptionally long stipe, measuring 60-130 mm, inamyloid, thin-walled. Pleurocystidia
and with Siccus-type broom cells that have abundant, also scattered along lamellae edge
fewer and more blunt setulae (Pegler 1986).
among the cheilocystidia, 36-46 × 7-10 µm,
fusoid to clavate, usually attenuated and
for M. ruforotula by Wannathes et al. (2007)
strangulate at the apex, hyaline, inamyloid,
based on Thai specimens.
thin-walled. Pileipellis a hymeniform layer of
Globulares-type cells: 11-28 × 10-15 µm,
Section Globulares Kühner, Botaniste 25:
clavate to broadly clavate or pyriform, hyaline,
100. 1933 (ut Globularineae).
inamyloid, thin-walled. Pileus trama inter-
Type: Marasmius globularis Fr. (= M. wynneae
woven, dextrinoid. Lamellar trama hyphae 7-
Berk. & Broome).
20 µm diam., regular, cylindrical to inflated,
smooth, hyaline, dextrinoid, thin-walled, non-
18. Marasmius calvus Berk. & Broome, J.
Linn. Soc. Bot. 14: 34. 1873 (Figs 22, 30-1)
subparallel, cylindrical, yellow, smooth, weakly
Pileus 42-87 mm diam., convex to plano-
dextrinoid, thin-walled, non-gelatinous. Stipe
convex, sulcate to plicate, glabrous, dull,
trama hyphae 6-9(-11) µm diam., parallel,
greyish orange (5B5) to light brown (6D5)
cylindrical, hyaline, smooth, dextrinoid, thin-
overall. Context brownish orange (5C5), thin.
Lamellae free to narrowly adnexed, subdistant
240
Fungal Diversity
Habit, habitat and known distribution.

leaves, Indonesia, northern Thailand and Sri
Lanka.
Province, Doi Suthep-Pui National Park, Mokfa
m., 25 June 2005, N. Wannathes 331 (CMU, SFSU).
Discussion: Diagnostic features of
Marasmius calvus include a sulcate to plicate, Fig. 23. Marasmius albimyceliosus (N. Wannathes 422).
greyish orange to light brown pileus, subdistant 1. Basidiomes 2. Basidioles 3. Basidiospores 4.
Pileipellis, Scale bar 1 = 10 mm, 2-4 = 10 µm
(14-19), broad lamellae, a pallid, glabrous stipe,
narrowly ellipsoid basidiospores with mean 8.9 basidiole-like, clavate cells. Pleurocystidia
× 3.9 µm, numerous cheilocystidia and absent. Pileipellis a hymeniform layer of
pleurocystidia, and an absence of caulocystidia. Globulares-type cells: 15-30 × 9-18 µm,
The Thai specimen matches the material clavate to broadly clavate or pyriform, hyaline
described from Indonesia and Sri Lanka quite to light brown, inamyloid, thin-walled. Pileus
well. Absent from the description of the trama interwoven, dextrinoid. Lamellar trama
Indonesian specimen (Desjardin et al., 2000) hyphae 4-10 (-16) µm diam., regular to
are the sterile lamellae edge composed of short, interwoven, cylindrical to inflated, smooth,
broadly clavate cheilocystidia as reported hyaline, dextrinoid, thin-walled, non-gelatinous.
herein. Re-examination of Retnowati 145 Stipitipellis hyphae 4-8 µm diam., subparallel,
(SFSU) confirmed their presence. Marasmius cylindrical, brown to dark brown, smooth,
calvus is the only known member of sect. inamyloid, thick-walled (up to 2 µm), non-
Globulares from Thailand that forms distinct gelatinous. Stipe trama hyphae 7-12 µm diam.,
pleurocystidia. subparallel, cylindrical, pale yellow to yellow,
smooth, inamyloid, thin- to slightly thick-
19. Marasmius albimyceliosus Corner, Beih. walled (up to 0.5 µm), non-gelatinous.
Nova. Hedwigia. 111: 28. 1996 Caulocystidia absent. Clamp connections
(Figs 23, 30-2) present in all tissues.
Pileus 15-38 mm diam., convex when Habit, habitat and known distribution:
young, plano-convex to plano-concave and Solitary on dicotyledonous leaves, northern
subumbilicate in age, reticulate, glabrous, dull, Thailand and Singapore.
disc and striae brown, elsewhere pale cream to Material examined: Thailand. Phrae Province,
grayish brown. Context light brownish cream, Muang District, Pa Dang, Nong Cham Village, 18
August 2005, N. Wannathes 422 (CMU, SFSU).
thin. Lamellae adnate, subdistant (12-16) with
3-4 series of lamellulae, narrow (2-3 mm), Discussion: Marasmius albimyceliosus is
cream, non-marginate, strong intervenose to distinguished by a reticulate, pale cream to
greyish brown pileus with darker disc and
poroid-reticulate. Stipe 23-53 × 1-4 mm,
striae, subdistant, poroid-reticulate lamellae, a
central, cylindrical or tapering downward to a
glabrous, brown to reddish brown stipe, small
subbulbous base, hollow, glabrous, non-
insititious, apex light greyish brown, base basidiospores with mean 6.8 × 4 µm, poorly
brown to reddish brown. Odor and taste not differentiated cheilocystidia, and an absence of
distinctive. pleurocystidia and caulocystidia. The Thai
material matches quite well with the protologue
Basidiospores 6-7(-8) × (3-)3.5-4(-4.5)
based on specimens collected from Singapore
µm [xm = 6.8 ± 0.5 × 4 ± 0.3 µm, Q = 1.5 - 2.0,
(Corner, 1996). Marasmius albimyceliosus is
Qm = 1.7, n = 25 spores, s = 1 specimen],
similar to M. cohortalis Berk. described from
ellipsoid, smooth, hyaline, inamyloid, thin-
South America, but the latter differs in forming
walled. Basidia not observed. Basidioles
well differentiated, subglobose to pyriform
cylindrical. Cheilocystidia poorly differentiated,
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 campan-
ulate, 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,
Fig. 24. Marasmius pellucidus (N. Wannathes 022). 1.
wavy or undulate; surface hygropha-nous, Basidiomes 2. Basidia and basidioles 3. Basidiospores 4.
glabrous, moist to dry, dull; disc ivory, cream Cheilocystidia 5. Pileipellis 6. Caulocystidia, Scale bar 1
(4A3), pale orange white (5A2-3), or pale = 10 mm, 2-6 = 10 µm
brownish grey (6C3); margin white, ivory, buff,
pale yellowish white (4A2) or cream (4A3), in 12-30(-40) × 4.5-8(-12) µm, versiform, ranging
age entire pileus often dingy white and pellucid. from irregularly cylindrical to fusoid,
Context thin (0.5-1 mm), pliant, white. ventricose, clavate or boot-shaped, hyaline,
Lamellae adnate to shallowly adnexed, non- inamyloid, thin-walled. Pleurocystidia absent.
collariate, close to subdistant or distant (12- Pileipellis a hymeniform layer of Globulares-
20(-26)) with 3-5 series of lamellulae, narrow type cells: cells 10-25(-32) × (6-)8-18(-25) µm,
(0.5-2 mm, rarely up to 4 mm broad), subglobose to pyriform, clavate or broadly
sometimes not reaching pileus margin, often clavate, sometimes in short chains, hyaline,
anastomosing and intervenose, white to ivory, inamyloid, thin-walled; arising from a subcutis
pale yellowish white (4A2) or cream (4A3); up to 30 µm thick formed of irregularly
edges even, non-marginate. Stipe (20-)40-112 × isodiametric or puzzle-shaped cells 5-12 µm
(1-)2-3 mm, central, terete or sometimes once- diam that are tightly packed, dextrinoid, thin-
cleft, ±equal, sometimes wavy, cartilaginous or walled. Pileus trama of loosely interwoven
pliant, tough, fistulose, sometimes twisted- hyphae 2.5-8(-12) µm diam, cylindrical to
fibrous, dull, dry, apex minutely pruinose, base irregularly cylindrical, thin-walled or thick-
pruinose or furfuraceous to appressed-fibrillose, walled (up to 0.5 µm), hyaline, dextrinoid, non-
non-insititious, arising from dense creamy gelatinous. Lamellar trama regular; hyphae
white mycelium; apex white to pale yellowish 2.5-10 µm diam, cylindrical, hyaline,
white (4A2), centrally greyish orange (5B4), dextrinoid, thin-walled, non-gelatinous.
brownish yellow (5C5-6) or brownish orange Stipitipellis hyphae 5-10 µm diam, parallel,
(6C5-7), base brown (7D-E5-8), reddish brown cylindrical, smooth, non-incrusted, yellow to
(8D-E4-8), or dark brown (7-8F5-8). Odor tawny in H2O, pale yellowish green in 3%KOH,
mild, somewhat sweet, like Marasmius oreades. strongly dextrinoid, thick-walled (up to 0.5
Taste not distinctive. µm), non-gelatinous. Stipe trama hyphae 5-
Basidiospores (6-)6.5-8(-8.5) × (2.5-)3-4 16(-20) µm diam, subparallel, cylindrical,
µm [xmr = 6.9-7.3 × 3-3.6 µm, xmm = 7.1 ± 0.1 hyaline, dextrinoid thin-walled. Caulocystidia
× 3.4 ± 0.2 µm, Qmr = 1.7-2.3, Qmm = 2.1 ± 0.2, 15-60(-85) × (4-)6-14 µm, versiform, ranging
n = 20-25 spores, s = 11 specimens], subfusoid from irregularly cylindrical to fusoid,
to ellipsoid or amygdaliform, with a prominent ventricose, clavate or irregular in outline,
hilar appendix, smooth, hyaline, inamyloid, sometimes geniculate, hyaline, inamyloid to
dextrinoid, thin-walled to thick-walled (0.5-1.5
thin-walled. Basidia 16-30 × 4.5-7 µm, clavate,
µm). Clamp connections present in all tissues.
4-spored. Basidioles clavate to subfusoid.
Cheilocystidia abundant, lamellar edge sterile,
Gregarious to densely gregarious, typically in
242
Fungal Diversity
cespitose clusters on leaf mulch or on woody grey (14F3-4) or dark brown to dark greyish
debris or rotten logs of various dicotyledonous brown (7-9F3-5) overall when young,
plants, sometimes amongst bamboo leaves. becoming greyish magenta (14D3) to pale
Borneo, Java, Malaysia, New Caledonia, brownish grey (8-9D-E3) on disc and plicae in
Singapore, Sri Lanka and Thailand. age with the ridges paler to pale greyish brown
Material examined: Thailand. Chiang Mai (10E3) or pale violet brown. Context pale
Province, Doi Inthanon National Park, Mae Chem jct., greyish white to white, thin. Lamellae subfree
elev. 1600 m, N18°31.420’, E98°29.216’, 26 June 2003,
N. Wannathes 007 (CMU, SFSU); Chiang Mai Province,
to adnexed, distant (13-18) with 0-1 series of
Pathummikaram temple on Hwy 1095, 13 Jul. 2003, N. lamellulae, broad (4-10 mm), greyish white
Wannathes 022 (CMU, SFSU); Chiang Mai Province, (9B1) to grey (9C-D2), greyish magenta (14D3)
Mushroom Research Centre, 27 km marker on Hwy or brownish grey (8D2), non-marginate, non-
1095, 2 Jul. 2003, N. Wannathes 016 (CMU, SFSU);
intervenose. Stipe 85-180 × 3-7 mm, central,
Chiang Mai Province, Doi Suthep National Park, Sangra
Sabhasri Lane to Huai Kok Ma Village, elev. 1145 m, cylindrical with a narrowly clavate base,
N18°48.402’, E98°54.617’, 14 Aug. 2003, N. Wannathes twisted-fibrous,
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, pellucid-
striate pileus, very narrow and intervenose
lamellae, and cespitose, reddish brown to Fig. 25. Marasmius mokfaensis (D.E. Desjardin 7726). 1.
brown stipes. It forms large, dense clusters on Basidiomes 2. Basidia 3. Cheilocystidia 4. Pileipellis 5.
leaf mulch, woody debris or on rotten logs. Basidiospores, Scale bar 1 = 10 mm, 2-6 = 10 µm
Microscopically, diagnostic features include: a
hymeniform pileipellis of subglobose to hollow, glabrous, base tomentose, apex greyish
broadly clavate, non-setulose cells; versiform, violet (17D4-6) to greyish white, base greyish
non-setulose cheilocystidia and caulocystidia; a yellow (4B3-4) to brownish grey (6-8D2-3);
lack of pleurocystidia; relatively small, stipe context yellow in age. Odor faintly
subfusoid basidiospores; and dextrinoid tissues. raphanoid to rancid. Taste unpleasant.
Collectively, these characters indicate that M. Basidiospores 27-33 × 5-6 µm [xmr =
pellucidus belongs in sect. Globulares, where it 28.6-30.7 × 5.0-5.6 µm, xmm = 30.0 ± 1.0 × 5.3
is allied with a number of species from Africa ± 0.3 µm, Q = 4.7-6.6, Qmm = 5.7 ± 0.2, Qmr =
and South America. A redescription of M. 5.5-5.9, n = 25 spores, s = 4 specimens],
pellucidus based on examination of the type clavate to subfusoid, often curved in profile,
specimens of M. pellucidus, M. papyraceus and hyaline, inamyloid, thin- walled. Basidia 41-45
C. elegans, plus numerous newly collected × 11-12 µm, clavate, 4-spored. Basidioles
specimens throughout Southeast Asia was cylindrical to clavate. Cheilocystidia common,
published by Wannathes et al. (2004). 15-33 × (3-)5-11(-17) µm, irregularly clavate
to ventricose, hyaline, inamyloid, thin-walled.
21. Marasmius mokfaensis Wannathes, Pleurocystidia absent. Pileipellis a hymeniform
Desjardin & Lumyong, Fungal Diversity 36: layer of Globulares-type cells, 11-31(-39) × 7-
160. 2009. (Figs 25, 30-4) 15 µm, clavate to broadly clavate or pyriform,
MycoBank: MB511934 hyaline, inamyloid, thin-walled. Pileus trama
Pileus 30-90 mm diam., obtusely conical interwoven, dextrinoid. Lamellar trama hyphae
when young, convex to campanulate in age, 4-8 µm diam., interwoven, cylindrical to
dull, dry, glabrous, rugulose-plicate, hygro- inflated, smooth, hyaline, dextrinoid, thin-
phanous, blackish blue (19F4-6) to purplish walled, non-gelatinous. Stipitipellis hyphae 5-
243
15 µm diam., parallel, cylindrical, smooth, subdistant (9-13) with 0-1 series of lamellulae,
brown to dark greenish brown, inamyloid to broad (1-3 mm), pale yellow (2A3), non-
weakly dextrinoid, thin- to thick-walled (up to marginate, non-intervenose. Stipe 52-103 ×
1 µm), non-gelatinous. Stipe trama hyphae 3-8 0.5-1.5 mm, central, cylindrical with or without
µm diam., parallel, cylindrical, smooth, hyaline, a subbulbous base, hollow, glabrous, non-
dextrinoid, thin-walled, non-gelatinous. insititious, apex purplish grey (16D2), base
Caulocystidia absent. Clamp connections brown to reddish brown (E8E). Odor and taste
present in all tissues. not distinctive.
Solitary on bamboo and dicotyledonous leaves,
Northern Thailand.
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
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 Fig. 26. Marasmius purpureostriatus (N. Wannathes
Marasmius mokfaensis include: a very large, 318). 1. Basidiomes 2. Basidioles 3. Basidiospores 4.
Cheilocystidia 5. Pileipellis, Scale bar 1 = 10 mm, 2-6 =
plicate pileus 30-90 mm diam., colored 10 µm
blackish blue to purplish grey with paler ridges;
distant, broad greyish brown lamellae; a large Basidiospores (19-)21-30 × (4-)5-6(-7)
(85-180 × 3-7 mm), pale colored stipe; clavate µm [xmr = 21.3-28.2 × 4.6-6.2 µm, xmm = 23.8
basidiospores with mean 30 x 5.3 µm and mean ± 3.9 × 5.3 ±0.8 µm, Qmr = 3.6-6.2, Qmm = 4.7,
Q = 5.7; irregularly clavate to ventricose n = 25 spores, s = 8 specimens], clavate to
cheilocystidia, and absence of pleurocystidia fusoid, often curved in profile, smooth, hyaline,
and caulocystidia. This is the largest known inamyloid, thin-walled. Basidia not observed.
Marasmius in Thailand and is phenetically Basidioles clavate to cylindrical. Cheilocystidia
similar to two large African species. abundant, 10-31 × 7-15 µm, cylindrical to
Marasmius zenkeri Henn. differs in forming broadly clavate or pyriform, hyaline, inamyloid,
pilei with more lilac-violet to lilac-pink pilei, thin-walled. Pleurocystidia absent. Pileipellis a
smaller basidiospores (15.5-27 × 4.2-5.6 µm hymeniform layer of Globulares-type cells, 12-
with mean Q = 4.0), and more consistently 27 × 8-14 µm, broadly clavate to pyriform,
clavate cheilocystidia (Antonín 2007). hyaline to light brown, inamyloid, thin- to
Marasmius bekolacongoli Beeli differs in thick-walled (up to 0.5 µm). Pileus trama
forming pilei that in age have dull red to interwoven, dextrinoid. Lamellar trama hyphae
greyish red disc and sulcae with yellowish 3-10(-16) µm diam., interwoven, cylindrical to
white to lemon yellow ridges, smaller inflated, smooth, hyaline, dextrinoid, thin- to
basidiospores (17.5-26 × 3.8-5.4 µm with mean thick-walled (up to 1.5 µm), non-gelatinous.
Q = 4.8), and more consistently clavate Stipitipellis hyphae 4-11 µm diam., subparallel
cheilocystidia (Antonín 2007). to parallel, cylindrical, yellowish brown to light
22. Marasmius purpureostriatus Hongo, J. brown, smooth, weakly dextrinoid to inamyloid,
Jap. Bot. 33: 344. 1958. (Figs 26, 30-8) thin- to thick-walled (up to 1µm), non-
Pileus 13-20 mm diam., obtusely conical gelatinous. Stipe trama hyphae 3-8(-12) µm
to convex with a small rugulose umbo, sulcate diam., subparallel, cylindrical, hyaline, smooth,
to plicate, glabrous, dull, striped, disc dark dextrinoid to weakly dextrinoid, thin-walled,
violet (18F8), sulcae greyish violet (18E6), non-gelatinous. Caulocystidia absent. Clamp
greyish yellow (2B4) elsewhere. Context connections present in all tissues.
greyish violet (18E6), thin. Lamellae adnexed, Habit, habitat and known distribution:
Solitary to gregarious on dicotyledonous leaves,
244
Fungal Diversity
Japan, Malaysia, Thailand and Papua New

Guinea.
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, Fig. 27. Marasmius pseudopurpureostriatus (N.
SFSU). Wannathes 286). 1. Basidiomes 2. Basidioles 3.
Discussion: Marasmius purpureostriatus is Pileipellis 4. Cheilocystidia 5. Basidiospores, Scale bar 1
distinguished by a relatively large, plicate and = 10 mm, 2-6 = 10 µm
striped pileus with greyish violet disc and
plicae and greyish yellow ridges, subdistant, Pleurocystidia absent. Pileipellis a hymeniform
broad lamellae, a purplish grey to reddish layer of Globulares-type cells, 14-23 × 10-11
brown stipe, large clavate basidiospores with µm, clavate to broadly clavate, hyaline,
mean range of 21.3-28.2 × 4.6-6.2 µm, and an inamyloid, thin- to thick-walled (up to 1 µm).
absence of pleurocystidia and caulocystidia. Pileus trama interwoven, dextrinoid. Lamellar
The Thai material matches quite closely with trama hyphae 4-10 µm diam., interwoven,
the descriptions of specimens collected from cylindrical to inflated, hyaline, smooth,
Papua New Guinea (Desjardin & Horak, 1997) dextrinoid, thin-walled, non-gelatinous.
except that the Thai specimens formed smaller Stipitipellis hyphae (3-) 5-10 µm diam.,
basidiomes (pilei 13-20 mm diam. vs 15-50 subparallel, cylindrical, brown to light brown,
mm diam.). smooth, dextrinoid, thin- to thick-walled (up to
1 µm), non-gelatinous. Stipe trama hyphae 5-
23. Marasmius pseudopurpureostriatus 12 µm diam., subparallel, cylindrical, hyaline,
Wannathes, Desjardin & Lumyong, Fungal smooth, dextrinoid, thin-walled. Caulocystidia
Diversity 36: 162. 2009. (Figs 27, 30-36) absent. Clamp connections present in all tissues.
MycoBank: MB511935 Habit, habitat and known distribution:
Pileus 14-38 mm broad, convex to Scattered to gregarious on dicotyledonous
depressed, dull, glabrous, plicate, disc and leaves, Northern Thailand.
plicae dark purple (14F5) to dark magenta Material examined: Thailand, Chiang Rai
(13F7), striped with pale yellow (3A3) to Province, Weing Papoa District, Khun Chae National
Park, N19°4.405', E99°23.543', 963 m alt., 10 June 2005,
greyish yellow (4B4) ridges. Context greyish N. Wannathes 286 (CMU: holotype; SFSU: isotype).
yellow (4B4), thin. Lamellae adnexed, distant Discussion: Basidiomes of Marasmius
(9-11) with 0-1 series of lamellulae, narrow, pseudopurpureostriatus look like a robust form
greyish yellow (4B4), non-marginate, non- of M. purpureostriatus Hongo, and the two
intervenose. Stipe 62-80 × 1.5-3 mm, central, species may be confused in the field. M.
cylindrical to tapering upwards, hollow, purpureostriatus differs from the new species
glabrous, non-insititious, apex greyish magenta in forming smaller pilei (13-20 mm diam.) with
(14E5), base brown (7E7). Odor and taste not darker violet disc and plicae and more whitish
distinctive. ridges, has a thinner stipe (< 1.5 mm), and
Basidiospores 20-25 × 5-6.2 µm [xm = slightly longer basidiospores (21-30 µm with
22.8 ± 2.4 × 5.6 ± 0.6, Q = 3.9-4.4, Qm = 4.1, n mean length about 24 µm). Marasmius
= 4 spores, s = 1 specimen], clavate to pseudopurpureostriatus is also similar to M.
subfusoid, often curved in profile, hyaline, bekolacongoli, a species widespread in tropical
inamyloid, thin-walled. Basidia not observed. Africa, but the latter differs in forming pilei
Basidioles cylindrical to clavate. Cheilocystidia that are violaceous brown to dull red or greyish
common, 12-25 × 6-12 µm, clavate to broadly red with lemon yellow stripes, and a much
clavate, hyaline, inamyloid, thin-walled. larger stipe (50-150 x 2.5-10 mm).
245
24. Marasmius grandiviridis Wannathes, to dextrinoid, thick-walled (up to 2 µm), non-
Desjardin & Lumyong, Fungal Diversity 36: gelatinous. Stipe trama hyphae 4-12 µm diam.,
156. 2009. (Figs 28, 30-7) parallel, cylindrical, hyaline, smooth, dex-
MycoBank: MB511932 trinoid, thin-walled, non-gelatinous. Caulo-
Pileus 37-88 mm diam., conical to cystidia absent. Clamp connections present in
hemispherical when young, broadly conical all tissues.
with a slight depression in age, glabrous, dull, Habit, habitat and known distribution:
plicate, yellowish green with olive green plicae. Scattered to gregarious on dicotyledonous
Context yellowish green, thin. Lamellae leaves; Northern Thailand.
narrowly adnate, distant (10-13) with 1 series Material examined: Thailand. Chiang Mai
of lamellulae, broad (2-9 mm), yellowish green, 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 (37-
88 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
Fig. 28. Marasmius grandiviridis (N. Wannathes 152). 1. cheilocystidia, and an absence of pleurocystidia
Basidiomes 2. Basidioles 3. Cheilocystidia 4. and caulocystidia. It is similar to M. viridis
Basidiospores 5. Pileipellis, Scale bar 1 = 10 mm, 2-6 = Desjardin & E. Horak, described from Papua
10 µm New Guinea, but the latter differs in forming
non-marginate, non-intervenose. Stipe 133-180 smaller basidiomes (pilei 10-15 mm diam, stipe
× 3-7 mm, central, cylindrical, hollow, 30-50 × 1 mm) with more lamellae (12-16), a
glabrous, non-insititious, apex yellowish light dark reddish brown stipe, smaller basidiospores
brown, base brownish green to brown. Odor and (20-25 × 4-5 µm) and cheilocystidia that are
taste not distinctive. more regularly clavate (Desjardin & Horak
Basidiospores (23-)26-30 × 4-5 µm [xm = 1997).
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 25. Marasmius laticlavatus Wannathes,
subfusoid, often curve in profile, smooth, Desjardin & Lumyong, Fungal Diversity 36:
hyaline, inamyloid, thin-walled. Basidia not 158. 2009. (Figs 29, 30-5)
observed. Basidioles cylindrical to clavate. MycoBank: MB511933
Cheilocystidia abundant, lamellae edge sterile, Pileus 15-19(-33) mm diam., convex to
18-43 × 5-12 µm, irregularly cylindrical to plano-convex or plano-infundibuliform with a
clavate, lageniform with a broad mucro, rarely wrinkled umbo, dull, glabrous, sulcate to
forked, hyaline, inamyloid, thin-walled. plicate, disc brown, margin yellowish grey to
Pleurocystidia absent. Pileipellis a hymeniform greyish cream. Context greyish yellow, thin.
layer of Globulares-type cells, 15-42 × 11-15 Lamellae adnexed to adnate, distant (10-13)
µm, clavate to broadly clavate or pyriform, with 1-2 series of lamellulae, broad (2-6 mm),
hyaline to pale yellow, inamyloid, thin-walled. buff to cream or white, non-marginate, non-
Pileus trama interwoven, dextrinoid. Lamellar intervenose. Stipe 30-80 × 1-2 mm, central,
trama hyphae (3-) 5-10 µm diam., regular to cylindrical with a narrow subbulbous at base,
interwoven, cylindrical to inflated, smooth, hollow, glabrous, apex yellowish grey to buff,
hyaline, weakly dextrinoid to inamyloid, thin- base brownish orange to reddish brown. Odor
walled, non-gelatinous. Stipitipellis hyphae 4-8 and taste not distinctive.
(-10) µm diam., subparallel, cylindrical, yellow Basidiospores 26-35 × 5-6(-7) µm [xmr =
to greenish yellow, smooth, weakly dextrinoid 28.7-32 × 5.1-6 µm, xmm = 30.4 ± 1.9 × 5.6 ±
246
Fungal Diversity
0.5 µm, Q = 4.1-6.9, Qmr = 5.0-6.0, Qmm = 5.5 Discussion: The new Thai species is
± 0.5, n = 25 spores, s = 4 specimens], clavate characterized by the following features:
to subfusoid, often curved in profile, hyaline, relatively small, plicate pilei with brown disc
inamyloid, thin-walled. Basidia not observed. 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
Fig. 29. Marasmius laticlavatus (N. Wannathes 231). 1. brown disc, broader lamellae (8-10 mm), a
Basidiomes 2. Basidioles 3. Cheilocystidia 4. Pileipellis much thicker stipe (4-6 mm), smaller
5. Basidiospores, Scale bar 1 = 10 mm, 2-6 = 10 µm
basidiospores (21-30 × 5.5-6.5 µm with mean
Basidioles cylindrical to clavate. Cheilocystidia Q = 4.2), and growth on wood. Marasmius
tshopoensis Antonín differs in forming larger
abundant, 13-30 × 9-16 µm, clavate to broadly
pilei (up to 60 mm diam.), strongly intervenose
clavate, hyaline, inamyloid, thin-walled.
Pleurocystidia absent. Pileipellis a hymeniform lamellae, smaller basidiospores (19-26 × 4.8-6
µm with mean Q = 4.3), and variably-shaped
layer of Globulares-type cells, 15-33 × 9-17(-
cheilocystidia some of which may be
23) µm, broadly clavate to pyriform, hyaline,
irregularly lobed, but not consistently broadly
inamyloid, thin-walled. Pileus trama
clavate. Marasmius brunneolus (Beeli) Singer
interwoven, dextrinoid. Lamellar trama hyphae
differs in forming larger pilei (30-80 mm diam.)
3-10 µm diam., interwoven, cylindrical,
that are more evenly pigmented brown to
smooth, hyaline, weakly dextrinoid to
reddish brown or striped white, a larger stipe
dextrinoid, thin-walled, non-gelatinous. Stipiti-
pellis hyphae 3-8 µm diam., subparallel, (70-180 × 3-6 mm), and smaller basidiospores
cylindrical, smooth, yellow to light brown, (15.5-25.5 × 3.8-5.4 µm with mean Q = 4.5).
dextrinoid, thick-walled (up to 2 µm), non-
gelatinous. Stipe trama hyphae 5-10 µm diam., Section Neosessiles Singer, Mycologia 50:
parallel, cylindrical, smooth, hyaline, weakly 104. 1958.
dextrinoid, thin-walled, non-gelatinous. Caulo- Type: Marasmius neosessilis Singer.
cystidia absent. Clamp connections present in
all tissues. 26. Marasmius tenuissimus (Jungh.) Singer,
Habit, habitat and known distribution: Fl. Neotrop. Monogr. 17: 258. 1976.
Scattered to gregarious on bamboo or (Figs 31, 38-1)
≡ Agaricus tenuissimus Jungh., Verh. Batav.
dicotyledonous leaves, Northern Thailand. Genootsch. 17: 84. 1838.
= Marasmius rufescens Berk. & Broome, J. Linn.
Province, 22 km marker on Hwy 1095, N19°07.57',
Soc. Bot. 14: 41. 1873.
E98°45.65', 750 m alt., 5 July 2004, T. Y. Shin 312
= Xerotus tener Berk. & Broome, J. Linn. Soc.
(CMU, SFSU); Chiang Mai Province, Mae Rim District,
Bot. 14: 45. 1873.
Huai Tung Taow Reservoir, 23 July 2004, N.Wannathes
= Marasmius campanella Holterm., Mykol.
231(CMU: holotype; SFSU: isotype); Chiang Rai
Unters.Tropen: 105. 1898.
Province, Muang District, Pong Prabath Waterfall, 11
=Marasmius campanella var. rufescens (Berk. &
June 2005, N.Wannathes 293(CMU, SFSU); Phrae
Br.) Petch apud Petch, in & Bisby, Peradeniya Manual 6:
Province, Muang District, Cherng Thong Waterfall, 16
59. 1950.
August 2005, N. Wannathes 412 (CMU, SFSU).
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
cylindrical, obtuse to subacute, hyaline, thin-

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, thin-
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
Fig. 31. Marasmius tenuissimus (N. Wannathes 199). 1. connections present in all tissues.
Basidiomes 2. Basidium 3. Basidiospores 4. Habit, habitat and known distribution:
Cheilocystidia 5. Pileipellis 6. Caulocystidia, Scale bar 1 Gregarious to densely gregarious, typically in
= 20 mm, 2-6 = 10 µm cespitose clusters on wood, Malaysia, Northern
Thailand and South America (the latter
Pileus 7-22 mm diam., convex to plano- reported by Singer, 1976).
convex, striate to sulcate, often subreticulate, Material examined: Thailand. Chiang Mai
glabrous, dull to shiny, greyish orange (6B5) to Province, Mushroom Research Centre, 27 km marker on
golden brown (6C5) or light brown (6D5). Hwy 1095, 28 July 2003, N. Wannathes 050 (CMU,
Context cream, thin. Lamellae adnate, remote SFSU); Chiang Mai Province, Mae Taeng District, Tung
Joaw Village, N 19º 8.07’ E 98º 38.9’ alt. 1423 m., 31
(5-9) with 0-1 series of lamellulae, narrow, July 2003, N. Wannathes 065 (CMU, SFSU); same
reticulate, intervenose, cream, non-marginate. location, 30 June 2004, N. Wannathes 192 (CMU,
Stipe 0.5-4 × 0.3-1 mm, eccentric to sublateral, SFSU); Chiang Mai Province, Mae Taeng District, New
cylindrical to tapering downwards, hollow, Waterfall, on 36 km. marker of Hwy1095, 2 July 2004,
N. Wannathes 199 (CMU, SFSU); same location, 20
pubescent to velutinous, insititious, dark brown August 2004, N. Wannathes 261 (CMU, SFSU).
(7F4) overall. Odor and taste not distinctive. Discussion: Marasmius tenuissimus is
Basidiospores (7-)9-10(-12) × 4-6(-7) µm characterized by a greyish orange to golden
[xmr = 8.1-10.1 × 4.5-6.1 µm, xmm = 9.4 ± 0.8 × brown or light brown, plicate-reticulate pileus,
5.3 ± 0.6 µm, Qmr = 1.7-1.9, Qmm = 1.8 ± 0.1, n remote, reticulate and intervenose lamellae, a
= 25 spores, s = 5 specimens], ellipsoid, short, eccentric to sublateral stipe, small
smooth, hyaline, inamyloid, thin-walled. basidiospores with mean 9.4 × 5.3 µm, and
Basidia 22-40 × 8-11 µm, cylindrical to clavate, growth on woody debris. Interestingly, the ITS
4-spored. Basidioles cylindrical to clavate. data indicate that M. tenuissimus is sister to M.
Cheilocystidia of Siccus-type broom cells; leveilleanus, the only known species in sect.
main body 11-25 × 5-14 µm, cylindrical to Leveilleani, and surprisingly distant from
clavate or pyriform, hyaline, inamyloid, thin- members of sect. Sicci where its morphology
walled; apical setulae 3-8 × 1-2 µm, conical to 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.
convex when young, convex to depressed in
age, umbonate, striate to sulcate, glabrous, dull
to shiny, disc dark brown (7F8) to brown (6E7-
8), 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, non-
marginate, non-intervenose. Stipe 22-90 × 0.5-
1 mm, central, cylindrical, wiry, tough, hollow,
glabrous, insititious, brown to dark brown Fig. 32. Marasmius leveilleanus (N. Wannathes 268). 1.
Basidiomes 2. Basidia 3. Basidiospores 4. Cheilocystidia
overall; rhizomorphs sometimes present. Odor
5. Pileipellis, Scale bar 1 = 20 mm, 2-5 = 10 µm
Basidiospores (8-)9-11(-12) × (3-)4(-5) non-gelatinous. Caulocystidia absent. Clamp
µm [xmr = 9.0-10.7 × 3.9-4.7 µm, xmm = 9.6 ± connections present in all tissues.
0.7 × 4.2 ± 0.3 µm, Qmr = 2.1-2.6, Qmm = 2.3 ± Habit, habitat and known distribution:
0.2, n = 25 spores, s = 6 specimens], ellipsoid, Scattered to gregarious on wood, Africa,
smooth, hyaline, inamyloid, thin-walled. Mexico, Malaysia and Northern Thailand.
Basidia 24-28 × 6-9 µm, cylindrical to clavate, Material examined: Thailand. Chiang Mai
4-spored. Basidioles cylindrical to clavate. Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
Cheilocystidia of Siccus-type broom cells; m., 28 August 2003, N. Wannathes 137 (CMU, SFSU);
main body 12-28(-37) × 4-15 µm, cylindrical to same location, 29 June 2004, Y. S. Tan 276 (CMU,
clavate, hyaline, inamyloid, thin-walled; apical SFSU); Chiang Mai Province, Chiang Mai University, 5
August 2004, N. Wannathes 248 (CMU, SFSU); Chiang
setulae 1-5 × 1-2(-3) µm, conical to cylindrical, Mai Province, Mae Taeng District, New Waterfall, on 36
obtuse, hyaline, thin- to thick-walled. km. marker of Hwy1095, 20 August 2004, N. Wannathes
Pleurocystidia absent. Pileipellis hymeniform, 264 (CMU, SFSU); same location, 30 June 2005, N.
mottled, composed of Siccus-type broom cells; Wannathes 381 (CMU, SFSU); Chiang Mai Province,
main body 9-24(-36) × 5-11 µm, clavate to Doi Suthep-Pui National Park, Mokfa Waterfall, on Hwy
1095, N19º 6.5’ E 98º 46.3’ alt. 1014 m., 21 August
broadly clavate or pyriform, yellow to hyaline, 2004, N. Wannathes 268 (CMU, SFSU); Chiang Rai
inamyloid, thin- to thick walled; apical setulae Province, Muang District, Khun Con Waterfall, 12 June
3-6 × 1.5-3 µm, conical to cylindrical, obtuse, 2005, N. Wannathes 308, (CMU, SFSU).
brown to light brown, thick-walled. Pileus Discussion: Marasmius leveilleanus is
trama interwoven, inamyloid. Lamellar trama distinguished by a relatively large sulcate
hyphae (3-)5-12(-18) µm diam., interwoven, pileus colored brown to brownish orange, free,
cylindrical to inflated, smooth, hyaline, non-collariate lamellae, a wiry, tough, dark
inamyloid, thin-walled, non-gelatinous. Stipiti- brown, insititious stipe, Siccus-type broom
pellis hyphae 3-8(-10) µm diam., subparallel, cells, moderately sized basidiospores with
cylindrical, brown to dark brown, smooth, mean 9.6 × 4.2 µm, non-dextrinoid tissues, and
weakly dextrinoid, thin- to thick- walled, non- lignicolous habit. Macromorphological features
gelatinous. Stipe trama hyphae (3-)5-8(-10) µm suggest that the species is allied with members
diam., subparallel, cylindrical, hyaline to pale of sect. Marasmius (e.g., wiry, insititious stipe)
yellow, smooth, weakly dextrinoid, thin-walled, although the lamellae are free and not collariate,
250
Fungal Diversity
while micromorphological features suggest that µm, ellipsoideae, laeves, hyalinae, inamyloideae,
it is allied with members of sect. Sicci (e.g., tenuitunicatae. Basidia non observata. Basidiolae
clavatae. Cheilocystidia abundantia, 2 typorum: a)
Siccus-type broom cells). Because of the cellulis dispersis typi Sicci 12-23 × 5-7(-10)µm,
unusual combination of features and especially cylindratis usque clavatis, hyalinis, inamyloideis,
the free lamellae and inamyloid tissues, Singer tenuitunicatis; setulis apicalibus 5-15 × 1-2 µm,
(1964) erected the monotypic sect. Leveilleani cylindratis, obtusis uque subacutis, hyalinis usque pallide
to accommodate the species. The ITS data flavis, tenui- usque crassetunicatis; b) cheilosetis 25-57 ×
presented here support its distinction from 4-6 µm, fusoideis usque lanceolatis, subacutis usque
acutis, hyalinis, inamyloideis, tenui- usque
members of both sects. Marasmius and Sicci, crassetunicatis. Pleurosetae nullae vel rarae, 30-60 × 4-6
and suggest that it is more closely allied with µm, fusoideae usque lanceolatae, subacutae usque acutae,
M. tenuissimus of sect. Neosessiles. It should hyalinae, inamyloideae, tenui- usque crassetunicatae.
be noted that M. leveilleanus is phenetically Pileipellis hymeniformis, leniter maculosus, 3 typorum:
very similar to M. nodulocystis Pegler, a) cellulis typi Sicci 11-20 × 6-10 µm, cylindratis usque
described from Africa. A comparison of the clavatis, flavis, inamyloideis, tenui- usque crassetunicatis;
setulis apicalibus 3-7(15) × 1-2 µm, coarctatis,
descriptions of both species as reported by cylindratis usque conicis, obtusis usque subacutis,
Pegler (1977) and Antonín (2007) indicate that brunneis, tenuitunicatis; b) pileosetis 34-60 × 5-8 µm,
they differ primarily in M. nodulocystis having vulgaribus, dispersis, lanceolatis usque fusoideis,
dextrinoid tissues, and M. leveilleanus having interdum lobatis, subacutis usque acutis, hyalinis usque
an insititious stipe base and sometimes pallide flavo-brunneis, inamyloideis, crassetunicatis; c)
rhizomorphs. The stipe basal attachment of M. intermediis inter cellulas scopiformes et setas 24-39 ×7-
11 µm; paucis setularum apicalium 7-37 × 2-4 µm,
nodulocystis was not described although it has crassetunicatis. Trama pilei intertexta, dextrinoidea.
been interpreted as non-insititious because the Trama lamellae regularis usque intertexta, hyphis 4-14(-
species is accepted in sect. Sicci. 20) µm diametro, cylindratis, laevibus, hyalinis,
dextrinoideis, tenuitunicatis, haud gelatinosis.
Section Sicci Singer, Mycologia 50: 106. 1958. Stipitipellis subparallelus, hyphis 6-10(-20) µm diametro,
Type: Marasmius siccus (Schwein.) Fr. cylindratis, flavis usque laete brunneis, laevibus,
dextrinoideis, tenui- usque crassetunicatis (usque ad 1
µm), haud gelatinosis. Trama stipitis parallela, hyphis
Subsect. Siccini, ser. Spinulosi (Clémençon) (8-)14-18 µm diametro, cylindratis, hyalinis, laevibus,
Desjardin, in Antonín & Noordeloos, Lib. dextrinoideis, tenuitunicatis, haud gelatinosis.
Bot. 8: 179. 1993. Caulocystidia 2 typorum: a) cellulis typi Sicci 6-11 × 5-7
Type: Marasmius cohaerens (Pers.) µm, cylindratis vel adumbratim inaequabilibus, flavis,
Cooke & Quél. [Bas. Agaricus cohaerens inamyloideis, tenuitunicatis; 3-5 setulis apicalibus, 5-26
Pers.]. × 1-3 µm, cylindratis, subacutis usque acutis, flavis
usque laete brunneis, tenuitunicatis; b) caulosetis 38-68
= series Actinopus Singer, pro parte, Fl. Neotrop.
Monogr. 17: 236. 1976. × 6-9 µm, lanceolatis usque fusoideis, raro furcatis,
flavis, inamyloideis, crassetunicatis. Fibulae praesentes
in omnibus texturis.
28. Marasmius coarctatus Wannathes, Typus: Chiang Mai Province, Doi Suthep-Pui
Desjardin & Lumyong, sp. nov. National Park, trail opposite with the way to Huai Kok
(Figs 33, 38-3) Ma Village, solitarius in folis plantae dicotyledoneae vel
MycoBank: MB512417 ligno, 2 July 2005 N. Wannathes 385 (CMU: holotypus;
Etymology: ‘coarctatus’ (L) = close-set, crowded; SFSU: isotypus).
referring to the lamellae spacing. Pileus 10-27 mm diam., convex to plano-
Pileus 10-27 mm diametro, convexus usque convex, often subumbonate, smooth or
plano-convexus, saepe subumbonatus, laevis usque pellucid-striate, velutinous, dull, brown with a
pellucido-striatus, velutinus, hebetatus, brunneus cum hint of olive to brownish beige (6F3) or
perpaulo coloris olivacei usque brunneo-ravus vel
brunneo-aurantiacus. Contextus laete flavo-brunneus,
brownish orange (5C5). Context light yellowish
tenuis. Lamellae anguste adnatae usque annexae, brown (5D4), thin. Lamellae narrowly adnate
coarctatae (33-36) cum 3-5 seriebus lamellularum, to adnexed, crowded (33-36) with 3-5 series of
angustae, obcure flavae, haud marginatae vel lamellulae, narrow, dull yellow (3B3) with or
brunneomarginatae, haud intervenosae. Stipes 41-60 × 1- without brown edges, non-intervenose. Stipe
2 mm, centralis, cylindratus, cavus, hispidulus, haud
insititius, apice pallide flavo, basi atrobrunnea. Odor 41-60 × 1-2 mm, central, cylindrical, hollow,
saporque non propria. Basidiosporae 6-8(-10) × 2.5-3.5 hispidulous, non-insititious, apex pale yellow
251
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. Stipiti-
pellis 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. Caulo-
cystidia composed of 2 types of cells: a)
5-7 µm, cylindrical or irregular in outline,
Fig. 33. Marasmius coarctatus (N. Wannathes 385). 1. yellow, inamyloid, thin-walled; apical setulae
Basidiomes 2. Basidiole 3. Basidiospores 4a. Siccus-type
cheilocystidia 4b. Cheilosetae 5a. Siccus-type pileipellis
5-26 × 1-3 µm, ranging from 3-5 setulae per
5b. Cells transitional between broom cells and setae 5c. cell, cylindrical, subacute to acute, yellow to
5c. Pileosetae 6a. Siccus-type caulocystidia 6b. light brown, thin-walled; b) caulosetae 38-68 ×
Caulosetae, Scale bar 1 = 10 mm, 2-6 = 10 µm 6-9 µm, lanceolate to fusoid, seldom forked,
yellow, inamyloid, thick-walled. Clamp
(4A3), base dark brown (9F6). Odor and taste connections present in all tissues.
not distinctive. Habit, habitat and known distribution:
Basidiospores 6-8(-10) × 2.5-3.5 µm Solitary on dicotyledonous leaves or on wood,
[xmr = 6.4-6.8 × 3.0 µm, xmm = 6.6 ± 0.4 × 3.0 ± Northern Thailand.
0.2 µm, Qmr = 2.1, Qmm = 2.1, n = 15-25 spores, Material examined: Thailand. Chiang Mai
s = 2 specimens], ellipsoid, smooth, Province, Doi Suthep-Pui National ParkMedicianl plant
garden, 16 June 2005, N. Wannathes 315 (CMU, SFSU);
hyaline,inamyloid, thin-walled. Basidia not
Chiang Mai Province, Doi Suthep-Pui National Park,
observed. Basidioles clavate. Cheilocystidia trail opposite with the way to Huai Kok Ma Village, 2
abundant, composed of 2 types of cells: a) July 2005 N. Wannathes 385 (CMU: holotype; SFSU:
scattered Siccus-type broom cells with main isotype).
body 12-23 × 5-7(-10) µm, cylindrical to Discussion: Marasmius coarctatus is
clavate, hyaline, inamyloid, thin-walled; apical distiguished by the following features: a
setulae 5-15 × 1-2 µm, cylindrical, obtuse to medium size pileus that is smooth or pellucid-
subacute, hyaline to pale-yellow, thin- to thick- striate and coloured brown with a hint of olive;
walled; b) cheilosetae 25-57 × 4-6 µm, fusoid crowded (33-36), narrow, dull yellow lamellae;
to lanceolate, subacute to acute, hyaline, a hispidulous stipe; small basidiospores with
inamyloid, thin- to thick-walled. Pleurosetae mean 6.6 × 3.0 µm; a lack of pleurocystidia
absent or seldom present, 30-60 × 4-6 µm, and pleurosetae (rarely present); and presence
fusoid to lanceolate, subacute to acute, hyaline, of distinctive pileosetae, cheilosetae and
inamyloid, thin- to thick-walled. Pileipellis caulosetae. The new Thai species is similar to
hymeniform, weakly mottled, composed of 3 M. cohaerens var. lachnophyllus (Berk.)
types of cells: a) Siccus-type broom cells with Gilliam described from North America, but the
main body 11-20 × 6-10 µm, cylindrical to latter species differs in forming larger pilei (10-
clavate, yellow, inamyloid, thin- to thick- 60 mm diam.) that are dark reddish brown,
walled; apical setulae 3-7(15) × 1-2 µm, broader (1-2 mm) lamellae with reddish brown
crowded, cylindrical to conical, obtuse to edges, slightly larger basidiospores (6.5-9 ×
subacute, brown, thick-walled; b) pileosetae, 3.2-4.8 µm, xm = 7.6 × 4.1 µm), and brownish
common, scattered, 34-60 × 5-8 µm, lanceolate orange to reddish brown, dextrinoid setae up to
252
Fungal Diversity
20 µm diam. (Desjardin, 1989). Marasmius type broom cells; main body (9-)15-28 × 6-15
coarctatus is also similar to M. coklatus µm, cylindrical to clavate, hyaline, inamyloid,
Desjardin, Retn. & E. Horak, described from thin-walled; apical setulae 2-5(-8) × 1(-2) µm,
Indonesia, but the latter differs in forming conical to cylindrical, obtuse to subacute,
larger (15-60 mm diam.) chocolate brown pilei, yellow, thick-walled. Pleurocystidia absent.
remote to distant (10-15) lamellae 4-12 mm Pileipellis hymeniform, mottled, composed of 2
broad, and larger basidiospores in the range 10- types of cells: a) Siccus-type broom cells with
11 × 4.5-6 µm with mean 10.3 × 5.3 µm. Other main body 14-26(-41) × 6-10 µm, cylindrical to
micromorphological features are similar except clavate, seldom branched, hyaline to yellow,
for occasional dextrinoid setae; they are all inamyloid, thin- to thick-walled; apical setulae
inamyloid in the Thai species (Desjardin et al., (2-)4-8(-14) × 1-1.5 µm, crowded, cylindrical,
2000). Two additional phenetically similar obtuse to subacute, brownish yellow, thick-
species are M. nexus Desjardin & E. Horak and walled; b) pileosetae, common, scattered, 60-
M. acanthocheilus Desjardin & E. Horak, both 300 × 5-15 µm, lanceolate to fusoid, subacute
described from Papua New Guinea (Desjardin to acute, golden to golden brown, inamyloid,
and Horak, 1997), Marasmius nexus differs in thick-walled (up to 2 µm). Pileus trama hyphae
forming Globulares-type cell in the pileipellis, 3-6(-9) µm, interwoven, cylindrical to inflated,
broader hymenial setae (up to 13 µm), and smooth, hyaline, dextrinoid, thin-walled.
grows on wood. Marasmius acanthocheilus Lamellar trama hyphae 3-8(-10) µm diam.,
differs in forming a rusty orange pileus, larger regular to intervowen, cylindrical to inflated,
basidiospores (9-10.5 × 4-4.5 µm), and grows smooth, hyaline, dextrinoid, thin-walled, non-
on grass or bamboo leaves. gelatinous. Stipitipellis hyphae 3-5 µm diam,
subparallel, cylindrical, yellowish brown to
29. Marasmius trichotus Corner, Beih. Nova
brown, smooth, inamyloid to weakly dextrinoid,
Hedwigia 111: 102. 1996. (Figs 34, 38-4)
thick-walled (up to 1.5 µm), non-gelatinous.
Pileus 6-23 mm diam., broadly conical to
Stipe trama hyphae 4-7(-11) µm diam.,
convex when young, plano-convex with or
subparallel, cylindrical, hyaline, smooth,
without an umbo in age, smooth (non-striate),
dextrinoid, thin-walled, non-gelatinous.
hispidulous with erect golden hairs, dull, pale
orange (5A5) to orange (6A-B7-8), golden Caulosetae abundant, 22-213 × 4-7(-15) µm,
yellow (5B7-8) or pale brownish orange (5C4- fusoid to lanceolate, sometimes cylindrical,
5), sometimes orangish white (4-5A3) at the brown, inamyloid, thick-walled. Clamp
margin. Context yellowish white, thin. connections present in all tissues.
Lamellae adnexed to narrowly adnate,
subdistant to close (12-22) with 2-4 series of
lamellulae, narrow, yellowish white, non-
marginate, non-intervenose. Stipe 15-90 × 0.5-
1 mm, central, cylindrical, hollow, hispid with
brown hairs overall, non-insititious, apex
yellowish white (4-5A2) to pale orange (5A3-
5), base brownish orange (6C5-6), rarely to
brown (7E7-8) or reddish brown (8E-F8). Odor
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, Fig. 34. Marasmius trichotus (N. Wannathes 262). 1.
thin-walled. Basidia 14-16 × 8-9 µm, clavate, Basidiomes 2. Basidia 3. Cheilocystidia 4. Basidiospores
5a. Siccus-type pileipellis 5b. Pileosetae 6. Caulosetae,
4-spored. Basidioles fusoid to clavate. Scale bar 1 = 10 mm, 2-6 = 10 µm
Cheilocystidia abundant, composed of Siccus-
253
Habit, habitat and known distribution: intervenose. Stipe 20-45 × < 1 mm, central,
Scattered to gregarious on dicotyledonous cylindrical, hollow, velutinous to hispid, non-
leaves, Northern Thailand, Papua New Guinea insititious, apex brownish orange (6C6-8), base
and Singapore. reddish brown (8E7). Odor and taste not
Material examined: Thailand. Chiang Mai distinctive.
Basidiospores 12-14 × 3-4.5 µm [xmr =
98º 29.6’ alt. 1,703 m., 26 June 2003, N. Wannathes 006 12.9-13.6 × 3.8-4.0 µm, xmm = 13.2 ± 0.5 × 3.9
(CMU, SFSU); same location, 25 June 2004, N. ± 0.1 µm, Qmr = 3.3-3.6, Qmm = 3.4 ± 0.2, n =
Wannathes 178 (CMU, SFSU); same location, 3 July 25 spores, s = 2 specimens], fusoid to narrowly
2004, Y.S. Tan 295 and 296 (CMU, SFSU); same
location, 27 June 2005, N. Wannathes 355 (CMU,
SFSU); Chiang Mai Province, Mushroom Research walled. Basidia not observed. Basidioles fusoid
Centre, 27 km marker on Hwy 1095, 13 July 2003, N. to clavate. Cheilocystidia abundant, composed
Wannathes 024 (CMU, SFSU); same location, 27 July of Siccus-type broom cells; main body 14-21 ×
2003, N. Wannathes 037 and N. Wannathes 039(CMU, 5-13 µm, cylindrical to clavate or broadly
SFSU); same location, 28 July 2003, N. Wannathes 045
(CMU, SFSU); same location, 10 August 2003, N.
clavate, hyaline, inamyloid, thin-walled; apical
Wannathes 098 (CMU, SFSU); Chiang Mai Province, setulae 2-7 × 1 µm, cylindrical, obtuse to
Doi Suthep-Pui National Park, Mokfa Waterfall, on Hwy subacute, dark brown to black, thick-walled.
1095, N19º 6.5’.E 98º 46.3’ alt. 1014 m., 1 August 2003, Pleurocystidia absent or rare, of Siccus-type
N. Wannathes 074 (CMU, SFSU), same location, 25
broom cells like the cheilocystidia; pleurosetae
June 2005, N. Wannathes 326 (CMU, SFSU); Chiang
Mai Province, Mae Taeng District, New Waterfall, on 36 absent. Pileipellis hymeniform, mottled,
km. marker of Hwy1095, 20 August 2004, N. Wannathes composed of 2 types of cells: a) Siccus-type
262 and N. Wannathes263 (CMU, SFSU); Phrae broom cells with main body (5-)14-28 × 7-18
Province, Muang District, Suen Keun, Na Koo Haa µm, cylindrical to clavate or turbinate, hyaline
Waterfall, 15 August 2005, N. Wannathes 408 (CMU,
SFSU); Phrae Province, Muang District, Pa Dang, Nong
to yellow, inamyloid, thin- to thick-walled;
Kam Village, near Nong Kam reservoir, 18 August 2005, apical setulae (2-)3-9(-13) × 1-1.5 µm,
N. Wannathes 429 (CMU, SFSU). crowded, cylindrical, obtuse to subacute,
Discussion: Marasmius trichotus is easily brown to reddish brown, thick-walled; b)
recognized in the field by the orange to pileosetae, absent or rare, scattered, 100-115 ×
brownish orange pileus stipe, both covered 8-13 µm lanceolate to fusoid, subacute to acute,
with erect golden brown hairs. In addition, the hyaline to reddish brown, inamyloid, thick-
species has close (16-22) lamellae, narrowly walled. Pileus trama interwoven, dextrinoid.
ellipsoid basidiospores with mean 12.1 × 4.1 Lamellar trama regular; hyphae 4-9 µm diam.,
µm, and lacks hymenial setae. Thai populations cylindrical, smooth, hyaline, dextrinoid, thin-
differ from Papua New Guinea and Singapore walled, non-gelatinous. Stipitipellis hyphae 3-6
populations only in forming larger pilei (6-23 µm diam., subparallel, cylindrical, yellow to
mm diam. vs 4-12 mm diam.) that are slightly light brown, smooth, inamyloid, thick-walled
paler. All pertinent diagnostic features are (up to 1 µm), non-gelatinous. Stipe trama
indistinguishable. hyphae 4-7 µm diam., subparallel, cylindrical,
hyaline, smooth, dextrinoid, thin-walled, non-
30. Marasmius nummularius Berk. & gelatinous. Caulocystidia composed of 2 types
Broome, J. Linn. Soc. Bot. 14: 33. 1873. of cells: a) Siccus-type broom cells with main
(Figs 35, 38-5) body 8-18(-36) × 5-10(-15) µm, cylindrical to
Pileus 6-20 mm diam., broadly conical to clavate or irregular in outline, hyaline to yellow,
convex when young, plano-convex to inamyloid, thin-walled; apical setuale 2-9(-23)
depressed and often umbonate in age, pruinose ×1-2 µm, cylindrical, subacute to acute, yellow,
to subvelutinous, dull; disc wrinkled, reddish thick-walled; b) caulosetae 45-110 × 9-16 µm,
brown (8E7); margin striate, brown (6-7E6-8). fusoid to lanceolate, often apically or centrally
Context pale orange (6A3), thin. Lamellae setulose (2-4 setules per cell), hyaline to yellow,
adnexed to subadnate, subdistant (12-18) with inamyloid, thick-walled. Clamp connections
2-3 series of lamellulae, narrow, pale orange present in all tissues.
(6A3) with pinkish brown edge, non-
254
Fungal Diversity
Thai material matches nicely with Indonesian

material as described by Desjardin et al. (2000).
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, non-
intervenose. 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-)6-
7 µm [xm = 11.2 ± 0.6 × 6.1 ± 0.3 µm, Q = 1.6-
2.0, Qm = 1.8, n = 25 spores, s = 1 specimen],
walled. 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
Fig. 35. Marasmius nummularius (N. Wannathes 266). 1. 20-44 × 4-6 µm, fusoid to lanceolate, often
Basidiomes 2. Basidioles 3. Basidiospores 4. forked, subacute to acute, brown, inamyloid,
Cheilocystidia 5a. Siccus-type pileipellis 5b. Pileosetae thick-walled. Pleurosetae abundant, 20-44 × 4-
6a. Siccus-type caulocystidia 6b. Caulosetae, Scale bar 1
= 10 mm, 2-6 = 10 µm 6 µm, fusoid to lanceolate, often forked,
subacute to acute, brown, inamyloid, thick-
Habit, habitat and known distribution: walled. Pileipellis hymeniform, mottled,
Scattered to gregarious on dicotyledonous composed of 2 types of cells: a) Siccus-type
leaves, Sri Lanka, Java and Northern Thailand. broom cells with main body (9-)16-37 × 6-11
Material examined: Thailand. Chiang Mai µm, cylindrical to clavate, hyaline to yellow,
Province, Doi Suthep-Pui National Park, Mokfa inamyloid, thick-walled; apical setulae 4-20×
m., 21 August 2004, N. Wannathes 226, (CMU, SFSU);
1.5-3 µm, crowded, cylindrical to conical,
Chiang Mai Province, Doi Suthep-Pui National Park, subacute, brown to dark brown, thick-walled; b)
Medicinal Plant Garden, 3 August 2005, N. Wannathes pileosetae, common, scattered, 35-38 × 7-9 µm
396 (CMU, SFSU). lanceolate to fusoid, conical to subacute,
Discussion: Marasmius nummularius is yellow to yellowish brown, inamyloid, thick-
characterized by a reddish brown rugulose- walled. Pileus trama interwoven, inamyloid.
striate pileus, subdistant (12-18) lamellae with Lamellar trama hyphae 6-12 µm diam.,
pinkish brown edges, narrowly ellipsoid interwoven, cylindrical to inflated, smooth,
basidiospores with mean 13.2 × 3.9 µm, hyaline, inamyloid, thin-walled, non- gelatinous.
scattered pileosetae, no pleurosetae, and a Stipitipellis hyphae 6-9(-12) µm diam., parallel,
velutinous to hispid stipe covered with cylindrical, brown to light brown, smooth,
caulosetae that are often apically setulose. The 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
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, pellucido-
striatus 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 brunneo-
Fig. 36. Marasmius coklatus (S.Y. Tan.301) 1. cinereo, basi laete brunnea. Odor saporque non propria.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4. Basidiosporae 6-7 (-8) × 3.5-4 µm, ellipsoideae, laeves,
Hymenial setae 5a. Siccus-type pileipellis 5b. Pileosetae hyalinae, inamyloideae, tenuitunicatae. Basidia non
6a. Siccus-type caulocystidia 6b. Caulosetae, Scale bar 1 observata. Basidiolae fusoideae usque clavatae.
= 10 mm, 2-6 = 10 µm Cheilocystidia abundantia, 2 typorum: a) cellulis typi
Sicci 13-22 × 5-7 µm, cylindratis vel adumbratim
non-gelatinous. Stipe trama hyphae 7- 17 µm inaequabilibus, hyalinis, inamyloideis, tenui- usque
diam., parallel, cylindrical, hyaline, smooth, crassetunicatis (usque ad 0.5 µm); 2-4 setulis apicalibus,
3-16 × 1-3 µm, cylindratis usque conicis, obtusis usque
weakly dextrinoid to inamyloid, thin-walled,
subacutis, flavis, tenui- usque crassetunicatis (usque ad
non-gelatinous. Caulocystidia composed of 2 0.5 µm); b) cheilosetis 19-22 × 4-6 µm, cylindratis vel
types of cells: a) Siccus-type broom cells with adumbratim inaequabilibus, obtusis usque subacutis,
main body 18-26 × 7 µm, cylindrical or hyalinis, inamyloideis, tenui- usque crassetunicatis
irregular in outline, often lobed, yellow, (usque ad 0.5 µm). Pleurosetae vulgares, cheilosetis
similes, 25-38 × 4-6 µm, fusoideae usque ventricoso-
inamyloid, thick-walled; apical setuale 15-26 × fusoideae, raro clavatae, obtusae usque subacutae,
2-3 µm, ranging from 2-4 setulae per cell, interdum furcatae, pallide flavae, inamyloideae, tenui-
cylindrical, subacute to acute, yellow to light usque crassetunicatae. Pileipellis hymeniformis, haud
brown, thick-walled; b) caulosetae 39-52 × 6-7 maculosus, 2 typorum: a) cellulis typi Sicci 13-18 × 6-10
µm, fusoid to lanceolate, seldom lobed, yellow, µm, cylindratis usque clavatis vel late clavatis, hyalinis,
inamyloideis, tenuitunicatis; setulis apicalibus 2-5 (-23)
inamyloid, thick-walled. Clamp connections
× 1-2 (-4) µm, coarctatis, cylindratis usque conicis,
present in all tissues. obtusis usque subacutis, purpureo-brunneis, tenui-
Habit, habitat and known distribution: tunicatis; b) cellulis araneae similibus, dispersis, 16-38 ×
Scattered to gregarious on dicotyledonous 7-8 µm, cylindratis vel adumbratim inaequabilibus,
leaves, Java and Northern Thailand. flavis usque laete brunneis, inamyloideis, crassetunicatis;
Material examined: Thailand. Chiang Mai 3-6 setulis apicalibus, (4-) 30-70 × 2-3 µm, cylindratis,
Province, Doi Inthanon National Park Junction of subacutis usque acutis, flavis usque laete brunneis,
Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E inamyloideis, crassetunicatis. Trama pilei intertexta,
98º 29.6’ alt. 1,703 m., 3 July 2004, T.Y. Shin 301 dextrinoidea. Trama lamellae intertexta, hyphis 2-7 µm
(CMU, SFSU). diametro, cylindratis, laevibus, hyalinis, dextrinoideis,
Discussion: Marasmius coklatus, tenuitunicatis, haud gelatinosis. Stipitipellis subparallelus,
hyphis 4-8 µm diametro, cylindratis, flavis usque laete
described originally from material collected in brunneis, laevibus, dextrinoideis, tenui- usque
Indonesia by Desjardin et al. (2000), is crassetunicatis (usque ad 1 µm), haud gelatinosis. Trama
characterized by a dark chocolate brown, stipitis parallela, hyphis 8-13 µm diametro, cylindratis,
velutinous pileus, distant to subdistant, broad, hyalinis, laevibus, dextrinoideis, tenuitunicatis, haud
lamellae with yellowish brown edges, a brown, gelatinosis. Caulocystidia 2 typorum: a) cellulis typi
256
Fungal Diversity
Sicci 12-31 × 5-6 µm, cylindratis, hyalinis, inamyloideis, inamyloid, thick-walled; apical setulae(4-) 30-
tenui- usque crassetunicatis; 2-3 setulis apicalibus, 10-20 70 × 2-3 µm, ranging from 3-6 setulae per cell,
× 1.5-2 µm, cylindratis, obtusis usque subacutis, hyalinis
cylindrical, subacute to acute, yellow to light
usque flavis, crassetunicatis; b) caulosetis 40-50 × 4-6
µm, polymorphis, cylindratis usque clavatis vel brown, inamyloid, thick-walled. Pileus trama
adumbratim inaequabilibus, saepae furcatis, hyalinis, interwoven, dextrinoid. Lamellar trama hyphae
inamyloideis, crassetunicatis. Fibulae praesentes in 2-7 µm diam., interwoven, cylindrical, smooth,
omnibus texturis. hyaline, dextrinoid, thin-walled, non-gelatinous.
Typus: Chiang Mai Province, Doi Inthanon
Stipitipellis hyphae 4-8 µm, subparallel,
Mae Chaem, N 18º 31.6’ E 98º 29.6’ alt. 1,703 m., cylindrical, yellow to light brown,smooth,
dispersus usque gregarius in folis plantae dicotyledoneae dextrinoid, thin- to thick-walled (up to 1 µm),
vel ligno, 27 June 2005, N. Wannathes 358 (CMU: non-gelatinous. Stipe trama hyphae 8-13 µm,
holotypus; SFSU: isotypus). parallel, cylindrical, hyaline, smooth, dextrioid,
Pileus 20-42 mm diam., convex with a thin-walled, non-gelatinous. Caulocystidia
slightly depressed, rugulose disc, pellucid- composed of 2 types of cells: a) Siccus-type
striate to venose-reticulate, glabrous, dull, disc broom cells with main body 12-31 × 5-6 µm,
dark purplish brown (14F4), margin greyish cylindrical, hyaline, inamyloid, thin- to thick-
brown (7E3). Context reddish grey (12C2), thin. walled; apical setulae 10-20 × 1.5-2 µm,
Lamellae adnate, subdistant (10-12) with 3 ranging from 2-3 setulae per cell, cylindrical,
series of lamellulae, broad (3-5 mm), brownish obtuse to subacute, hyaline to yellow, thick-
orange (6C3) with dark purple edges, strong walled; b) caulosetae 40-50 × 4-6 µm,
intervenose to reticulate. Stipe 27-45 × 1-2 mm, polymorphic, cylindrical to clavate or irregular
central, cylindrical, hollow, hispid with dark in outline, often forked, hyaline, inamyloid,
purple (14F4) hairs, non-insititious, apex thick-walled. Clamp connections present in all
brownish grey (11C2), base light brown. Odor tissues.
and taste not distinctive. Habit, habitat and known distribution:
Basidiospores 6-7 (-8) × 3.5-4 µm [xm = Scattered to gregarious on dicotyledonous
6.6 ± 0.6 × 3.9 ± 0.3 µm, Q = 1.2 - 2.0, Qm = leaves or on wood, Northern Thailand.
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, 25-
38 × 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
Fig. 37 Marasmius araneocephalus (N. Wannathes358) 1.
of 2 types of cells: a) Siccus-type broom cells Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
with main body 13-18 × 6-10 µm, cylindrical Pleurocystidia 5a. Siccus-type pileipellis 5b. Spider-like
to clavate or broadly broom cells, scattered, pileosetae 6. Caulocystidia, Scale bar 1 = 10 mm, 2-6 =
with main body 16-38 × 7-8 µm, cylindrical or 10 µm
irregular in outline, yellow to light brown,
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 coklatus, but the latter species differs in
forming larger basidiospores (xm = 11.2 × 6.1
98º 29.6’ alt. 1,703 m., 27 June 2005, N. Wannathes 358 µm), a smooth, dark brown pileus, non-
(CMU: holotype; SFSU: isotype). intervenose-reticulate lamellae, and lacks the
Discussion: Marasmius araneocephalus distinctive spider-like pileipellis cells. In the
has the following diagnostic features: a venose- field M. araneocephalus look like a dark form
reticulate, greyish brown pileus with dark of M. albimyceliosus, but the latter is a member
purplish brown disc; subdistant (10-12), of sect. Globulares with distinctly different
strongly intervenose to reticulate lamellae; a micromorphological features.
light brown stipe covered with dark purple
hairs; small basidiospores with mean 6.6 × 3.9 Section Sicci, subsect. Siccini, ser.
µm; and scattered spider-like broom cells on Atrorubentes Desjardin & Horak, Biblio.
the pileus surface with setulae 30-70 µm long. Mycol. 168: 27. 1997.
Many micromorphological features of M. Type: Agaricus atrorubens Berk.,
araneocephalus are similar to those of M. London J. Bot. 1: 138. 1842.
258
Fungal Diversity
[= Marasmius atrorubens (Berk.) Berk., lamellulae, narrow, cream, non-marginate, non-

Hooker's J. Bot. Kew. Gard. Misc. 8: 137. intervenose. Stipe 35-68 × 1.5-2 mm, central,
1856]. cylindrical, hollow, hispidulose to hispid, non-
insititious, apex yellowish white, base dark
33. Marasmius auratus Wannathes, Desjardin reddish brown. Odor and taste not distinctive.
& Lumyong, sp. nov. Basidiospores 11-14 × 4-5 µm [xmr =
(Figs 39, 49-1) 11.9-12.6 × 4.0-4.3, xmm = 12.2 ± 0.4 × 4.1 ±
MycoBank: 512419 0.1 µm, Qmr = 2.8 - 3.1, Qmm = 3.0 ± 0.2, n =
Etymology: ‘auratus’ = gold; referring to the gold
color of the pileus.
25 spores, s = 3 specimens], ellipsoid, curved
Pileus 5-30 mm diametro, late conicus, saepe in profile, smooth, hyaline, inamyloid, thin-
margine erecto, laevis usque striatulatus, pruinosus, walled. Basidia not observed. Basidioles
hebetatus, disco auruto, margine aurantiaco usque flavo- clavate.
aurantiaco 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, tenui-
usque 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, Fig. 39. Marasmius auratus (N. Wannathes351) 1.
cylindratis, hyalinis, laevibus, dextrinoideis, tenuituni- Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
catis, haud gelatinosis. Caulocystidia numerosa, 22-12 × Pileipellis 5. Caulocystidia, Scale bar 1 = 20 mm, 2-5 =
5-7 µm, cylindrata usque adumbratim inaequabilia, 10 µm
hyalina, inamyloidea usque leniter dextrinoidea,
crassetunicata (usque ad 1 µm). Fibulae praesentes in
omnibus texturis. Cheilocystidia absent or simple, 22-28 × 4-6
Typus: Chiang Mai Province, Doi Inthanon µm, cylindrical to clavate or irregular in outline,
National Park Junction of Highway 1,009 and road to seldom lobed, hyaline, inamyloid, thin-walled.
Mae Chaem, N 18º 31.6’ E 98º 29.6’ alt. 1,703 m., Pleurocystidia absent. Pileipellis hymeniform,
dispersus usque gregarius in folis plantae dicotyledoneae,
raro in ligno, 25 June 2004, N. Wannathes 175 (CMU: mottled, composed of Siccus-type broom cells;
holotypus; SFSU: isotypus). main body 9-26 × 5-11 µm, clavate to broadly
Pileus 5-30 mm diam., broadly conical, clavate, turbinate or irregular in outline,
often with upturned margin, smooth to hyaline, inamyloid, thin- to thick-walled; apical
striatulate, pruinose, dull, disc golden yellow setulae 3-9 × 1-2 µm, crowded, cylindrical to
(5B7), margin orange (5A6-7) to yellowish irregular in outline, obtuse to subacute, yellow,
orange (4B6-7) or greyish yellow (4B5). thick-walled. Pileus trama interwoven, dex-
Context light orange (5A4), thin. Lamellae trinoid. Lamellar trama hyphae (2-)4-8(-12)
adnate, close (17-24) with 3-4 series of µm diam., interwoven, cylindrical to inflated,
259
smooth, hyaline, dextrinoid, thin- walled, non- Marasmius auratus is also similar to two
gelatinous. Stipitipellis hyphae 5-10(-15) µm American species that are members of this
diam., parallel, cylindrical, yellow to brown, series. Marasmius actinopus Mont. sensu
smooth, weakly dextrinoid to dextrinoid, thin- Singer from South America differs in forming
to thick-walled (up to 0.5 µm), non-gelatinous. a sulcate pileus colored ochraceus yellow to
Stipe trama hyphae (4-)6-9(-11) µm diam., dull olive brown, slightly narrower
parallel, cylindrical, hyaline, smooth, dex- basidiospores (3-4 µm diam.), and has Siccus-
trinoid, thin-walled, non-gelatinous. Caulocys- type cheilocystidia (Singer, 1976; see also
tidia numerous, simple, 22-12 × 5-7 µm, Desjardin and Horak, 1997: 27). Marasmius
cylindrical to irregular in outline, obtuse, ciliatomarginatus Desjardin, described from
hyaline, inamyloid to weakly dextrinoid, thick- North America, differs in forming a reddish
walled (up to 1 µm). Clamp connections brown pileus, larger basidiospores with mean
present in all tissues. 15.7 × 4.0 µm, and red-marginate lamellae with
Habit, habitat and known distribution: cheilocystidia having reddish orange contents
Solitary, scattered to gregarious on dicotyle- (Desjardin and Petersen, 1989).
donous leaves, or rarely on wood. Northern
Thailand. 34. Marasmius pseudopellucidus Wannathes,
Material examined: Thailand. Chiang Mai Desjardin & Lumyong, sp. nov.
Province, Doi Inthanon National Park Junction of (Figs 40, 49-3)
98º 29.6’ alt. 1,703 m., 2 August 2003, N. Wannathes
MycoBank: MB512420
076 (CMU, SFSU); same location, 25 June 2004, N. Etymology: ‘pseudo’ = false; referring to the
Wannathes 175 (CMU: holotype; SFSU: isotype); same basidiomes which look like M. pellucidus.
location, 27 June 2005, N. Wannathes 351 (CMU, Pileus 10-40 mm diametro, late conicus usque
SFSU). convexus cum disco leviter depresso et margine reflexo
ubi vetus, pellucido-striatus, pruinosus, hebetatus, primo
Discussion: Marasmius auratus is disco pallide flavo et margine albo demum ubique albus.
characterized by the following features: a Contextus albidus, tenuis. Lamellae adnatae, arctae (19-
golden yellow, smooth to striatulate pileus; 22) cum 2-3 seriebus lamellularum, angustae, albae,
close lamellae; dark reddish brown stipe; haud marginatae, intervenosae ubi vetus. Stipes 16-50 ×
slender, moderately long basidiospores with 1-2 mm, centralis, cylindratus, cavus, subvelutinus usque
velutinus, haud insititius, apice albo, basi laete
mean 12.2 × 4.1 µm; cheilocystidia that are aurantiaca. Odor suavis. Sapor non proprius.
absent or only simple and basidiole-like; Basidiosporae 10-12 × 3.5-4.5 µm, ellipsoideae, oblique
abundant, simple, cylindrical, obtuse, thick- curvae, laeves, hyalinae, inamyloideae, tenuitunicatae
walled, inamyloid caulocystidia; and growth on Basidia 33-35 × 7-8 µm, clavata, 4-spora. Basidiolae
dicotyledonous leaves. This new Thai species clavatae. Margo lamellae sterilis; cheilocystidia
is most phenetically similar to several simplicia, basidiolae similibus, 13-18 × 3.5-10 µm,
cylindrata usque clavata, raro adumbratim inaequabilia,
Malaysian species (Tan et al., 2009) that have hyalina, inamyloidea, tenuitunicata. Pleurocystidia nulla.
simple caulocystidia. Marasmius musicolor Pileipellis hymeniformis, maculosus, typi Sicci; (9-)13-
Y.S. Tan & Desjardin differs in forming fewer 23 × 7-13µm, clavatus usque late clavatus vel turbinatus,
lamellae (12-14), larger basidiospores with saepe ramosus, hyalinus, inamyloideus, tenui- usque
mean 16.1 × 3.9 µm, has distinct Siccus-type crassetunicatus; setulis apicalibus 2-12 × 1-1.5µm,
coartatis, cylindratis, obtusis usque subacutis, flavis,
cheilocystidia, and dextrinoid caulocystidia crassetunicatis. Trama pilei intertexta, dextrinoidea.
formed in clusters. Marasmius ochropoides Trama lamellae intertexta, hyphis 4-9(-14) µm diametro,
Y.S. Tan & Desjardin differs in forming a cylindratis usque inflatis, laevibus, hyalinis,
smaller pileus (5-6 mm diam.), paler stipe dextrinoideis, tenuitunicatis, haud gelatinosis.
(orangish white), smaller basidiospores with Stipitipellis parallelus, hyphis (4-)6-10 µm diametro,
cylindratis, flavis usque brunneo-flavis, laevibus,
mean 10.6 µm, well-developed Siccus-type dextrinoideis, crassetunicatis (usque ad 1 µm), haud
cheilocystidia, and dextrinoid caulocystidia. gelatinosis. Trama stipitis parallela, hyphis 6-14 µm
Marasmius iras Y.S. Tan & Desjardin differs diametro, cylindratis, hyalinis, laevibus, dextrinoideis,
in forming a brown pileus, slightly longer tenuitunicatis, haud gelatinosis. Caulocystidia numerosa,
basidiospores with mean 14.1 × 3.7 µm, well- 23-87 × 6-9 µm, cylindrata usque adumbratim
inaequabilia, raro lobata, hyalina usque pallide flava,
developed Siccus-type cheilocystidia, and leniter dextrinoidea, tenui- usque crassetunicata (usque
smaller, lobed or forked caulocystidia. ad 2 µm). Fibulae praesentes in omnibus texturis.
260
Fungal Diversity
Typus: Chiang Rai Province, Muang District, dicotyledonous leaves or woody debris,
Khun Con Waterfall, dispersus usque gregarius, saepe Northern Thailand.
caespitosus in putredine e bambusa, raro in folis plantae Material examined: Thailand. Chiang Mai
dicotyledoneae vel putridine ligneus , 12 June 2005, N. Province, Huai Nam Dang National Park, North 19º
Wannathes 305 (CMU: holotypus; SFSU: isotypus). 18.3’ E 98º 35.8’ alt. 1,538 m., 28 June 2004, N.
Pileus 10-40 mm diam., broadly conical Wannathes 186 (CMU, SFSU); Chiang Rai Province,
to convex with a slightly depressed disc and Muang District, Khun Con Waterfall, 12 June 2005, N.
reflexed margin in age, pellucid-striate, Wannathes 305 (CMU: holotype; SFSU: isotype).
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, thin-
walled. Basidia 33-35 × 7-8 µm, clavate, 4-
spored. 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 Fig. 40. Marasmius pseudopellucidus (N. Wannathes305)
hymeniform, mottled, composed of Siccus-type 1. Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
broom cells; main body (9-)13-23 × 7-13 µm, Pileipellis 5. Caulocystidia, Scale bar 1 = 20 mm, 2-5 =
10 µm
clavate to broadly clavate or turbinate, often
branched, hyaline, inamyloid, thin- to thick- Discussion: Basidiomes of Marasmius
walled; apical setulae 2-12 × 1-1.5 µm, pseudopellucidus look like M. pellucidus, and
crowded, cylindrical, obtuse to subacute, the two species may be confused in the field.
yellow, thick-walled. Pileus trama interwoven, Marasmius pellucidus is a member of sect.
dextrinoid. Lamellar trama hyphae 4-9(-14) Globulares and is characterized by having
µm diam., interwoven, cylindrical to inflated, smooth, clavate to subglobose cells forming the
smooth, hyaline, dextrinoid, thin-walled, non- pileipellis, rather than Siccus-type broom cells
gelatinous. Stipitipellis hyphae (4-)6-10 µm as in M. pseudopellucidus. The new Thai
diam., parallel, cylindrical, yellow to brownish species is most similar to two white African
yellow, smooth, dextrinoid, thick-walled (up to species. Marasmius haediniformis Singer
1 µm), non-gelatinous. Stipe trama hyphae 6- differs in forming fewer lamellae (12-18 vs 19-
14 µm diam., parallel, cylindrical, hyaline, 22), a stipe colored deep brown at the base,
smooth, dextrinoid, thin-walled, non-gelatinous. numerous Siccus-type cheilocystidia, and lacks
Caulocystidia numerous, 23-87 × 6-9 µm, caulocytidia (Antonín, 2007). Marasmius
cylindrical to irregular in outline, rarely lobed, subarborescens Singer differs in forming
hyaline to pale yellow, weakly dextrinoid, thin- smaller basidiospores (xm = 6.3 × 3 µm vs 11 ×
to thick-walled (up to 2 µm). Clamp 4.1 µm), Siccus-type cheilocystidia, and a
connections present in all tissues. chestnut brown base stipe (Pegler, 1977;
Habit, habitat and known distribution: Antonín, 2007). Marasmius pseudopellucidus
Scattered to gregarious, often in cespitose also phenetically similar to two white species
clusters on bamboo debris or seldom on
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)
convex, umbonate, smooth to striate, pruinose,
Fig. 41. Marasmius ochroleucus (N. Wannathes 299) 1.
dull, hygrophanous, disc light yellow (4A2) to Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
cream (4A3-4), margin paler. Context off-white, Pileipellis 5. Caulocystidia, Scale bar 1 = 20 mm, 2-5 =
thin. Lamellae adnexed, close to crowded (20- 10 µm
24) with 3-4 series of lamellulae, narrow,
intervenose, off-white to cream, non-marginate. thin- to thick-walled (up to 1 µm), non-
Stipe 35-58 × 1-1.2 mm, central, cylindrical, gelatinous. Stipe trama hyphae 6-8(-13) µm
hollow, hispidulous, non-insititious, apex off- diam., parallel, cylindrical, hyaline, smooth,
white to light yellow (4A4), base yellowish dextrinoid, thin-walled, non-gelatinous,
orange (4A6) to light brown. Odor sweet. Taste Caulocystidia numerous, 14-53 × 5-12 µm,
not distinctive. cylindrical to fusoid-ventricose or irregular in
Basidiospores 8-12 × 3.5-4 µm [xm = 9.7 outline, obtuse, hyaline, inamyloid, thin- to
± 1.0 × 3.9 ± 0.2 µm, Q = 2.0-3.1, Qm = 2.5, n thick-walled. Clamp connections present in all
= 25 spores, s = 1 specimen], ellipsoid, curved tissues.
in profile, smooth, hyaline, inamyloid, thin- Habit, habitat and known distribution:
walled. Basidia not observed. Basidioles fusoid Scattered to gregarious on dicotyledonous
to clavate. Cheilocystidia common, of Siccus- leaves, New Caledonia and Northern Thailand.
Material examined: Thailand. Chiang Rai
type broom cells; main body 12-27 × 4-6 µm,
Province, Muang District, Khun Con Waterfall, 12 June
cylindrical or irregular in outline, hyaline, 2005, N. Wannathes 299 (CMU, SFSU).
inamyloid, thin-walled; apical setulae 3-10 × 1- Discussion: Marasmius ochroleucus,
2 µm, cylindrical, seldom branched, obtuse to originally described from material collected in
subacute, hyaline, thin-walled. Pleurocystidia New Caledonia, is characterized by a smooth,
absent. Pileipellis hymeniform, weakly mottled, hygrophanous pileus colored light yellow to
composed of Siccus-type broom cells; main cream, close to crowded, emarginate lamellae,
body 13-20 × (3-)8-9 µm, clavate to broadly a pale yellowish orange to light brown
clavate or turbinate, hyaline to pale yellow, hispidulous stipe, numerous irregularly
inamyloid, thin- to thick-walled; apical setulae cylindrical, obtuse, thin-walled caulocystidia,
2-7 × 1 µm, crowded, cylindrical to irregular in and an absence of pleurocystidia. The Thai
outline, obtuse to subacute, yellow, thick-walled. material is nearly indistinguishable from the
Pileus trama interwoven, dextrinoid. Lamellar protologue (differing only in forming fewer
trama hyphae 3-6 µm diam., interwoven, lamellae) and the species is most phenetically
cylindrical to inflated, smooth, hyaline, similar to M. auratus (described above) and M.
dextrinoid, thin-walled, non-gelatinous. musicolor Y.S.Tan & Desjardin from Malaysia.
Stipitipellis hyphae 4-10 µm diam., parallel, Marasmius auratus differs in forming a darker,
cylindrical, yellow, smooth, weakly dextrinoid, golden yellow pileus, larger basidiospores (xm
262
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)
convex, 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 × Fig. 42. Marasmius luteolus (N. Wannathes 138) 1.
1 mm, central, cylindrical with a subbulbous Basidiomes 2. Basidiospores 3. Hymenial cystidia 4.
Pileipellis 5a. Non-setulose caulocystidia 5b. Siccus-type
base, hollow, pruinose, non-insititious, apex caulocystidia, Scale bar 1 = 20 mm, 2-5 = 10 µm
yellowish white to buff, base brown. Odor and
taste not distinctive. 15 µm diam., parallel, cylindrical, hyaline,
Basidiospores 9-12× 3.5-5 µm [xmr = 9.7- smooth, dextrinoid, thin-walled, non-gelatinous.
11 × 3.7-4.3 µm, xmm = 10.2 ± 0.5 × 4.1 ± 0.2 Caulocystidia composed of two types of cells:
µm, Qmr = 2.4-3.0, Qmm = 2.6 ± 0.2, n = 25 a) Siccus-type broom cells with main body 8-
spores, s = 6 specimens], ellipsoid, curved in 18 × 6-7 µm, scattered, uncommon, cylindrical
profile, smooth, hyaline, inamyloid, thin- or irregular in outline, hyaline; apical setulae 2-
walled. Basidia not observed. Basidioles fusoid 6 × 1 µm, cylindrical to conical, wavy in
to clavate. Cheilocystidia and pleurocystidia outline, subacute, hyaline to pale-yellow, thick-
similar, common on the sides and edges of walled; b) non-setulose cells, common, 11-60 ×
lamellae, composed of Siccus-type broom cells; 5-10 µm, polymorphic, cylindrical to clavate or
main body (9-)12-33 × (5-)7-10 µm, cylindrical irregular in outline, often lobed, obtuse, hyaline,
to clavate or broadly clavate, hyaline, inamyloid, thin- to thick-walled (up to 1 µm).
inamyloid, thin- to thick-walled; apical setulae Clamp connections present in all tissues.
3-12 × 1-2 µm, cylindrical to conical, wavy in Habit, habitat and known distribution:
outline, subacute, pale yellow to yellow, thick- Scattered to gregarious on bamboo debris or on
walled. Pileipellis hymeniform, mottled, dicotyledonous leaves. Japan (Bonin Islands),
composed of Siccus-type broom cells; main Northern Thailand.
body 10-20 × 6-10µm, clavate to broadly Material examined: Thailand. Chiang Mai
clavate, hyaline, inamyloid, thin-to thick- Province, Doi Suthep-Pui National Park, Mokfa
walled; apical setulae 3-11 × 1-2 µm, crowded, m., 28 June 2003, N. Wannathes 008 (CMU, SFSU);
cylindrical to conical, wavy in outline, same location, 28 August 2003, N. Wannathes 138(CMU,
subacute, pale yellow to yellow, thick-walled. SFSU); Chiang Mai Province, Doi Suthep-Pui National
Pileus trama interwoven, dextrinoid. Lamellar Park, Sangasabhasri Land on the way to Huai Kok Ma
trama hyphae 3-8(-10) µm diam., interwoven, Village, N 18º 48.4’ E 98º 54.6’ alt. 1,146 m., 29 August
2003, N. Wannathes 143 and N. Wannathes 144 (CMU,
cylindrical, smooth, hyaline, dextrinoid, thin- SFSU); Chiang Mai Province, Huai Nam Dang National
walled, non-gelatinous. Stipitipellis hyphae 3- Park, North 19º 18.3’ E 98º 35.8’ alt. 1,538 m., 28 June
12 µm diam., parallel, cylindrical, yellowish 2004, N. Wannathes 185 (CMU, SFSU); Chiang Rai
brown, smooth, dextrinoid, thick-walled (up to Province, Muang District, Khun Con Waterfall, 12 June
2 µm), non-gelatinous. Stipe trama hyphae 4- 2005, N. Wannathes 304 (CMU, SFSU).
263
Discussion: Marasmius luteolus is flavo-brunneis usque brunneis, crassetunicatis. Trama
characterized in forming striate to sulcate pilei pilei intertexta, fortiter dextrinoidea. Trama lamellae
intertexta, hyphis (4-)8-15(-25) µm diametro, cylindratis
colored pale cream-orange to yellowish orange usque inflatis, laevibus, hyalinis, fortiter dextrinoideis,
with bright brownish orange to golden orange tenuitunicatis, haud gelatinosis. Stipitipellis parallelus,
disc, subdistant, narrow lamellae, a relatively hyphis (3-)5-8 µm diametro, cylindratis, flavo-brunneis
short (10-35 mm), pruinose stipe covered with usque brunneis, laevibus, dextrinoideis, crassetunicatis
two types of caulocystidia (Siccus-type broom (usque ad 1 µm), haud gelatinosis. Trama stipitis
parallela, hyphis 4-8 µm diametro, cylindratis, hyalinis,
cells and non-setulose cells), medium-sized laevibus, dextrinoideis, tenuitunicatis, haud gelatinosis.
basidiospores, and growth on monocotyle- Caulocystidia 2 typorum: a) cellulis typi Sicci (5-)15-25
donous and dicotyledonous debris. A × 4-7 µm, dispersis, raris, cylindratis vel adumbratim
distinctive feature of M. luteolus are inaequabilibus, hyalinis; setulis apicalibus (2-)4-11 × 1-
pleurocystidia of Siccus-type broom cells, a 1.5 µm, cylindratis usque conicis vel undulatis, pallide
feature rare in Marasmius. The type was flavis, tenui- usque crassetunicatis; b) cellulis haud
setulosis, abundantia, 11-42 × 4-10 µm, cylindratis vel
collected on palm leaves in the Bonin Islands adumbratim inaequabilibus, raro ramosis, obtusis usque
(isotype FH!) and this is the first report of the subacutis, hyalinis, inamyloideis, tenui- usque
species outside of Japan. Marasmius luteolus is crassetunicatis (usque ad 1 µm). Fibulae praesentes in
similar to M. setulosifolius Singer, another omnibus texturis.
species with Siccus-type pleurocystidia, but the Typus: Chiang Mai Province, Doi Suthep-Pui
National Park, Mokfa Waterfall, on Hwy 1095, N19º
latter species differs in lacking simple, non- 6.5’.E 98º 46.3’ alt. 1014 m., dispersus usque gregarius
setulose caulocystidia (Singer, 1976). in folis plantae dicotyledoneae vel ligno, 1 August 2003,
N. Wannathes 067 (CMU: holotypus; SFSU: isotypus).
37. Marasmius jasminodorus Wannathes, Pileus 10-40 mm diam., broadly convex
Desjardin & Lumyong, sp. nov. to campanulate, sometimes with a small umbo,
(Figs 43, 49-4) pruinose, dull; disc rugulose, dark reddish
MycoBank: 512421 brown (7-8D6-8); margin rugulose-striate, light
Etymology: ‘jasmin’ = jasmine, ‘odorus’ = odor, brown to brownish orange (7C6-8). Context
referring to the jasmine tea odor of the basidiomes. yellowish white, thin. Lamellae narrowly
Pileus 10-40 mm diametro, late convexus usque
campanulatus, interdum umbone parvo, pruinosus,
adnexed, subdistant (12-17) with 3-4 series of
hebetatus, disco ruguloso, rubro-atrobrunneo, margine lamellulae, broad (2-5 mm), pale yellowish
rugoloso-striato, laete brunneo usque brunneo-aurantiaco. white, non-marginate or with pale brownish
Contextus flavo-albus, tenuis. Lamellae annexae, orange edges, non-intervenose. Stipe 20-60 ×
subdistantes (12-17) cum 3-4 seriebus lamellularum,
1-2 mm, central, cylindrical, tough, hollow,
latae (2-5 mm), pallide flavo-albae, haud marginatae vel
pallide brunneo-aurantiacomarginatae, haud intervenosae. glabrous to minutely velutinous, non-insititious,
Stipes 20-60 × 1-2 mm, centralis, cylindratus, tenax, base with strigose, brownish orange mycelium;
cavus, glaber usque minute velutinus, haud insititius, apex pale yellowish white (4A2), base brown
mycelio strigosum et brunneo-aurantiacum ad basim, (7E7-8) to reddish brown (8E7-8) or dark
apice pallide flavo-albo, basi brunnea usque rubro- reddish brown (8F5-8). Odor strong, sweet and
brunnea vel rubro-atrobrunnea. Odor fortis, suavis et
fragrans, iasmino sicco similis. Sapor leviter amarus. fragrant, like jasmine tea. Taste mildly bitter.
Basidiosporae 9-12(-14) × 3-4(-4.5) µm, ellipsoideae, Basidiospores 9-12(-14) × 3-4(-4.5) µm
oblique curvae, laeves, hyalinae, inamyloideae, [xmr = 9.4-11.5 × 3.1-4.1 µm, xmm = 10.1 ± 0.5
tenuitunicatae Basidia 23-25 × 5-6 µm, cylindrata usque × 3.6 ± 0.3 µm, Qmr = 2.6-3.4, Qmm = 2.8 ± 0.1,
clavata, 4-spora. Basidiolae fusoideae usque clavatae.
n = 25 spores, s = 18 specimens], ellipsoid,
Cheilocystidia vulgaria, typi Sicci; 9-19(26) × 6-10 µm,
cylindrata usque clavata, hyalina usque pallide flava, curved in profile, smooth, hyaline, inamyloid,
inamyloidea, tenui- usque crassetunicata; setulis thin-walled. Basidia 23-25 × 5-6 µm,
apicalibus (2-)4-9(-11) × 1-2 µm, cylindratis vel cylindrical to clavate, 4-spored. Basidioles
adumbratim inaequabilibus, obtusis usque subacutis, fusoid to clavate. Cheilocystidia common, of
flavis usque brunneo-flavis, crassetunicatis. Pleurocys- Siccus-type broom cells; main body 9-19(-26)
tidia nulla. Pileipellis hymeniformis, maculosus, typi
Sicci; 10-23(-29) × 6-9(-15) µm, clavatus usque late × 6-10 µm, cylindrical to clavate, hyaline to
clavatus, saepe ramosus, hyalinus, inamyloideus, tenui- pale yellow, inamyloid, thin- to thick-walled;
usque crassetunicatus; setulis apicalibus (2-)4-9(-15) × apical setulae (2-)4-9(-11) × 1-2 µm, cylin-
1-2(-3) µm, coartatis, cylindratis vel adumbratim drical or irregular in outline, obtuse to subacute,
inaequabilibus, saepe ramosis, obtusis usque subacutis,
yellow to brownish yellow, thick-walled.
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, thick-
walled. 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,
walled, non-gelatinous. Caulocystidia com-
posed 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, Fig. 43 Marasmius jasminodorus (N. Wannathes067) 1.
cylindrical or irregular in outline, seldom Basidiomes 2. Basidioles 3. Basidiospores 4.
brunched, obtuse to subacute, hyaline, Cheilocystidia 5. Pileipellis 6a. Siccus-type caulocystidia
6b. Caulocystidia, Scale bar 1 = 20 mm, 2-6 = 10 µm
inamyloid, thin- to thick-walled (up to 1 µm).
Clamp connections present in all tissues. location,25 June 2004, N. Wannathes 172 (CMU, SFSU);
Habit, habitat and known distribution: same location, 27 June 2005, N. Wannathes 365 (CMU,
Scattered to gregarious on dicotyledonous SFSU); Chiang Mai Province, Doi Inthanon National
leaves or on wood, Northern Thailand. Park Junction of Highway 1,009 and road to Mae Chaem,
Material examined: Thailand. Chiang Mai N 18º 31.6’ E 98º 29.6’ alt. 1,703 m., 3 July 2004, N.
Province, Doi Suthep-Pui National Park, Mokfa Wannathes 210 (CMU, SFSU); Chiang Mai Province,
Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014 Doi Suthep-Pui National Park, trial to 9 channel station,
m., 28 June 2003, D.E. Desjardin 7589 (CMU, SFSU); 24 July 2004, N. Wannathes 238 (CMU, SFSU); Chiang
same location, 25 July 2003, N. Wannathes 028(CMU, Rai Province, Khun Chae National Park, N 19º 4.4' E 99º
SFSU); same location, 1 August 2003, N. Wannathes 23.5' alt. 963 m., 10 June 2005, N. Wannathes 287
067 (CMU: holotype; SFSU: isotype); same location, 29 (CMU, SFSU); Chiang Rai Province, Muang District,
June 2004, Y.S. Tan 283(CMU, SFSU); same location, 2 Doo Village, Pong Prabath Waterfall, N 19º 53.2' E 100º
August 2004, N. Wannathes 219 (CMU, SFSU); Chiang 07.3' alt. 408 m., 11 Jume 2005, N. Wannathes 294
Mai Province, Mae Taeng District, “Water Conservation (CMU, SFSU);Chiang Mai Province, Doi Suthep-Pui
Area” Highway 1,095 at 22 km marker, N. 19º 7.5’ E 98º National Park, the trail opposite with the way to Huai
45.7’ alt. 724 m., 2 July 2003, D.E. Desjardin 7604 Kok Ma Village, 2 July 2005, N. Wannathes 387 (CMU,
(CMU, SFSU); same location, 4 August 2003, N. SFSU); Chiang Mai Province, Mae Taeng District, Sri
Wannathes 084 (CMU, SFSU); same location, 2 June Lanna National Park, near Mae Ngad Dam, 9 August
2004, N. Wannathes 169 (CMU, SFSU); Chiang Mai 2005 , N. Wannathes 404 (CMU, SFSU); Phrae Province,
Province, Mae Taeng District, Mushroom Research Muang District, Cherng Thong Waterfall, 16 August
Centre, 27 km marker on Hwy 1095, 9 June 2004, N. 2005, N. Wannathes 414 (CMU, SFSU).
Wannathes 151 (CMU, SFSU); Chiang Mai Province, Discussion: Distinctive features of this
Doi Inthanon National Park, At 25 km marker on new species include the following: a rugulose-
Highway 1,009, N 18º 32.5’ E 98º 33.5’ alt. 1,076 m., 10 striate pileus colored light brown to brownish
June 2004, N. Wannathes 156 (CMU, SFSU); same
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, Siccus-
type broom cells and simple cylindrical non-
setulose 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).
Fig. 44. Marasmius araucariae var. araucariae (N.
38A. Marasmius araucariae Singer var. Wannathes 300) 1. Basidiomes 2. Basidiospores 3.
Cheilocystidia 4. Pileipellis 5. Caulocystidia, Scale bar 1
araucariae, Sydowia 18: 333. 1965. (Fig. 44) = 20 mm, 2-5 = 10 µm
convex, slightly depressed in age, pruinose, clavate to broadly clavate, turbinate or irregular
dull; disc rugulose, reddish brown; margin in outline, hyaline, inamyloid, thin- to thick-
striate, brownish orange. Context off-white, walled; apical setulae 3-13 × 1-1.5 µm,
thin. Lamellae narrowly adnate, subdistant (14- crowded, cylindrical to irregular in outline,
15) with 3-4 series of lamellulae, broad (1-3 obtuse to subacute, brownish yellow, thick-
mm), cream, non-marginate, non-intervenose. walled. Pileus trama interwoven, dextrinoid.
Stipe 30-45 × 1-1.2 mm, central, cylindrical, Lamellar trama hyphae 4-10(-16) µm diam.,
hollow, subvelutinous, non-insititious, apex interwoven, cylindrical, smooth, hyaline,
off-white, base brown. Odor and taste not dextrinoid, thin-walled, non-gelatinous.
distinctive. Stipitipellis hyphae 4-8 µm diam., parallel,
Basidiospores 9-11(-12) × 3-4 µm [xm = cylindrical, yellowish brown, smooth,
10.5 ± 0.8 × 3.9 ± 0.3 µm, Q = 2.3-3.1, Qm = dextrinoid, thin- to thick-walled (up to 1 µm),
2.7, n = 25 spores, s = 1 specimen], ellipsoid, non-gelatinous. Stipe trama hyphae 6-13 µm
curved in profile, smooth, hyaline, inamyloid, diam., parallel, cylindrical, hyaline, smooth,
thin-walled. Basidia not observed. Basidioles dextrinoid, thin-walled, non-gelatinous.
fusoid to clavate. Cheilocystidia common, of Caulocystidia numerous, 24-60 × 5-8 µm,
Siccus-type broom cells; main body 11-15 × 5- polymorphic, cylindrical to clavate or irregular
7 µm, subcylindrical to clavate or irregular in in outline, obtuse, hyaline, inamyloid, thin- to
outline, seldom lobed, hyaline, inamyloid, thin- thick-walled (up to 1.5 µm); Siccus-type broom
walled; apical setulae 3-8 × 1-2 µm, cylindrical cells absent on stipe surface. Clamp
to conical, sometimes wavy in outline, obtuse connections present in all tissues.
to subacute, yellow to brownish yellow, thin- Habit, habitat and known distribution:
walled. Pleurocystidia absent. Pileipellis Scattered to gregarious, often in cespitose
hymeniform, mottled, composed of Siccus-type clusters on dicotyledonous leaves. Argentina,
broom cells; main body 12-21 × 7-12 µm, Northern Thailand, Papua New Guinea.
266
Fungal Diversity

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, basidio-
spores 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 Fig. 45. Marasmius araucariae var. siccipes (N.
Wannathes 364) 1. Basidiomes 2. Basidiospores 3.
diam.).
Cheilocystidia 4. Pleurocystidia 5. Pileipellis 6a. Siccus-
type caulocystidia 6b. Caulocystidia, Scale bar 1 = 20
38B. Marasmius araucariae var. siccipes mm, 2-6 = 10µm
Desjardin, Retn. & E. Horak, Sydowia 52: 173.
2000. (Figs 45, 49-4) to conical, sometimes wavy in outline, obtuse
Pileus 10-33 mm diam., convex to to subacute, light brown to brownish yellow,
campanulate when young, expanding to plano- thin-walled. Pleurocystidia absent. Pileipellis
convex with upturned margin in age, striate to hymeniform, mottled, composed of Siccus-type
subsulcate, pruinose, dull; disc dark brown broom cells; main body (9-)14-26 × 7-10 µm,
(7F7-9F4), margin brown (6E8) to brownish clavate to broadly clavate, turbinate or irregular
orange (6C6). Context thin, greyish orange in outline, hyaline, inamyloid, thin- to thick-
(6B4) to light brown (6D5). Lamellae adnexed walled; apical setulae (2-)4-10 × 1-2 µm,
to adnate, subdistant to close (13-20) with 3-4 crowded, cylindrical to irregular in outline,
series of lamellulae, broad (2-4 mm), brownish obtuse to subacute, brown to dark brown,
orange (5C4) to greyish brown, non-marginate, thick-walled. Pileus trama interwoven,
non-intervenose. Stipe 13-82 × 1-2 mm, central, dextrinoid. Lamellar trama hyphae (3-)5-12
cylindrical, hollow, pruinose to subglabrous, µm diam., interwoven, cylindrical, smooth,
non-insititious, brown to dark brown overall. hyaline, dextrinoid, thin-walled, non-gelatinous.
Odor and taste not distinctive. Stipitipellis hyphae 4-10 µm diam., parallel,
Basidiospores (8-)9-12 × 3.5-4 µm [xmr = cylindrical, yellowish brown to brown, smooth,
9.8-11.3 × 3.3-4.0 µm, xmm = 10.4 ± 0.8 × 3.6 ± dextrinoid, thick-walled (up to 1.5 µm), non-
0.4 µm, Qmr = 2.8 - 3.1, Qmm = 2.9 ± 0.2, n = gelatinous. Stipe trama hyphae 6-14 (-20) µm
25 spores, s = 3 specimens], ellipsoid, curved diam., parallel, cylindrical, hyaline, smooth,
in profile, smooth, hyaline, inamyloid, thin- dextrinoid, thin-walled, non-gelatinous.
walled. Basidia not observed. Basidioles fusoid Caulocystidia composed of two types of cells:
to clavate. Cheilocystidia common, of Siccus- a) Siccus-type broom cells with main body 3-
type broom cells; main body 11-18(-28) × 5-9 12 × 4-7 µm, scattered, uncommon, cylindrical
µm, subcylindrical to clavate or irregular in or irregular in outline, hyaline; apical setulae 4-
outline, seldom lobed, hyaline, inamyloid, thin- 11 × 1-1.5 µm, cylindrical to conical or wavy,
walled; apical setulae 3-8 × 1-2 µm, cylindrical pale-yellow, thin-walled; b) abundant non-
267
setulose cells, 13-40 × 4-6 µm, polymorphic, cylindrical to clavate, broadly clavate or
cylindrical to clavate or irregular in outline, irregular in outline, often lobed, hyaline,
sometimes branched, obtuse, hyaline, inamy- inamyloid, thin-walled; apical setulae 3-10 × 1-
loid, thin- to thick-walled (up to 1 µm). Clamp 1.5 µm, cylindrical, obtuse to subacute, pale
connections present in all tissues. yellow to yellow, thick-walled. Pleurocystidia
Habit, habitat and known distribution: absent. Pileipellis hymeniform, weakly mottled
Scattered to solitary on dicotyledonous leaves, to non-mottled, composed of Siccus-type
Java, Malaysia and Northern Thailand. broom cells; main body (9-)12-24 × 5-11µm,
Material examined: Thailand, Chiang Mai clavate to broadly clavate, often branched,
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, 2-5 × 1-1.5 µm, crowded, cylindrical, obtuse to
Doi Inthanon National Park, At 25 km marker on subacute, yellowish brown, thick-walled.
Highway 1,009, N 18º 32.5’ E 98º 33.5’ alt. 1,076 m, 25 Pileus trama interwoven, dextrinoid. Lamellar
June 2004, N. Wannathes 173 (CMU, SFSU); same trama hyphae 3-10 µm diam., interwoven,
location, 27 June 2005, N. Wannathes 364 (CMU,
SFSU).
Discussion: This variety described by
Desjardin et al. (2000) from Java differs from
the type variety in forming numerous Siccus-
type broom cells on the stipe surface amongst
the simple, cylindrical caulocystidia. The Thai
populations are micromorphologically indis-
tinguishable 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)
convex, 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 Fig. 46. Marasmius xestocephalus (N. Wannathes 344) 1.
subvelutinous, non-insititious, apex light Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
yellow (3A4), base light brown (6D6) to Pileipellis 5. Caulocystidia, Scale bar 1 = 20 mm, 2-5 =
10 µm
reddish brown (8E7). Odor and taste not
distinctive. cylindrical, smooth, hyaline, dextrinoid, thin-
Basidiospores (10-)11-14(-15) × 3-4 µm walled, non-gelatinous. Stipitipellis hyphae
[xmr = 11.8-13 × 3.5-3.8 µm, xmm = 12.5 ± 0.6 (2-)4-8 µm diam., subparallel, cylindrical,
× 3.7 ± 0.2 µm, Qmr = 3.4, Qmm = 3.4 ± 0.0, n = brown to yellowish brown, smooth, weakly
25 spores, s = 3 specimens], ellipsoid to fusoid, dextrinoid, thick-walled (up to 1 µm), non-
smooth, hyaline, inamyloid, thin-walled. gelatinous. Stipe trama hyphae 3-8(-10) µm
Basidia not observed. Basidioles fusoid to diam., subparallel, cylindrical, hyaline, smooth,
clavate. Cheilocystidia abundant, of Siccus- dextrinoid, thin-walled, non-gelatinous. Caulo-
type broom cells; main body 10-18 × 6-9 µm, cystidia numerous, usually apically forked;
268
Fungal Diversity
main body 25-54 × 6-10 µm, cylindrical to white (3A2), base reddish brown (8E7). Odor
irregular in outline, hyaline to pale yellow, and taste not distinctive.
weakly dextrinoid, thin- to thick-walled (up to Basidiospores (11-)12-14(-15) × 3-4(-5)
1 µm); apical fork 10-20 × 4-7 µm, cylindrical, µm [xmr = 12.6-13.4 × 3.4-4.2, xmm = 12.8 ± 0.5
obtuse, hyaline to pale yellow, thin- to thick- × 3.7 ± 0.4 µm, Qmr = 3.0-3.7, Qmm = 3.5 ± 0.4,
walled; Siccus-type broom cells absent on stipe n = 25 spores, s = 3 specimens], fusiod, smooth,
surface. Clamp connections present in all hyaline, inamyloid, thin-walled. Basidia not
tissues. observed. Basidioles fusoid to clavate.
Habit, habitat and known distribution: Cheilocystidia common, of Siccus-type broom
Scattered to gregarious on dicotyledonous cells; main body (8-)13-21 × 5-7(-10) µm,
leaves, Africa and Northern Thailand. cylindrical to clavate or irregular in outline,
Material examined: Thailand. Chiang Mai seldom branched, hyaline, inamyloid, thin-
Province, Mae Taeng District, “Water Conservation
Area” Highway 1,095 at 22 km marker, N. 19º 7.5’ E 98º walled; apical setulae (2-)4-9(-11) × 1-1.5(-2)
45.7’ alt. 724 m., 5 July 2004, Y.S. Tan 313 (CMU, µm, cylindrical, obtuse to subacute, yellow to
SFSU); Chiang Mai Province, Doi Suthep-Pui National brown, thick-walled. Pleurocystidia absent.
Park, Mokfa Waterfall, on Hwy 1095, N19º 6.5’.E 98º Pileipellis hymeniform, mottled to weakly
46.3’ alt. 1014 m., 7 July 2004, J.F. Kerekes 69 (CMU, mottled, composed of Siccus-type broom cells;
SFSU); Chiang Mai Province, Mae Taeng District, Mae
Sae Village, Pang Sa Ded Water Conservation Area, N main body 12-25 × 5-9 µm, cylindrical to
19º 14.6', E 98º 38.5' alt. 962 m., 26 June 2005, N. clavate or turbinate, seldom branched, hyaline,
Wannathes 344 (CMU, SFSU). inamyloid, thin-walled; apical setulae 3-8 × 1
Discussion: Diagnostic features of µm, crowded, cylindrical to irregular in outline,
Marasmius xestocephalus include: a greyish obtuse to subacute, dark brown to black, thick-
yellow to yellowish orange pileus with walled. Pileus trama interwoven, dextrinoid.
yellowish brown disc; subdistant (15-18), Lamellar trama hyphae 3-8(-10) µm diam.,
narrow, pale yellow lamellae; a pubescent to interwoven, cylindrical, smooth, hyaline,
subvelutinous stipe; fusoid basidiospores with dextrinoid, thin-walled, non-gelatinous. Stipiti-
mean 12.5 × 3.7 µm; and apically forked pellis hyphae 3-7(-10) µm diam., subparallel,
caulocystidia. The Thai material is cylindrical, yellow to brown, smooth,
indistinguishable from the protologue (Singer, inamyloid to weakly dextrinoid, thick-walled
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)
Fig. 47. Marasmius iras (N. Wannathes 276) 1.
with brown edges, non-intervenose. Stipe 17- Basidiomes 2. Basidioles 3. Basidiospores 4.
43 × 0.5-1 mm, central, cylindrical, hollow, Cheilocystidia 5. Pileipellis 6. Caulocystidia, Scale bar 1
subvelutinous, non-insititious, apex yellowish = 20 mm, 2-6 = 10 µm
269
(up to 1 µm), non-gelatinous. Stipe trama cylindratus, cavus, subvelutinus usque hispidulus, haud
hyphae 4-10(-13) µm diam., parallel, cylin- insititius, apice flavo-albo, basi rubro-atrobrunnea. Odor
saporque non propria. Basidiosporae 8-12 × 3-4 µm,
drical, hyaline, smooth, dextrinoid, thin-walled, ellipsoideae usque fusoideae, oblique curvae, laeves,
non-gelatinous. Caulocystidia numerous, 23-43 hyalinae, inamyloideae, tenuitunicatae Basidia non
× 6-10 µm, cylindrical to clavate or irregular in observata. Basidiolae fusoideae usque clavatae.
outline, rarely lobed, hyaline, inamyloid, thin- Cheilocystidia vulgaria, typi Sicci; 13-23 × 4-9 µm,
walled; Siccus-type broom cells absent on stipe subcylindrata usque clavata vel adumbratim inaequabilia,
raro ramosa, hyalina, inamyloidea, tenuitunicata; setulis
surface. Clamp connections present in all
apicalibus 2-6 × 1-1.5 µm, cylindratis usque conicis, raro
tissues. ramosis, obtusis usque subacutis, hyalinis usque flavis,
Habit, habitat and known distribution: tenuitunicatis. Pleurocystidia nulla. Pileipellis
Scattered to solitary on dicotyledonous leaves, hymeniformis, maculosus, typi Sicci; 15-23 × 6-11 µm,
Malaysia and Northern Thailand. clavatus usque late clavatus, turbinatus vel adumbratim
Material examined: Thailand. Chiang Mai inaequabilis, hyalinus, inamyloideus, tenui- usque
Province, Mae Taeng District, Tung Joaw Village, N 19º crassetunicatus; setulis apicalibus 3-5 × 1 µm, coartatis,
8.07’ E 98º 38.9’ alt. 1423 m., 30 August 2003, N. cylindratis vel adumbratim inaequabilibus, obtusis usque
Wannathes 147 (CMU, SFSU); same location, 22 subacutis, flavis usque brunneo-flavis, crassetunicatis.
August 2004, N. Wannathes 276 (CMU, SFSU); Chiang Trama pilei intertexta, dextrinoidea. Trama lamellae
Mai Province, Huai Nam Dang National Park, North 19º intertexta, hyphis (3-)9-13 µm diametro, cylindratis,
18.3’ E 98º 35.8’ alt. 1,538 m., 29 June 2005, N. laevibus, hyalinis, dextrinoideis, tenuitunicatis, haud
Wannathes 375 (CMU, SFSU). gelatinosis. Stipitipellis subparallelus, hyphis 4-6(-9) µm
Discussion: Marasmius iras, described diametro, cylindratis, brunneis usque olivaceo-brunneis,
laevibus, inamyloideis usque leniter dextrinoideis,
from specimens collected in Malaysia, is crassetunicatis (usque ad 1 µm), haud gelatinosis. Trama
characterized by a yellowish brown pileus with stipitis parallela, hyphis 4-13 µm diametro, cylindratis,
dark brown disc, subdistant, yellowish white hyalinis, laevibus, dextrinoideis, tenuitunicatis, haud
lamellae, a subvelutinous stipe, fusoid gelatinosis. Caulocystidia numerosa, 24-48 × 6-8 µm,
basidiospores, non-setulose caulocystidia, and cylindrata, adumbratim inaequabilia, hyalina,
inamyloidea, tenui- usque crassetunicata; sine cellulis
an absence of pleurocystidia and Siccus-type
type Sicci super superficiem stipitis. Fibulae praesentes
broom cells on the stipe surface. The Thai in omnibus texturis.
specimens match quite closely the Malaysian Typus: Chiang Mai Province, Doi Inthanon
type except for having slightly smaller National Park Junction of Highway 1,009 and road to
basidiospores with mean 12.8 × 3.7 µm vs. Mae Chaem, N 18º 31.6’ E 98º 29.6’ alt. 1,703 m.,
14.1 × 3.7 µm, and longer setulae of 27 June 2005, N. Wannathes 353 (CMU: holotypus;
cheilocystidia (4-11µm vs 4-7 µm long). SFSU: isotypus).
Marasmius iras is phenetically similar to M. Pileus 7-42 mm diam., convex when
xestocephalus (see above), but the latter young, expanding to plano-convex with a
species differs in forming pilei with yellow slightly depressed disc and reflexed margin in
tones and apically forked, thicker-walled age, smooth to striate, pruinose, dull, at first
caulocystidia. olive brown overall, fading through brownish
orange (5C4-5) to yellowish brown (5E4-5) in
41. Marasmius inthanonensis Wannathes, age. Context pale orange (5A3) to greyish
Desjardin & Lumyong, sp. nov. brown (5E3), thin. Lamellae adnexed to adnate,
(Figs 48, 49-7) close (12-20) with 3-4 series of lamellulae,
MycoBank: 512422 broad (3-6 mm), pale-yellow to yellowish grey
Etymology: ‘inthanon’ referring to Doi Inthanon (3B2) with light brown edges, non-intervenose.
National Park, the site where the holotype specimen was
collected. Stipe 40-102 × 1-3 mm, central, cylindrical,
Pileus 7-42 mm diametro, convexus ubi iuvenis, hollow, subvelutinous to hispidulous, non-
expansus et plano-convexus, disco leviter depresso et insititious, apex yellowish white, base dark
margine reflexo ubi vetus, laevis usque striatus, reddish brown. Odor and taste not distinctive.
pruinosus, hebetatus, primo olivaceo-brunneus demum
brunneo-aurantiacus usque flavo-brunneus. Contextus
Basidiospores 8-12 × 3-4 µm [xmr = 9.8-
pallide aurantiacus usque cinereo-brunneus, tenuis. 10.9 × 3.9-4.0, xmm = 10.3 ± 0.8 × 3.9 ± 0.1 µm,
Lamellae annexae usque adnatae, arctae (12-20) cum 3-4 Qmr = 2.5 - 2.7, Qmm = 2.6 ± 0.1, n = 25 spores,
seriebus lamellularum, latae (3-6 mm), pallide flavae s = 2 specimens], ellipsoid to fusoid, curved in
usque flavo-cinereae, laete brunneomarginatae, haud
profile, smooth, hyaline, inamyloid, thin-
intervenosae. Stipes 40-102 × 1-3 mm, centralis,
270
Fungal Diversity
walled. Basidia not observed. Basidioles fusoid merous, 24-48 × 6-8 µm, cylindrical, irregular
to clavate. Cheilocystidia common, of Siccus- in outline, hyaline, inamyloid, thin- to thick-
type broom cells; main body 13-23 × 4-9 µm, walled; Siccus-type broom cells absent on stipe
subcylindrical to clavate or irregular in outline, surface. Clamp connections present in all
seldom branched, hyaline, inamyloid, thin- tissues.
walled; apical setulae 2-6 × 1-1.5 µm, Habit, habitat and known distribution:
cylindrical to conical, seldom branched, obtuse Scattered to gregarious on dicotyledonous
to subacute, hyaline to yellow, thin- leaves, Northern Thailand.
(CMU, SFSU); same location, 27 June 2005, N.
Wannathes 353 (CMU: holotype; SFSU: isotype).
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
Fig. 48. Marasmius inthanonensis (N. Wannathes 353) 1.
Basidiomes 2. Basidiospores 3. Cheilocystidia 4. include M. auratus (No. 33), M. ochroleucus
Pileipellis 5. Caulocystidia, Scale bar 1 = 20 mm, 2-5 = (No. 35), and M. araucariae var. siccipes (No.
10 µm 38B). Marasmius auratus differs in forming a
golden pileus that lacks olive tones, slightly
walled. Pleurocystidia absent. Pileipellis larger basidiospores with mean 12.2 × 4.1 µm,
hymeniform, mottled, composed of Siccus-type only cylindrical cheilocystidia, and longer
broom cells; main body 15-23 × 6-11 µm, caulocystidia (up to 112 µm long). Marasmius
clavate, to broadly clavate, turbinate or ochroleucus differs primarily in forming a
irregular in outline, hyaline, inamyloid, thin- to paler pileus that is light yellow to cream and
thick-walled; apical setulae 3-5 × 1 µm, lacks olive tones; basidiospores size and
crowded, cylindrical to irregular in outline, cysitidia are indistinguishable. Marasmius
obtuse to subacute, yellow to brownish yellow, araucariae var. siccipes differs in lacking olive
thick-walled. Pileus trama interwoven, tones on the pileus, and forms brown lamellae
dextrinoid. Lamellar trama hyphae (3-)9-13 and two types of caulocystidia. ITS sequence
µm diam., interwoven, cylindrical, smooth, data confirm the distinction of these four
hyaline, dextrinoid, thin-walled, non-gelatinous. species.
Stipitipellis hyphae 4-6(-9) µm diam.,
subparallel, cylindrical, brown to olive brown, Section Sicci, subsect. Siccini, ser. Leonini
smooth, inamyloid to weakly dextrinoid, thick- Singer, Fl. Neotrop. Monogr. 17: 160. 1976.
walled (up to 1 µm), non-gelatinous. Stipe Type: Marasmius leoninus Berk.
trama hyphae 4-13 µm diam., parallel,
cylindrical, hyaline, smooth, dextrinoid, thin- 42. Marasmius cremeus Wannathes, Desjardin
walled, non-gelatinous. Caulocystidia nu- & 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 cylindrical to cylindrical or irregular in outline,

when young, expanding to convex, plano- rarely fork, subacute to acute, hyaline to pale
convex or applanate with or without a broad, yellow, inamyloid, thin-walled. Pleurocystidia
rugulose umbo in age; margin striate to sulcate; absent. Pileipellis hymeniform, weakly mottled
surface dull, dry, glabrous; cream (4A3) overall to mottled, composed of Siccus-type broom
when young, in age disc cream (4A3) to light cells; main body (8-)12-21 × 7-16 μm, clavate
yellow (4A2), margin white, to broadly clavate, cylindrical, subglobose,
pyriform or turbinate, sometimes irregular in
outline, hyaline, inamyloid, thin- to thick-
walled; 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 thick-
walled (> 1 μm), non-gelatinous. Stipe trama
hyphae (2.5-)4-8(-10) μm diam, parallel,
Fig. 50. Marasmius cremeus (N. Wannathes 118). 1. cylindrical, smooth, hyaline, inamyloid to weak
Pileipellis, Scale bar 1 = 20 mm, 2-4 = 10 µm dextrinoid, thin-walled. Caulocystidia absent.
discoloring pale grayish orange in splotches. Habit, habitat, and known distribution:
Context thin, white. Lamellae adnate, Scattered to gregarious, seldom solitary, on
subdistant (8-15) with 0-1 series of lamellulae, undetermined dicotyledonous leaves. Northern
non-intervenose, narrow to moderately broad Thailand.
Material examined. Thailand. Chiang Mai
(0.5-1.0 mm), light buff to cream-colored, non- Province, Doi Suthep-Pui National Park, Mok Fa
marginate. Stipe 11-30 × 0.2-0.5 mm, central, waterfall, on Hwy 1095, N19º 6.581’, E98º 46.353’,
cylindrical, terete, filiform, tough, pliant, 1014 meters, 1 Aug. 2003, N. Wannathes 069 (CMU,
glabrous, non-insititious, base with short, tufted, SFSU); Chiang Mai Province, Mae Taeng District,
white to cream (4A3) basal tomentum; “Water Conservation Area” Highway 1,095 at 22 km
marker, N. 19º 7.5’ E 98º 45.7’ alt. 724 m., 4 August
yellowish white to dark brownish orange (7C5- 2003, N. Wannathes 085 (CMU); same location, 13 June
7) at the apex, dark brown (7-8F4-6) to nearly 2004, N. Wannathes 171 (CMU); Chiang Mai Province,
black at the base; rhizomorphs absent. Odor Doi Suthep-Pui National Park, Montathan Falls trail, 650
and taste not distinctive. meters, 4 Jul. 2002, D.E. Desjardin 7455 (CMU, SFSU);
same location, 6 Jul. 2004, Y. S. Tan 320 (CMU, SFSU);
Basidiospores (11-)12-14(-16) × 3-4(-5) Chiang Mai Province, Doi Suthep-Pui National Park,
μm [xmr = 11.7-13.5 × 3.1-4.6 μm, xmm = 12.7 ± Sangasabhasri Land on the way to Huai Kok Ma Village,
0.6 × 3.8 ± 0.5 μm, Q = 1.8-5.0, Qmr = 2.7-4.0, N 18º 48.4’ E 98º 54.6’ alt. 1,146 m., 2 July 2005, N.
Qmm = 3.4 ± 0.5, n = 25 spores, s = 8 Wannathes 383 (CMU, SFSU); Chiang Mai Province,
Doi Inthanon National Park, at 25 km marker on
specimens], narrowly ellipsoid to fusoid, Highway 1009, N18º 32.54’, E98º 33.51’, 1076 meters,
smooth, hyaline, inamyloid, thin-walled. 10 Jun. 2004, N. Wannathes154 (CMU, SFSU), same
Basidia 20-26 × 5-7 μm, clavate to cylindrical, location, 27 June 2005, N. Wannathes 366 (CMU,
4-spored. Basidioles clavate to subfusoid. SFSU); Chiang Mai Province, Mae Taeng District, Pa
Cheilocystidia common, of Siccus-type broom Deng Village, Pathummikaram temple, on Hwy 1095,
N19º 06' 28.8", E98º 44' 47.3", 1050 meters, 22 Aug.
cells; main body 9-19 × 6-12 μm, cylindrical to 2003, N. Wannathes 118 (CMU: holotype; SFSU:
pyriform, clavate, broadly clavate or irregular isotype); same location, 7 Aug. 2003, N. Wannathes 093
in outline, hyaline, thin-walled; apical setulae (CMU, SFSU); Chiang Mai Province, Mushroom
2-8(-11)× 0.5-1.0 μm, crowded, narrowly Research Centre, 27 km marker on Hwy 1095, 27 Jul.
273
2003, N. Wannathes 042(CMU, SFSU); same location, Pileus 17-56 mm diam., convex to plano-
10 Aug. 2003, N. Wannathes 099 (CMU, SFSU). convex or applanate, often subumbonate,
Discussion: Marasmius cremeus is rugulose, striate to sulcate, glabrous to
distinguished by the following features: a subvelutinous, dull, disc brown (7E8) to dark
relatively small (3–14 mm diam), sulcate, brown (7F7-8), margin brownish orange (5C5)
cream-colored pileus; subdistant (8–15), to yellowish brown (5E6). Context yellowish
narrow, cream-colored, non-collariate, non- white to cream, thin. Lamellae free to adnexed,
marginate lamellae; a filiform (0.2–0.5 mm distant (12-16) with 3-4 series of lamellulae,
diam), glabrous, non-insititious, dark brown broad (2-10 mm), often intervenose, cream
stipe with white to cream-colored basal (4B2) with brownish orange edges. Stipe 38-95
tomentum; clavate spores with mean range × 1-3 mm, central, cylindrical, wiry, hollow,
11.7–13.5 × 3.1–4.6 µm; no pleurocystidia and glabrous, non-insititious, apex buff, base
no caulocystidia; Siccus-type cheilocystidia and yellowish brown (5D8) to reddish brown (8E7).
pileipellis cells with pale yellow setulae; and Odor and taste not distinctive.
growth on dicotyledonous leaves. The new Basidiospores (8-)9-12(-13) × 5-6(-8)µm
Thai species is most closely allied M. similis
[xmr = 8.9-10.8 × 5.1-6.2 µm, xmm = 9.9 ± 0.7 ×
Berk. & M.A. Curtis, a species described from
5.6 ± 0.4 µm, Qmr = 1.6-2.0, Qmm = 1.8 ± 0.1, n
South Carolina, USA in 1849 and one that is
= 25 spores, s = 6 specimens], ellipsoid,
known only from the type specimen (K!;
isotype FH!). Marasmius similis is nearly
Basidia not observed. Basidioles cylindrical to
indistinguishable from M. cremeus, differing
clavate. Cheilocystidia common, of Siccus-type
only in forming a white pileus (not cream-
colored) and in growing on twigs. A study of broom cells; main body (8-)10-25 × 4-10 µm,
the type specimen of M. similis shows no cylindrical to clavate or pyriform, hyaline,
differences in micromorphology when inamyloid, thin-walled; apical setulae 4-23(-32)
compared to the Thai specimens. However, × 1-1.5 µm, cylindrical, inamyloid, subacute,
because M. similis is known from only a single brown to light brown, thin- to thick-walled.
specimen collected on twigs in a temperate Pleurocystidia absent. Pileipellis hymeniform,
forest in eastern North America, until more mottled, composed of Siccus-type broom cells;
material from that region is discovered and main body 8-19(-26) × 5-11 µm, clavate to
compared with the Thai material, we choose to broadly clavate or pyriform, often branched,
recognize the Asian organism as a distinct hyaline, inamyloid, thin- to thick walled; apical
species. We conclude that M. cremeus is what setulae 3-11(-18) × 1-2 µm, crowded,
van Overeem and van Overeem-de Haas (1922) cylindrical, subacute to acute, brown to dark
reported as M. similis from Java (cf. Desjardin brown, thick-walled. Pileus trama interwoven,
et al. 2000:190). dextrinoid to weakly dextrinoid. regular to
Marasmius cremeus is also phenetically interwoven, cylindrical, smooth, hyaline,
similar to M. bellus Berk., a cream-colored to weakly dextrinoid to dextrinoid, thin-walled,
yellow species from the neotropics, and to M. non-gelatinous. Stipitipellis hyphae 3-6 µm
haediniformis Singer, a pure white species diam., subparallel, cylindrical, light brown to
from Africa and Ecuador. The latter two brown, smooth, dextrinoid, thin- to thick-
species differ from M. cremeus in forming walled, non-gelatinous. Stipe trama hyphae 7-
larger pilei (13–37 mm diam), thicker stipes 12(-18) µm diam., subparallel, cylindrical,
(1.0–1.5 mm), and slightly smaller hyaline, smooth, dextrinoid, thin-walled, non-
basidiospores on average, ca 11.3 × 3.3 µm gelatinous. Caulocystidia absent. Clamp
(Singer 1976; Pegler 1977, 1983). connections present in all tissues.
A bipolar mating system was reported for Habit, habitat and known distribution:
M. cremeus by Wannathes et al. (2007). Scattered to solitary on dicotyledonous leaves
or on wood. Java, Malaysia, Northern Thailand
43. Marasmius hypochroides Berk. & and Sri Lanka.
Broome, J. Linn. Soc., Bot. 14: 35. 1873. Material examined: Thailand. Chiang Mai
(Figs 51, 57-2)
274
Fungal Diversity
MycoBank: MB512423
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 cinereo-
albae, 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 × 7-
9 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; 13-
22 × 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,
Fig. 51. Marasmius hypochoides (N. Wannathes 136). 1. hyphis 3-8 µm diametro, cylindratis, laevibus, hyalinis,
Basidiomes 2. Basidioles 3. Basidiospores 4. dextrinoideis, tenuitunicatis, haud gelatinosis.
Cheilocystidia 5. Pileipellis, Scale bar 1 = 20 mm, 2-5 = Stipitipellis subparallelus, hyphis 3-4 µm diametro,
10 µm cylindratis, brunneis, laevibus, inamyloideis,
crassetunicatis (usque ad 1.5 µm), haud gelatinosis.
Trama stipitis parallela, hyphis 3-7 µm diametro,
cylindratis, hyalinis, laevibus, dextrinoideis,
m., 28 August 2003, N. Wannathes 136 (CMU, SFSU);
tenuitunicatis, haud gelatinosis. Caulocystidia nulla.
same location, 14 June 2004, N. Wannathes 160 (CMU,
Fibulae praesentes in omnibus texturis.
SFSU); same location, 25 June 2005, N. Wannathes 335
Typus: Chiang Mai Province, Doi Inthanon
(CMU, SFSU); Chiang Mai Province, Mae Taeng
District, Tung Joaw Village, N 19º 8.07’ E 98º 38.9’ alt.
Mae Chaem, N 18º 31.6’ E 98º 29.6’ alt. 1,703 m.,
1423 m., 30 June 2004, N. Wannathes 190 (CMU,
SFSU); Chiang Mai Province, Mushroom Research
3 July 2004, N. Wannathes 201 (CMU: holotypus;SFSU:
Centre, 27 km marker on Hwy 1095, 28 July 2004, N.
isotypus).
Wannathes 247 (CMU); Chiang Mai Province, Mae
Taeng District, Sri Lanna National Park, near Mae Ngad Pileus 3-8 mm diam., conical, striate,
Dam, 9 August 2005, N. Wannathes 405 (CMU, SFSU). pruinose, dull, disc dark grayish brown (5F3),
Discussion: Marasmius hypochroides is margin olive (3E-F3-5). Context olive grey,
characterized by a relatively large, rugulose, thin. Lamellae free, distant (12), broad (1-2
striate to sulcate pileus colored brownish mm), pale greyish white, non-marginate, non-
orange to yellowish brown with dark brown intervenose. Stipe 28-45 × 0.3-0.5 mm, central,
disc, distant (12-16), broad (2-10 mm) lamellae, cylindrical, wiry, glabrous, non-insititious,
moderately broad basidiospores with mean 9.9 apex yellowish white, base dark ruby (12F4).
× 5.6 µm, and mean Q = 1.8, cheilocystidia Odor and taste not distinctive.
with very long setulae (4-32 µm), and an Basidiospores (14-)15-17 × 3-4 µm [x =
absence of pleurocystidia and caulocystidia. 15.8 ± 0.9 × 3.4 ± 0.4 µm, Q = 3.8-5.7, Qm =
The Thai specimens match nicely those from 4.7, n = 25 spores, s = 1 specimen], narrowly
Java reported by Desjardin et al. (2000). For a fusoid to clavate, curved in profile, smooth,
comparison with similar species, refer to hyaline, inamyloid, thin-walled. Basidia not
Desjardin et al. (2000). observed. Basidioles fusoid. Cheilocystidia
common, of Siccus-type broom cells; main
44.Marasmius makok Wannathes, Desjardin & body 13-22 × 7-9 µm, cylindrical to clavate or
Lumyong, sp. nov. (Figs 52, 57-3)
275
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 olive-
coloured 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 (16-
20), a brown to fulvous stipe (not ruby red),
grows on woody debris, and has smaller
Fig. 52. Marasmius makok (N. Wannathes 201). 1. basidiospores in the range 8.3-12.5 × 2.7-4 µm
Basidiomes 2. Basidioles 3. Basidiospores 4. Cheilocystidia
5. Pileipellis, Scale bar 1 = 20 mm, 2-5 = 10 µm
(Singer, 1976). Marasmius makok is sister to M.
bambusiniformis Singer (No. 48 below) in the
irregular in outline, hyaline, inamyloid, thin- ITS phylogenetic tree, and the latter species
walled; apical setulae 2-4 × 1 µm, cylindrical differs only in pileus shape and color
to conical, subacute, hyaline, thin-walled. (campanulate-convex with reddish brown and
Pleurocystidia absent. Pileipellis hymeniform, brownish orange tones), and a brown stipe.
not mottled, composed of Siccus-type broom
45. Marasmius cupreostipes Wannathes,
cells; main body 13-22 × 6-9 µm, cylindrical to
Desjardin & Lumyong, sp. nov. (Fig. 53)
clavate or irregular in outline, hyaline,
MycoBank: MB512424
inamyloid, thin- to thick-walled; apical setulae Etymology: ‘cupreo’ = copper, ‘stipes’ = stipe;
2-5 × 1 µm, crowded, cylindrical, subacute, referring to the copper coloured stipe of dried
brown, thick-walled. Pileus trama interwoven, basidiomes.
dextrinoid. Lamellar trama hyphae 3-8 µm Pileus 3-17 mm diametro, campanulatus, sulcatus
diam., interwoven, cylindrical, smooth, hyaline, usque plicatus, glaber, hebetatus, disco aurantiaco-
brunneo, margine laete aurantiaco-brunneo. Contextus
dextrinoid, thin-walled, non-gelatinous. creameus, tenuis. Lamellae annexae, distantes (10-12)
Stipitipellis hyphae 3-4 µm diam., subparallel, sine lamellula, latae (1-2 mm), creameae, haud
cylindrical, brown, smooth, inamyloid, thick- marginatae, haud intervenosae. Stipes 67-250 × 0.5-0.8
walled (up to 1.5 µm), non-gelatinous. Stipe mm, centralis, cylindratus, filo metallico similis, glaber,
trama hyphae 3-7 µm diam., parallel, insititius usque haud insititius, apice aurantiaco, basi
cuprea, ubique cuprea in sicco. Odor saporque non
cylindrical, hyaline, smooth, dextrinoid, thin- propria. Basidiosporae (17-)18-21(-26) × 4-5(7-) µm,
walled, non-gelatinous. Caulocystidia absent. cylindratae usque fusoideae, oblique curvae, laeves,
Clamp connections present in all tissues. hyalinae, inamyloideae, tenuitunicatae Basidia non
Habit, habitat and known distribution: observata. Basidiolae fusoideae usque clavatae.
Scattered to gregarious on dicotyledonous Cheilocystidia vulgaria, typi Sicci; 13-20 × 6-11 µm,
leaves, Northern Thailand. cylindrata usque clavata vel adumbratim inaequabilia,
plerumque ramosa, hyalina, inamyloidea, tenuitunicata;
Province, Doi Inthanon National Park Junction of setulis apicalibus 2-6 × 1 µm, cylindratis usque conicis,
Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E plerumque undulatis, subacutis, pallide flavis, tenui-
98º 29.6’ alt. 1,703 m., 3 July 2004, N. Wannathes 201 usque crassetunicatis. Pleurocystidia nulla. Pileipellis
(CMU: holotype; SFSU: isotype) hymeniformis, haud maculosus, typi Sicci; 10-18(-30) ×
6-15 µm, clavatus usque pyriformis vel adumbratim
276
Fungal Diversity
inaequabilis, saepe ramosus, hyalinus usque pallide Stipitipellis hyphae 3-5 µm diam., subparallel,
flavus, inamyloideus, tenuitunicatus; setulis apicalibus 2- cylindrical, brown, smooth, inamyloid, thin-
4 × 1-1.5 µm, coartatis, cylindratis, subacutis, brunneis
usque flavo-brunneis, tenui- usque crassetunicatis.
walled, non-gelatinous. Stipe trama hyphae 6-
Trama pilei intertexta, dextrinoidea usque leniter 10 µm diam., parallel, cylindrical, hyaline,
dextrinoidea. Trama lamellae intertexta, hyphis 5-8(-12) smooth, dextrinoid, thin-walled, non-gelatinous.
µm diametro, cylindratis usque inflatis, laevibus, Caulocystidia absent. Clamp connections
hyalinis, dextrinoidea usque leniter dextrinoidea, present in all tissues.
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, non-
marginate, 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, Fig. 53. Marasmius cupreostipes (N. Wannathes 150). 1.
Qm = 4.0, n = 25 spores, s = 1 specimen], Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
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 Material examined: Thailand. Chiang Mai
to clavate or irregular in outline, usually Province, Mushroom Research Centre, 27 km marker on
branched, hyaline, inamyloid, thin-walled; Hwy 1095, 3 June 2004, N. Wannathes 150 (CMU:
apical setulae 2-6 × 1 µm, cylindrical to conical, holotype;SFSU: isotype).
usually wavy, subacute, pale yellow, thin- to Discussion: This new species is
thick-walled. Pleurocystidia absent. Pileipellis characterized by a campanulate, plicate,
hymeniform, not mottled, composed of Siccus- orangish brown pileus, distant, cream-coloured
type broom cells; main body 10-18(-30) × 6-15 lamellae, clavate basidiospores with mean 19.4
µm, clavate to pyriform or irregular in outline, × 5.0 µm, and an absence of pleurocystidia and
often branched, hyaline to pale yellow, caulocystidia. The most distinctive feature is its
inamyloid, thin-walled; apical setulae 2-4 × 1- very long (up to 250 mm) copper-colored stipe.
1.5 µm, crowded, cylindrical, subacute, brown Marasmius aciebrunneus Corner, described
to yellowish brown, thin- to thick-walled. from Singapore, differs in forming brown-
Pileus trama interwoven, dextrinoid to weakly marginate lamellae, a fuscous brown stipe only
dextrinoid. Lamellar trama hyphae 5-8(-12) 20-30 mm long, and longer but narrower
µm diam., interwoven, cylindrical to inflated, basidiospores with mean 24.8 × 4.3 µm
smooth, hyaline, dextrinoid to weakly (Holotype E!; Corner 1996). Marasmius
dextrinoid, thin-walled, non-gelatinous. selangorensis Y.S. Tan & Desjardin, described
277
recently from Malaysia, differs in forming a irregular in outline, hyaline, thin-walled; apical
more brown to yellowish brown pileus, more setulae (2-)3-10 × 0.5-1.5(-2) μm, narrowly
numerous lamellae (12-18), a stipe only up to conical to cylindrical or irregular in outline,
43 mm long coloured brownish orange to dark subacute to acute, pale yellow, inamyloid, thick-
brown (brown when dried), and acerose apical walled. Pleurocystidia absent. Pileipellis
setulae on broom cells that are 4-15 µm long hymeniform, not mottled, composed of Siccus-
(Tan et al., 2007). Marasmius longistipitatus type broom cells; main body (7-)14-25(-30) ×
Antonín, described from Africa, differs in 5-7(-10) μm, clavate to broadly clavate,
forming paler pilei (pale greyish orange to cylindrical, pyriform or sometimes irregular in
orange white), and distinctive pleurocystidia. outline, hyaline, inamyloid, thin- to thick-
In all other characters the two species are walled; apical setulae (2-)3-10 × 0.8-1.5 μm,
similar (Antonín, 2004, 2007). cylindrical to conical or irregular in outline,
subacute to acute, pale yellow to tawny,
46. Marasmius corneri Wannathes, Desjardin inamyloid, thick-walled. Pileus trama and
& Lumyong, Mycol. Res. 111: 992. 2007. lamellar trama interwoven; hyphae (2-)4-8(-11)
(Figs 54) μm diam, cylindrical, smooth, hyaline,
≡ Marasmius incarnatus Corner, Beih. Nova
Hedwigia 111: 60. 1996., nom. illegit., non Marasmius
dextrinoid to strongly dextrinoid, thin-walled,
incarnatus Quél., Enchir. Fung.:142. 1886. non-gelatinous. Stipitipellis hyphae 5-6(-10)
Pileus 10-43 mm diam, obtusely conical μm diam, parallel, cylindrical, smooth, hyaline
to convex or broadly convex, with a shallow to yellowish brown, dextrinoid to strong
broad umbo, disc occasionally rugulose, dextrinoid, thin- to thick-walled (>1.5 μm),
margin striate to sulcate, dull, dry, minutely non-gelatinous. Stipe trama hyphae 3-7(-12)
velutinous, disc brownish orange (7C8) to light μm diam, parallel, cylindrical, smooth, hyaline,
brown (7D8), orange (6B8) or brownish yellow dextrinoid to strongly dextrinoid, thin- to thick-
(5C7-8), margin light orange (5A4), brownish walled, non-gelatinous. Caulocystidia absent.
orange (6C7), orange (5A6) or grayish orange Clamp connections present in all tissues.
(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), non-
marginate. Stipe 23-90 × 0.8-3 mm, central,
terete, cylindrical, with or without a
subbulbous base, hollow, glabrous, non-
insititious, 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
cylindrical to fusoid, often curved in profile,
Basidia not observed. Basidioles clavate to
cylindrical. Cheilocystidia common, of Siccus- Fig. 54 Marasmius corneri (N. Wannathes 066). 1.
type broom cells; main body 15-25 × 4-9 μm, Basidiomes 2. Basidioles 3. Basidiospores 4. Cheilocystidia 5.
cylindrical to clavate, broadly clavate or Pileipellis, Scale bar 1 = 20 mm, 2-5 = 10 µm
278
Fungal Diversity
Habit, habitat, and known distribution: inamyloidea, tenuitunicata; setulis apicalibus 3-9 × 1 µm,
Scattered to gregarious on undetermined cylindratis, plerumque undulatis, saepe furcatis,
inamyloideis, subacutis, hyalinis usque pallide flavis,
dicotyledonous leaves or wood, Singapore and crassetunicatis. Pleurocystidia nulla. Pileipellis
Northern Thailand. hymeniformis, leniter maculosus, typi Sicci; 10-18 × 7-
Material examined: Singapore. Gardens Jungle, 8 9(-12) µm, clavatus usque late clavatus vel pyriformis,
Aug. 1940, Corner s.n. (E: holotype). Thailand. Chiang hyalinus, inamyloideus, tenui- usque crassetunicatus;
Mai Province, Doi Suthep-Pui National Park, Mok Fa setulis apicalibus 3-5(-8) × 1-1.5 µm, coartatis,
waterfall on Hwy 1095, N19º 6.581’, E98º 46.353’, 1014 cylindratis, plerumque undulatis, subacutis, brunneis
meters, 25 Jul. 2003, N. Wannathes 026 (CMU, SFSU); usque atrobrunneis, crassetunicatis. Trama pilei
same location, 1 Aug. 2003, N. Wannathes 066 (CMU, intertexta, leniter dextrinoidea usque inamyloidea.
SFSU); same location, 13 Aug. 2003, N. Wannathes 103 Trama lamellae intertexta, hyphis 3-6(-8) µm diametro,
(CMU, SFSU); same location, 29 June 2004, Y.S. Tan cylindratis usque inflatis, laevibus, hyalinis, leniter
274 (CMU, SFSU); same location, 7 Jul. 2004, Y.S. Tan dextrinoidea usque inamyloidea, tenuitunicatis, haud
325 and N. Wannathes 218 (CMU, SFSU); same gelatinosis. Stipitipellis subparallelus, hyphis (2-)3-5(-6)
location, 21 Aug. 2004, N. Wannathes 269 (CMU, µm diametro, cylindratis, brunneis usque atrobrunneis,
SFSU). laevibus, inamyloideis usque leniter dextrinoideis,
Discussion: The name used by Corner crassetunicatis, haud gelatinosis. Trama stipitis
(1996) for this new species described recently subparallela, hyphis 3-5(-8) µm diametro, cylindratis,
from Singapore is a homonym (non Quélet hyalinis, laevibus, inamyloideis usque leniter
1886) and a new name was proposed by dextrinoideis, tenuitunicatis, haud gelatinosis.
Caulocystidia nulla. Fibulae praesentes in omnibus
Wannathes et al. (2007). Their choice of texturis.
epithet honors E.J.H. Corner who has Typus: Phrae Province, Muang District, Pa Dang,
published many new species of Marasmius Nong Kam Village, near Nong Kam reservoir, dispersus
from Malaysia. The material from northern usque gregarius in folis plantae dicotyledoneae vel ligno,
Thailand differs only slightly from the 18 August 2005, N. Wannathes 423 (CMU: holotypus;
SFSU: isotypus).
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.
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 Fig. 55. Marasmius imitarius (N. Wannathes 423). 1.
annexae, distantes (10-12), angustae, creameae, Basidiomes 2. Basidiospores 3. Cheilocystidia 4.
brunneomarginatae, haud intervenosae. Stipes 8-23(-80) Pileipellis, Scale bar 1 = 20 mm, 2-4 = 10 µm
× < 0.5 mm, centralis, cylindratus, filo metallico similis,
glaber, insititius usque haud insititius, apice albido, basi Pileus 2-15 mm diam., obtusely conical
brunnea usque atrobrunnea. Odor saporque non propria. to convex when young, expanding to convex
Basidiosporae 17-20(-21) × 4-5 µm, cylindratae usque and often slightly depressed in age, with a
fusoideae, oblique curvae, laeves, hyalinae, rugulose umbo, striate to sulcate, pruinose, dull,
inamyloideae, tenuitunicatae Basidia non observata. disc brown (6-7E5-8) to reddish brown (8E6-7),
Basidiolae cylindratae usque clavatae. Cheilocystidia
margin paler to light brown (7D5-6). Context
vulgaria, typi Sicci; 8-17(-23) × (3-)6-10(-15) µm,
cylindrata usque clavata vel pyriformia, hyalina, yellowish white, thin. Lamellae adnexed,
279
distant (10-12), narrow, cream with brown with darker brown to reddish brown disc,
edges, non-intervenose. Stipe 8-23(-80) × < 0.5 distant (10-12), cream-coloured lamellae with
mm, central, cylindrical, wiry, glabrous, brown edges, clavate basidiospores with mean
insititious to non-insititious, apex off-white, 18.6 × 4.4 µm, an absence of pleurocystidia
base brown to dark brown. Odor and taste not and caulocystidia, and growth mostly on
distinctive. woody sticks. The micromorphology of this
Basidiospores 17-20(-21) × 4-5 µm [xmr new species is very similar to that of many
= 17.8-19.2 × 4.1-4.7 µm, xmm = 18.6 ± 0.7 × other species, hence our choice of epithets.
4.4 ± 0.3 µm, Qmr = 4.1-4.4, Qmm = 4.4 ± 0.3, n Marasmius kanchingensis Y.S. Tan &
= 25 spores, s = 3 specimens], cylindrical to Desjardin, described recently from Malaysia,
fusoid, curved in profile, smooth, hyaline, differs in forming a striped pileus with more
inamyloid, thin-walled. Basidia not observed. brownish orange to greyish orange tones, more
Basidioles cylindrical to clavate. Cheilocystidia lamellae (14-16), slightly longer basidiospores
common, of Siccus-type broom cells; main with mean range 18.5-20.5 µm long, and
body 8-17(-23) × (3-)6-10(-15) µm, cylindrical growth on leaves. Marasmius mazatecus Singer,
to clavate or pyriform, hyaline, inamyloid, an orange-ferruginous to deep rusty-red species
thin-walled; apical setulae 3-9 × 1 µm, from Mexico, M. striaepileus Antonín, an
cylindrical, usually wavy, often forked, orangish brown species from Burundi, and M.
inamyloid, subacute, hyaline to pale yellow, sierraleonis Beeli, a dull yellowish to rusty
thick-walled. Pleurocystidia absent. Pileipellis brown species from Africa, differ primarily in
hymeniform, weakly mottled, composed of forming larger, more brightly pigmented pilei.
Siccus-type broom cells; main body 10-18 × 7- Marasmius bambusiniformis Singer (see below)
9(-12) µm, clavate to broadly clavate or differs in forming a brownish orange to greyish
pyriform, hyaline, inamyloid, thin- to thick- orange pileus, more lamellae (10-18), has
slightly smaller basidiospores with mean 16.2 x
walled; apical setulae 3-5(-8) × 1-1.5 µm,
3.7 µm, and grows on dicot leaves. In the ITS
crowded, cylindrical, usually wavy, subacute,
phylogenetic analyses, M. imitarius is not sister
brown to dark brown, thick-walled. Pileus
to M. bambusiniformis.
trama interwoven, weakly dextrinoid to
inamyloid. Lamellar trama hyphae 3-6(-8) µm 48. Marasmius bambusiniformis Singer, Fl.
diam., interwoven, cylindrical to inflated, Neotrop. Monogr. 17: 167. 1976.
smooth, hyaline, weakly dextrinoid to (Figs 56, 57-5)
inamyloid, thin-walled, non-gelatinous. Pileus 3-10 mm diam., obtusely conical,
Stipitipellis hyphae (2-)3-5(-6) µm diam, usually umbonate, striate to sulcate, pruinose,
subparallel, cylindrical, brown to dark brown, dull, disc reddish brown (9E8) to brownish
smooth, inamyloid to weakly dextrinoid, thick- orange (6C8), margin brown (7D8) to greyish
walled, non-gelatinous. Stipe trama hyphae 3- orange (6B4). Context thin, cream. Lamellae
5(-8) µm diam., subparallel, cylindrical, adnexed, subdistant (10-18) with 0-1 series of
hyaline, smooth, inamyloid to weakly lamellulae, narrow, cream with brown edges,
non- intervenose. Stipe 13-52 × 0.2-0.4 mm,
Caulocystidia absent. Clamp connections
central, cylindrical, wiry, glabrous, non-
present in all tissues.
insititious, apex yellowish white, base brown.
Basidiospores (15-)16-18(-19) × 3.5-4
leaves or on wood, Northern Thailand.
Material examined: Thailand. Chiang Rai µm [xmr = 15.7-17 × 3.6-4.0 µm, xmm = 16.2 ±
Province, Muang District, Khun Con Waterfall, 12 June 0.7 × 3.7 ± 0.2 µm, Qmr = 4.3-4.5, Qmm = 4.4 ±
2005, N. Wannathes 297 (CMU, SFSU); Phrae Province, 0.1, n = 25 spores, s = 3 specimens], narrowly
Muang District, Pa Dang, Nong Kam Village, near Nong fusoid, curved in profile, smooth, hyaline,
Kam reservoir, 18 August 2005, N. Wannathes 423
(CMU: holotype; SFSU: isotype) and N. Wannathes 425
inamyloid, thin-walled. Basidia not observed.
(CMU, SFSU). Basidioles fusoid to clavate. Cheilocystidia
Discussion: Marasmius imitarius is common, of Siccus-type broom cells; main
characterized by a light brown, sulcate pileus body (4-)10-18 × 6-9 µm, cylindrical to clavate,
280
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, non-
insititious 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. Section Sicci, subsect. Siccini, ser.
Pileipellis, Scale bar 1 = 20 mm, 2-4 = 10 µm Haematocephali Singer, Fl. Neotrop.
Monogr. 17: 201. 1976.
hyaline, inamyloid, thin-walled; apical setulae Type: Marasmius haematocephalus
2-5 × 1 µm, cylindrical to conical, subacute, (Mont.) Fr.
yellow, thick-walled. Pleurocystidia absent.
Pileipellis hymeniform, weakly mottled, 49. Marasmius confertus Berk. & Broome, J.
composed of Siccus-type broom cells; main Linn. Soc., Bot. 14: 34. 1873. (Figs 57-7, 58)
body 10-20 × (4-)7-13 µm, cylindrical to = Marasmius chondripes Berk. & Broome, J.
clavate or irregular in outline, seldom branched, Linn. Soc., Bot. 14: 36. 1873.
= Marasmius hemibaphus Berk. & Broome, J.
hyaline, inamyloid, thin- to thick-walled; apical Linn. Soc., Bot. 14: 39. 1873.
setulae 2-5(7-) × 1 µm, crowded, cylindrical, Pileus 18-24(-40) mm diam., convex to
subacute, brownish yellow to light brown, thick broadly convex, subumbonate, smooth to
walled. Pileus trama interwoven, dextrinoid. striatulate, glabrous, dull, disc brown to dark
Lamellar trama hyphae (3-)4-10(-15) µm diam., brown (7E-F8) or reddish brown (8E-F8),
interwoven, cylindrical, smooth, hyaline, margin brownish orange (5C4-5) to yellowish
dextrinoid, thin-walled, non-gelatinous. Stipiti- brown (5E8), hygrophanous, disc fading to
pellis hyphae (2-)4-7 µm diam., subparallel, light brown (6D6). Context thin, orangish grey
cylindrical, yellowish brown to light brown, (5B3). Lamellae adnexed to adnate, close (15-
smooth, dextrinoid, thick-walled (up to 1.5 µm), 18) with 3-4 series of lamellulae, broad (2-3
non-gelatinous. Stipe trama hyphae 3-10 µm mm), pale greyish yellow, non-marginate or
diam, parallel, cylindrical, hyaline, smooth, slightly pale brown-marginate, non-intervenose.
dextrinoid, thin-walled, non-gelatinous. Caulo- Stipe 53-83 × 1-2 mm, central, cylindrical, wiry,
cystidia absent. Clamp connections present in hollow, glabrous, non-insititious, light brown
all tissues. (5D6) overall. Odor of coconut juice. Taste not
Habit, habitat and known distribution: distinctive.
Basidiospores 8-10(-12) × 4-5(-6) µm
leaves, Malaysia, neotropics, Northern
[xmr = 9.3-9.8 × 4.3-5.1 µm, xmm = 9.6 ± 0.3 ×
Thailand and Papua New Guinea.
Material examined: Thailand. Chiang Mai 4.8 ± 0.4 µm, Qmr = 1.9-2.2, Qmm = 2.1 ± 0.1, n
Province, Doi Suthep-Pui National Park, Mokfa = 25 spores, s = 3 specimens], ellipsoid,
Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014 smooth, hyaline, inamyloid, thin-walled.
m., 25 June 2005, N. Wannathes 329 (CMU, SFSU); Basidia not observed. Basidioles cylindrical to
Chiang Mai Province, Doi Inthanon National Park, At 25
clavate. Cheilocystidia scattered, of Siccus-type
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, broom cells; main body 9-16 × 4-7 µm,
SFSU); Phrae Province, Muang District, Suen Keun, Na 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, dextrinoid, thin-walled, non-gelatinous. Caulo-
cylindrical, obtuse to subacute, hyaline, thin- cystidia absent. Clamp connections present in
walled. Pleurocystidia common, 38-52 × 7-13 all tissues.
µm, cylindrical to fusoid or irregular in outline, Habit, habitat and known distribution:
wavy to constricted at the apex, hyaline, Scattered to gregarious on dicotyledonous
inamyloid, thin-walled. Pileipellis hymeniform, leaves. South America (Bolivia, Brazil, Peru,
mottled, composed of Siccus-type broom cells; Venezuela), Africa (Kenya, Uganda), Sri
main body (5-)8-14 × 4-7 µm, cylindrical to Lanka, and Northern Thailand.
clavate or irregular in outline, often branched, Material examined. Thailand. Chiang Mai
light brown, inamyloid, thin- to thick-walled; Highway 1,009 and road to Mae Chaem, N 18º 31.6’ E
apical setulae 4-11(-20) × 1-1.5(-2) µm, 98º 29.6’ alt. 1,703 m., 3 July 2004, Y. S. Tan 294 and
crowded, cylindrical, obtuse to subacute, 300 (CMU, SFSU); same location, 27 June 2005, N.
brown, thick-walled. Pileus trama interwoven, Wannathes 359 (CMU).
dextrinoid. 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
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
Fig. 58. Marasmius confertus (Y.S. Tan 294). 1. aurantiaco-albus, tenuis. Lamellae annexae usque
Basidiomes 2. Basidioles 3. Basidiospores 4. adnatae, distantes (6-8), angustae, flavo-albae, haud
Cheilocystidia 5. Pleurocystidia 6. Pileipellis, Scale bar 1 marginatae, haud intervenosae. Stipes 100-195 × 0.5 mm,
= 20 mm, 2-6 = 10 µm centralis, cylindratus, filo metallico similis, tenax, glaber,
insititius usque haud insititius, apice flavo-albo, basi
Lamellar trama hyphae 3-10(-14) µm diam., brunnea. Odor saporque non propria. Basidiosporae 25-
interwoven to regular, cylindrical to inflated, 32 × 4-4.5 µm, cylindratae usque clavatae, laeves,
smooth, hyaline, dextrinoid to weakly hyalinae, inamyloideae, tenuitunicatae Basidia 36-40 ×
9-10 µm, cylindrata usque clavata, 4-spora. Basidiolae
dextrinoid, thin-walled, non-gelatinous. cylindratae usque clavatae. Cheilocystidia vulgaria, typi
Stipitipellis hyphae 3-7(-9) µm diam., parallel, Sicci; 9-19 × 4-8(-13) µm, cylindrata usque clavata vel
cylindrical, yellowish brown to light brown, adumbratim inaequabilia, hyalina, inamyloidea,
smooth, inamyloid, thick-walled (up to 1 µm), tenuitunicata; setulis apicalibus 3-9 × 1-2 µm, cylindratis,
non-gelatinous. Stipe trama hyphae 7-14 µm saepe undulatis, raro furcatis, subacutis, pallide flavis
usque flavis, crassetunicatis. Pleurocystidia vulgaria, 51-
diam., parallel, cylindrical, hyaline, smooth,
283
82 × 7-13 µm, cylindrata usque fusoidea, plerumque often branched, pale yellow, inamyloid, thin- to
constricta ad apicem, hyalina, inamyloidea, tenuitunicata, thick walled; apical setulae 2-8 × 1-2 µm,
raro crassetunicata ad apicem. Pileipellis hymeniformis,
maculosus, typi Sicci; (5-)10-19 × 5-12 µm, cylindratus
crowded, cylindrical, wavy, subacute to acute,
usque clavatus vel adumbratim inaequabilis, saepe brown to dark brown, thick-walled. Pileus
ramosus, pallide flavus, inamyloideus, tenui- usque trama interwoven, dextrinoid. Lamellar trama
crassetunicatus; setulis apicalibus 2-8 × 1-2 µm, coartatis, hyphae 3-7 µm diam., interwoven, cylindrical
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, tenui-
usque 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, non-
intervenose. 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 Fig. 59 Marasmius ganyao (N. Wannathes 183). 1.
± 0.4 µm, Qmr = 6.2-7.0, Qmm = 6.6 ± 0.6, n = Basidiomes 2. Basidia 3. Basidiospores 4. Cheilocystidia
25 spores, s = 2 specimens], cylindrical to 5. Pleurocystidia 6. Pileipellis, Scale bar 1 = 20 mm, 2-6
clavate, smooth, hyaline, inamyloid, thin- = 10 µm
walled. Basidia 36-40 × 9-10 µm, cylindrical to to inflated, smooth, hyaline, weakly dextrinoid,
clavate, 4-spored. Basidioles cylindrical to thin-walled, non-gelatinous. Stipitipellis hy-
clavate. Cheilocystidia common, of Siccus-type phae 2-5 µm diam., subparallel, cylindrical,
broom cells; main body 9-19 × 4-8(-13) µm, brown to light brown, smooth, inamyloid to
cylindrical to clavate or irregular in outline, weakly dextrinoid, thin- to thick-walled, non-
hyaline, inamyloid, thin-walled; apical setulae gelatinous. Stipe trama hyphae 3-8 µm diam.,
3-9 × 1-2 µm, cylindrical, often wavy, seldom parallel, cylindrical, hyaline, smooth, inamy-
forked, subacute, pale yellow to yellow, thick- loid, thin- to thick-walled, non-gelatinous.
walled. Pleurocystidia common, 51-82 × 7-13 Caulocystidia absent. Clamp connections
µm, cylindrical to fusoid, usually constricted at present in all tissues.
the apex, hyaline, inamyloid, thin-walled, Habit, habitat and known distribution:
rarely apically thick-walled. Pileipellis hymeni- Scattered to gregarious on bamboo leaves,
form, mottled, composed of Siccus-type broom Northern Thailand.
cells; main body (5-) 10-19 × 5-12 µm, Material examined: Thailand. Chiang Mai
cylindrical to clavate or irregular in outline, Province, Huai Nam Dang National Park, North 19º
284
Fungal Diversity
18.3’ E 98º 35.8’ alt. 1,538 m., 24 June 2003, N. clavata, hyalina, inamyloidea, tenuitunicata; setulis
Wannathes 005 (CMU: holotype; SFSU: isotype); same apicalibus 3-12(-16) × 1-1.5(-2) µm, cylindratis, saepe
location, 28 June 2004, N. Wannathes 183 (CMU, SFSU) undulatis, raro furcatis, subacutis, hyalinis, tenuitunicatis.
Discussion: The new species is Pleurocystidia dispersa usque rara, 45-47 × 8-10 µm,
distinguished by a small obtusely conical, cylindrata usque fusoidea, plerumque undulata vel
sulcate pileus with a black to dark brown disc constricta ad apicem, hyalina, inamyloidea, tenuitunicata.
and brown margin with paler sulcae, distant (6- 8-12 µm, clavatus usque late clavatus vel turbinatus,
8), narrow lamellae, a long (up to 195 mm hyalinus, inamyloideus, tenui- usque crassetunicatus;
long ), wiry, brown stipe that is often insititious setulis apicalibus 5-18 × 1-2 µm, coartatis, cylindratis,
and grows on bamboo leaves, long clavate subacutis, atrobrunneis, crassetunicatis. Trama pilei
basidiospores with mean 28.2 × 4.3 µm, and intertexta, hyphis 5-10 µm diametro, cylindratis usque
inflatis, laevibus, hyalinis, dextrinoideis, tenuitunicatis.
simple strangulate pleurocystidia. Marasmus Trama lamellae regularis usque intertexta, hyphis 5-12
ganyao is similar to the following species. µm diametro, cylindratis, laevibus, hyalinis, leniter
Marasmius davidii Antonín, a species dextrinoideis, tenuitunicatis, haud gelatinosis.
previously known as M. brunneolus (Berk. & Stipitipellis parallelus, hyphis 4-9(-11) µm diametro,
Broome) Pegler (non M. brunneolus (Beeli) cylindratis, flavis usque laete brunneis, laevibus,
inamyloideis, tenuitunicatis, haud gelatinosis. Trama
Singer; cf. Antonín, 2003) and described from stipitis parallela, hyphis 6-8 µm diametro, cylindratis,
Sri Lanka, differs in forming larger pilei (7- hyalinis, laevibus, dextrinoideis, tenuitunicatis, haud
11mm vs 2-5 mm), a shorter stipe (30-45 mm), gelatinosis. Caulocystidia abundantia, plerumque
smaller basidiospores (20-27 × 4-6 µm, xm = 22 caespitosa, typi Sicci; 6-28 × 7-8 µm, cylindrata, hyalina
usque pallide flava, inamyloidea, tenuitunicata; setulis
× 4.5 µm), and has a lignicolous habit (Pegler,
apicalibus 5-40 × 1.5-2 µm, cylindratis, saepe furcatis,
1986). Marasmius longistipitatus Antonín, conicis usque subacutis, flavis, crassetunicatis (usque ad
from Africa, differs in forming a pale greyish 1 µm). Fibulae praesentes in omnibus texturis.
orange to orangish white pileus with a distinct Typus: Chiang Mai Province, Doi Inthanon
conical papilla, more numerous and National Park, Junction of Highway 1,009 and road to
pseudocollariate lamellae (9-12), a reddish Mae Chaem, N 18º 31.6’ E 98º 29.6’ alt. 1,703 m.,
dispersus usque gregarius in ligno vel folis plantae
orange to brownish orange stipe, and slightly dicotyledoneae, 2 August 2003, N. Wannathes 078
smaller basidiospores (20-28 × 4.5-6 µm) (CMU: holotypus; SFSU: isotypus).
(Antonín, 2007). Marasmius megistosporus Pileus 8-23 mm diam., campanulate,
Singer, described from Bolivia, differs in striate, glabrous, dull, disc dark brown, margin
forming larger pilei (up to 35 mm diam.) with brownish grey. Context thin, brownish cream.
deep ferruginous to golden brown margin, Lamellae adnexed, close (20-24) with 3 series
possibly longer basidiospores (28-37.5 µm of lamellulae, broad, greyish white, non-
long), and has a lignicolous habit (Singer, 1965, marginate, non-intervenose. Stipe 35-45 × 1.5-
1976). 2.5 mm, central, cylindrical, pruinose, non-
insititious, apex brownish yellow, base brown.
51. Marasmius graminipes Wannathes, Odor and taste not distinctive.
Desjardin & Lumyong, sp. nov. (Fig. 60) Basidiospores 18-21(-22) × 4 µm [x =
MycoBank: MB512427 19.3 ± 1.0 × 4.0 ± 0 µm, Q = 4.5-5.5, Qm = 4.8,
Etymology: ‘graminipes’ (Latin) = grass-stem;
referring to the tufts of grass-like caulocystidia on the
n = 25 spores, s = 1 specimen], narrowly
stipe surface. clavate to cylindrical, smooth, hyaline,
Pileus 8-23 mm diametro, campanulatus, striatus, inamyloid, thin-walled. Basidia not observed.
glaber, hebetatus, disco atrobrunneo, margine brunneo- Basidioles clavate. Cheilocystidia common, of
cinereus. Contextus brunneo-creameus, tenuis. Lamellae Siccus-type broom cells; main body 10-15 × 5-
annexae, arctae (20-24) cum 3 seriebus lamellularum,
latae, cinereo-albae, haud marginatae, haud intervenosae.
6 µm, cylindrical to clavate, hyaline, inamyloid,
Stipes 35-45 × 1.5-2.5 mm, centralis, cylindratus, thin-walled; apical setulae 3-12(-16) × 1-1.5(-2)
pruinosae, haud insititius, apice brunneo-flavo, basi µm, cylindrical, often wavy, seldom forked,
brunnea. Odor saporque non propria. Basidiosporae 18- subacute, hyaline, thin-walled. Pleurocystidia
21(-22) × 4 µm, anguste clavatae usque cylindratae, scattered to rarely present, 45-47 × 8-10 µm,
laeves, hyalinae, inamyloideae, tenuitunicatae Basidia
non observata. Basidiolae clavatae. Cheilocystidia cylindrical to fusoid, usually wavy or
vulgaria, typi Sicci; 10-15 × 5-6 µm, cylindrata usque constricted at the apex, hyaline, inamyloid, thin-
285
often forked, conical to subacute, yellow,
slightly thick-walled (up to 1 µm). Clamp
connections present in all tissues.
Scattered to gregarious on wood or on
dicotyledonous leaves, Northern Thailand.
98º 29.6’ alt. 1,703 m., 2 August 2003, N. Wannathes
078 (CMU: holotype; SFSU: isotype).
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 brown-
colored 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,
Fig. 60. Marasmius graminipes (N. Wannathes 078). 1. described from Malaysia, differs in forming a
Basidiomes 2. Basidioles 3. Basidiospores 4. non-striate pileus coloured more reddish brown,
Cheilocystidia 5. Pleurocystidia 6. Pileipellis 7. fewer lamellae (16-18) with brown edges,
Caulocystidia, Scale bar 1 = 20 mm, 2-7 = 10 µm
smaller basidiospores (mean 16.7 × 4.1 µm),
and Siccus-type broom cells on the stipe
walled. Pileipellis hymeniform, mottled,
surface with more well-developed basal cells,
composed of Siccus-type broom cells; main
although they do form setulae up to 60 µm long
body 12-27 × 8-12 µm, clavate to broadly
(Tan et al., 2007). Marasmius bondoi (see
clavate or turbinate, hyaline, inamyloid, thin-
below) differs in forming slightly smaller
to thick-walled; apical setulae 5-18 × 1-2 µm,
basidiospores with mean range 14.2-18.3 × 3.5-
crowded, cylindrical, subacute, dark brown,
4.0 µm, fewer lamellae (12-18), and a golden
thick-walled. Pileus trama hyphae 5-10 µm
brown stipe that lacks caulocystidia. Although
diam., interwoven, cylindrical to inflated,
in the ML phylogenetic tree (Fig. 1) the
smooth, hyaline, dextrinoid, thin-walled.
presence of M. graminipes makes M. bondoi
Lamellar trama hyphae 5-12 µm diam., regular
paraphyletic, a separate branch and bound
to interwoven, cylindrical, smooth, hyaline,
analysis of a subset including only M. bondoi
weakly dextrinoid, thin-walled, non-gelatinous.
and M. graminipes isolates with M. plicatulus
Stipitipellis hyphae 4-9(-11) µm diam., parallel,
Peck as an outgroup (Fig. 74), indicates that M.
cylindrical, yellow to light brown, smooth,
graminipes is sister to all M. bondoi isolates
inamyloid, thin-walled, non-gelatinous. Stipe
and supports our recognition of M. graminipes
as a distinct taxon.
walled, non-gelatinous. Caulocystidia abundant, 52. Marasmius brunneoolivascens Wannathes,
usually present in cespitose clusters of many Desjardin & Lumyong, sp. nov.
cells, composed of Siccus-type broom cells (Figs 59-6, 61)
with main body 6-28 × 7-8 µm, cylindrical, MycoBank: MB512428
hyaline to pale yellow, inamyloid, thin-walled; Etymology: ‘brunneo’ (Latin) = brown,
apical setulae 5-40 × 1.5-2 µm, cylindrical, ‘olivascens’ (Latin) = olive; referring to the olive brown
color of the pileus and lamellae.
286
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 brunneo-
marginatae 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, crassetuni-
catis; b) cellulis haud setulosis 30-50 × 5-9 µm,
cylindratis usque clavatis, constrictis et interdum lobatis
ad apicem, hyalinis, inamyloideis, tenuitunicatis.
5-12 µm, cylindratus usque clavatus vel pyriformis,
saepe ramosus, hyalinus usque pallide flavus, Fig. 61. Marasmius brunneoolivascens (N. Wannathes 227).
inamyloideus, tenui- usque crassetunicatus; setulis 1. Basidiomes 2. Basidioles 3. Basidiospores 4.
apicalibus 2-7(-12) × 1-1.5(-2.5) µm, coartatis, Cheilocystidia 5a. Siccus-type pleurocystidia 5b. Non-
cylindratis, subacutis usque acutis, brunneis usque setulose pleurocystidia 6. Pileipellis, Scale bar 1 = 20
atrobrunneis, crassetunicatis. Trama pilei intertexta, mm, 2-6 = 10 µm
dextrinoidea usque fortiter dextrinoidea. Trama lamellae
intertexta usque regularis, hyphis 3-8(-22) µm diametro, and margin brown to greyish brown. Context
cylindratis usque inflatis, laevibus, hyalinis, greyish cream, thin. Lamellae adnexed to free,
dextrinoideis usque fortiter dextrinoideis, tenuitunicatis, subdistant (12-18) with 2-3 series of lamellulae,
haud gelatinosis. Stipitipellis subparallelus, hyphis 3-9(-
13) µm diametro, cylindratis, brunneis usque laete broad (1-2 mm), olive brown to greyish olive
brunneis, laevibus, dextrinoideis, crassetunicatis (usque or yellowish olive, sometimes with brown to
ad 3 µm), haud gelatinosis. Trama stipitis parallela, olive-brown edges, non-intervenose. Stipe 15-
hyphis (3-)5-13(-20) µm diametro, cylindratis, hyalinis 55 × 0.5-1 mm, central, cylindrical, wiry,
usque pallide flavis, laevibus, dextrinoideis, tenuituni-
catis, haud gelatinosis. Caulocystidia nulla. Fibulae
hollow, glabrous, non-insititious, apex
praesentes in omnibus texturis. yellowish white, base brown to reddish brown.
Typus: Chiang Mai Province, Doi Suthep-Pui Odor and taste not distinctive.
National Park, Sangasabhasri Land on the way to Huai Basidiospores (11-)12-14(-15) × 4-5(-6)
Kok Ma Village, N 18º 48.4’ E 98º 54.6’ alt. 1,146 m.,
µm [xmr = 12.6-13.8 × 4.0-5.0, xmm = 13.0 ± 0.3
dispersus usque gregarius in folis plantae dicotyledoneae
vel ligno, 14 August 2003, N. Wannathes 112 (CMU: × 4.4 ± 0.4 µm, µm, Qmr = 2.7-3.4, Qmm = 3.0 ±
holotypus; SFSU: isotypus). 0.2, n = 25 spores, s = 11 specimens], fusoid,
Pileus 6-23 mm diam., convex with a inequilateral, smooth, hyaline, inamyloid, thin-
rugulose umbo when young, expanding to walled. Basidia20-22 × 6-8 µm, cylindrical to
plano-convex and slightly depressed in age, clavate, 4-spored. Basidioles fusoid to clavate.
striate to sulcate, pruinose, dull, dark brown Cheilocystidia abundant, of Siccus-type broom
overall when young, in age disc dark brown cells; main body 12-28 × 5-9 µm, cylindrical to
287
clavate, sometimes catenulate, hyaline, Discussion: Marasmius brunneooliva-
inamyloid, thin-walled; apical setulae 2-9(-10) scens is characterized by a striate to sulcate
× 1-1.5 µm, cylindrical, subacute, yellow to pileus coloured brown to greyish brown with a
brown, thick-walled. Pleurocystidia composed dark brown disc, subdistant (12-18) broad
of 2 type of cells: a) Siccus-type broom cells lamellae colored olive brown to greyish olive
like the cheilocystidia, scattered and rare; main or yellowish olive with or without brown to
body 12-22 × 6-8 µm, cylindrical to clavate, olive brown edges, a glabrous stipe, fusoid
hyaline, inamyloid, thin-walled; apical setulae basidiospores with mean 13.0 × 4.4 µm and
2-7 × 1-1.5 µm, cylindrical, subacute, yellow, mean Q = 3, two types of pleurocystidia
inamyloid, thick-walled; b) non-setulose cells, (simple strangulate cells and scattered Siccus-
30-50 × 5-9 µm, cylindrical to clavate, type broom cells), and an absence of
constricted and sometimes lobed at the apex, caulocystidia. This new species is similar to
hyaline, inamyloid, thin-walled. Pileipellis two African species. Marasmius grandisetu-
hymeniform, mottled, composed of Siccus-type losus Singer differs in forming white to cream-
broom cells; main body 8-23 × 5-12 µm, coloured lamellae, longer basidiospores (17-23
cylindrical to clavate or pyriform, often µm long) and lacks olive tones (Singer, 1964;
branched, hyaline to pale yellow, inamyloid, Antonín, 2007). Marasmius elaeocephalus
thin-to thick-walled; apical setulae 2-7(-12) × Singer sensu Antonín (2007) differs in forming
1-1.5(-2.5) µm, crowded, cylindrical, subacute an olive brown pileus with yellow tones on the
to acute, brown to dark brown, thick-walled. margin, yellowish white lamellae with
Pileus trama interwoven, dextrinoid to strongly concolorous edges, and lacks two types of
dextrinoid. Lamellar trama hyphae 3-8(-22) pleurocystidia.
µm diam., interwoven to regular, cylindrical to
53. Marasmius suthepensis Wannathes,
inflated, smooth, hyaline, dextrinoid to strongly
Desjardin & Lumyong, sp. nov.
(Figs 59-8, 62)
Stipitipellis hyphae 3-9(-13) µm diam.,
MycoBank: MB512429
subparallel, cylindrical, brown to light brown, Etymology: ‘suthep’ = Doi Suthep-Pui National
smooth, dextrinoid, thick-walled (up to 3 µm), Park; referring to the place where the holotype specimen
non-gelatinous. Stipe trama hyphae (3-)5-13(- was collected.
20) µm diam., parallel, cylindrical, hyaline to Pileus 10-27 mm diametro, obtuse conicus usque
pale yellow, smooth, dextrinoid, thin-walled, convexus, cum vel sine umbone, laevis usque striatus,
glaber, hebetatus, disco clare brunneo, margine brunneo-
non-gelatinous. Caulocystidia absent. Clamp flavo usque brunneo-aurantiaco. Contextus flavo-albus,
connections present in all tissues. tenuis. Lamellae annexae usque liberae, subdistantes
Habit, habitat and known distribution: (12-18) cum 3-4 seriebus lamellularum, latae (1-2 mm),
Scattered to gregarious on dicotyledonous pallide flavo-albae, haud marginatae, haud intervenosae.
leaves or on wood, Northern Thailand. Stipes 18-55 × 1 mm, centralis, cylindratus, filo
Material examined: Thailand. Chiang Mai metallico similis, cavus, glaber, haud insititius, apice
Province, Doi Suthep-Pui National Park, Sangasabhasri flavo-albo, basi brunnea. Odor saporque non propria.
Land on the way to Huai Kok Ma Village, N 18º 48.4’ E Basidiosporae (10-)11-14(-15) × (3.5-)4-5 µm, anguste
98º 54.6’ alt. 1,146 m., 29 July 2003, N. Wannathes 055 ellipsoideae usque fusoideae, laeves, hyalinae,
(CMU, SFSU); same location, 14 August 2003, N. inamyloideae, tenuitunicatae Basidia 13-15 × 5-6 µm,
Wannathes 112 (CMU: holotype; SFSU: isotype) and cylindrata usque clavata, 4-spora. Basidiolae cylindratae
N.Wannathes 113 (CMU); same location, 16 July 2004, usque clavatae. Cheilocystidia abundantia, typi Sicci; (6-
N. Wannathes 228 (CMU); same location, 24 June )12-18 × (3-)7-11 µm, cylindrata usque clavata, hyalina,
2005,N. Wannathes 322 (CMU); Chiang Mai Province, inamyloidea, tenuitunicata; setulis apicalibus 2-6 × 1 µm,
Mae Taeng District, Tung Joaw Village, N 19º 8.07’ E cylindratis usque conicis, obtusis usque subacutis, flavis
98º 38.9’ alt. 1423 m., 31 July 2003, N. Wannathes 063 usque flavo-brunneis, tenuitunicatis. Pleurocystidia
(CMU, SFSU); same location, 22 August 2004, N. vulgaria, 27-43 × 5-7 µm, cylindrata usque fusoidea,
Wannathes 277 (CMU, SFSU); Chiang Mai Province, undulata usque constricta et interdum lobata ad apicem,
Huai Nam Dang National Park, North 19º 18.3’ E 98º hyalina, inamyloidea, tenuitunicata. Pileipellis hymeni-
35.8’ alt. 1,538 m., 29 June 2005, N. Wannathes 373 formis, maculosus, typi Sicci; (7-)10-17(-22) × 4-10 µm,
(CMU, SFSU); Chiang Mai Province, Doi Suthep-Pui cylindratus usque clavatus vel pyriformis, hyalinus
National Park, Medicinal Plant Garden, 3 August 2005, usque pallide flavus, inamyloideus, tenuitunicatus;
N. Wannathes 397 (CMU, SFSU). setulis apicalibus 3-8 × 1(-1.5) µm, coartatis, cylindratis,
subacutis, brunneis usque laete brunneis, crassetunicatis.
288
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,
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-)5-
10(-13) µm diametro, cylindratis, hyalinis, laevibus,
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 Fig. 62. Marasmius suthepensis (Y.S. Tan 280). 1.
to convex, with or without an umbo, smooth to Basidiomes 2. Basidia 3. Basidiospores 4. Cheilocystidia
striate, glabrous, dull, disc bright brown, 5. .Pleurocystidia 6. Pileipellis, Scale bar 1 = 20 mm, 2-6
margin brownish yellow to brownish orange. = 10 µm
Context thin, yellowish white. Lamellae
adnexed to free, subdistant (12-18) with 3-4 hyphae (4-)6-13(-17) µm diam., interwoven,
series of lamellulae, broad (1-2 mm), pale cylindrical to inflated, smooth, hyaline,
yellowish white, non-marginate, non- strongly dextrinoid, thin-walled. Lamellar
intervenose. Stipe 18-55 × 1 mm, central, hyphae 3-7(-12) µm diam., trama interwoven
cylindrical, wiry, hollow, glabrous, non- to regular, cylindrical to inflated, smooth,
insititious, apex yellowish white, base brown. hyaline, dextrinoid, thin-walled, non-gelatinous.
Odor and taste not distinctive. Stipitipellis hyphae 3-7(-9) µm diam.,
Basidiospores (10-)11-14(-15) × (3.5-)4- subparallel, cylindrical, brown to dark brown,
5 µm [xmr = 11.7-13.6 × 3.9-4.1 µm, xmm = smooth, inamyloid to weakly dextrinoid, thin-
walled, non-gelatinous. Stipe trama hyphae (3-
12.6 ± 1.0 × 4.0 ± 0.1 µm, Qmr = 2.9-3.8, Qmm
)5-10(-13) µm diam., parallel, cylindrical,
hyaline, smooth, dextrinoid, thin-walled, non-
narrowly ellipsoid to fusoid, smooth, hyaline,
gelatinous. Caulocystidia absent. Clamp
inamyloid, thin-walled. Basidia 13-15 × 5-6 connections present in all tissues.
µm, cylindrical to clavate, 4-spored. Basidioles Habit, habitat and known distribution.
cylindrical to clavate. Cheilocystidia abundant, Scattered to gregarious on dicotyledonous
of Siccus-type broom cells; main body (6-)12- leaves or on wood, Northern Thailand.
18 × (3-)7-11 µm, cylindrical to clavate, Material examined: Thailand. Chiang Mai
hyaline, inamyloid, thin-walled; apical setulae Province, Doi Suthep-Pui National Park, Sangasabhasri
2-6 × 1 µm, cylindrical to conical, obtuse to Land on the way to Huai Kok Ma Village, N 18º 48.4’ E
subacute, yellow to yellowish brown, thin-
walled. Pleurocystidia common, 27-43 × 5-7 Wannathes 057 (CMU, SFSU); Chiang Mai Province,
µm, cylindrical to fusoid, wavy to constricted Doi Suthep-Pui National Park, Mokfa Waterfall, on Hwy
and sometimes lobed at the apex, hyaline, 1095, N19º 6.5’.E 98º 46.3’ alt. 1014 m., 29 June 2004,
inamyloid, thin-walled. Pileipellis hymeniform, Y.S. Tan 280 (CMU: holotype; SFSU: isotype).
mottled, composed of Siccus-type broom cells; Discussion: Diagnostic features of
main body (7-)10-17(-22) × 4-10 µm, cylindrical Marasmius suthepensis include: a smooth to
to clavate or pyriform, hyaline to pale yellow, striate pileus coloured brownish yellow to
brownish orange with bright brown disc;
inamyloid, thin-walled; apical setulae 3-8 × 1(-
subdistant (12-18), broad lamellae that are pale
1.5) µm, crowded, cylindrical, subacute, brown
yellowish white and non-marginate; a glabrous,
to light brown, thick-walled. Pileus trama
brown stipe; basidiospores with mean 12.6 ×
289
4.0 µm and mean Q = 3.7; simple cylindrical haematocephalus forms with M. siccus as an
pleurocystidia, and an absence of caulocystidia. outgroup (Fig. 70) indicates that several forms
The Thai new species is similar to two brown- are well-supported by BS values 92-100%
coloured neotropical species. Marasmius (forms haematocephalus, “atrobrunneus”,
hinnuleus Berk. & M.A. Curtis, described from “violaceus”), whereas other forms show
Cuba, differs in forming smaller (4-12 mm vs invariability (forms “variabilis”, “luteocepha-
10-27 mm), sulcate to plicate pilei coloured lus”) or form a polytomy (“f. robustus”).
pale reddish brown to cinnamon-rufous, and Marasmius haematocephalus as it occurs in
slightly longer spores with mean 14 × 3.8 µm Southeast Asia will remain a complex of
(Singer, 1976; Pegler, 1983). Marasmius morphologically and genetically distinct
helvolus Berk., described from Brazil, differs entities without formal taxonomic recognition
in forming distinctly sulcate to plicate pilei, until further specimens are analyzed in the
fewer lamellae (9-11) and usually lacks hopes of clarifying their differences and
pleurocystidia (rare in some basidiomes) relationships. The forms recognized below will
(Singer, 1976; Pegler, 1983; Desjardin and not be formally proposed and are named herein
Ovrebo, 2006). Marasmius suthepensis is also only to aid in documenting their existence.
similar to M. brunneoolivascens (see above),
which may occur at the same site, but the latter Key to forms of Marasmius haematocephalus
species differs in forming olive toned lamellae
and Siccus-type broom cells are often present 1. Pileus yellowish orange or with a pale brownish
yellow disc and nearly white margin .........................
as pleurocystidia on the lamellar sides. ......... 54A. M. haematocephalus “f. luteocephalus”
1. Pileus with red, violet, olive, greyish blue or brown
54. Marasmius haematocephalus (Mont.) Fr., pigments................................................................... 2
Epicr. Syst. Mycol: 382. 1838.
≡ Agaricus haematocephalus Mont. Ann. Sci. 2. Pileus 1-5 mm diam, dark brown overall, lacking red,
Nat., Bot., sér. 2, 8: 369. 1837. violet or purple...........................................................
≡ Androsaceus haematocephalus (Mont.) Pat., J. ..........54B. M. haematocephalus “f. atrobrunneus”
Bot. (Morot) 3: 336. 1889. 2. Pileus > 5 mm diam, with red, violet, purple, olive, or
= Marasmius rhodocephalus Fr., Nova Acta greyish blue tones ................................................... 3
Regiae Soc. Sci. Upsal ser. 3, 1: 31. 1851.
≡ Androsaceus rhodocephalus (Fr.) Pat. & 3. Pileus olive to greyish blue overall when young, disc
Gaillard, Bull. Soc. Mycol. France 4: 20. 1888. often with red, orange or reddish purple tones in age
= Marasmius semipellucidus Berk. & Broome, J. .................54C. M. haematocephalus “f. variabilis”
Linn. Soc., Bot. 14: 36. 1873. 3. Pileus lacking olive and greyish blue tones ............. 4
= Marasmius sanguineus Cooke & Massee,
Grevillea 17: 59. 1889. 4. Pileus typically 10-20(-30) mm diam., dark reddish
= Marasmius atropurpureus Murrill, N. Amer Fl. brown to dark violet brown (10-11F6-8) when young,
9: 262. 1915. fading to reddish-greyish brown in age......................
= Marasmius vinosus Beeli, Bull. Soc. Roy. Bot. ..................54D. M. haematocephalus “f. robustus”
Belgique 60: 158. 1928. 4. Pileus typically <10 mm diam, deep reddish brown to
maroon or deep violet-purple................................... 5
There occur in Thailand six distinct
forms of M. haematocephalus, that share 5. Pileus magenta, deep violet-purple or dark purple
basidiospores in the range 19-25 × 3.5-5 µm (13-14E-F6-8); basidiospores normally with mean in
with xmm = 20.7 × 4.0 µm and Qmm = 5.2, well- the range 20.5-22 × 3.8-4.2 µm; basidiomes often
developed, simple and strangulate pleuro- associated with bamboo leaves .................................
.................54E. M. haematocephalus “f. violaceus”
cystidia, two types of cheilocystidia, lack 5. Pileus deep reddish brown to maroon (9-11E-F7-8);
caulocystidia, and fall into a well-supported basidiospores normally with mean in the range 17.5-
clade based on ITS sequences (Fig. 1). They 18 × 4.0-4.3 µm; basidiomes usually associated with
differ in pileus size and pigmentation, ranging dicot leaves ................................................................
from yellowish orange to deep reddish brown, ....... 54F. M. haematocephalus f. haematocephalus
maroon, dark violet, purple and purplish brown,
and in substrate preference. The ITS sequence 54A Marasmius haematocephalus “f.
data, however, are of limited use in luteocephalus” Wannathes, Desjardin &
distinguishing the forms. A branch and bound Lumyong, nom. prov. (Figs 63, 69-1)
analysis that includes sequences of all M.
290
Fungal Diversity

convex, 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
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,
clavate. Cheilocystidia composed of 2 types of
cells: a) Siccus-type broom cells with main
body 12-19 × 7-9 µm, cylindrical to clavate,
2-5 × 1 µm, cylindrical, often forked, subacute
to obtuse, hyaline to yellow, thick-walled; b)
non-setulose cells like the pleurocystidia, 36-41 Fig. 63. Marasmius haematocephalus “f. luteocephalus” (N.
× 11-13 µm, fusoid, usually attenuated and Wannathes 310). 1. Basidiomes 2. Basidiospores 3a.
strangulate at the apex, hyaline, inamyloid, Siccus-type cheilocystidia 3b. Non-setulose
thin-walled. Pleurocystidia common, 52-58(- cheilocystidia 4. Pleurocystidia 5. Pileipellis, Scale bar 1
= 10 mm, 2-5 = 10 µm
62) × 10-12 µm, fusoid to clavate, usually
attenuated and strangulate at the apex, hyaline, Material examined: Thailand. Chiang Rai
inamyloid, thin-walled. Pileipellis hymeniform, Province, Muang District, Khun Con Waterfall, 12 June
mottled, composed of Siccus-type broom cells; 2005, N. Wannathes 310 (CMU, SFSU).
main body 12-15× 8-14 µm, clavate to broadly Discussion: Marasmius haematocephalus
clavate or pyriform, sometime branched, “f. luteocephalus” differs from all other forms
yellow to hyaline, inamyloid, thin-walled; in having a yellowish orange to yellowish
apical setulae 2-4 × 1 µm, cylindrical, subacute, white pileus with pale brownish yellow disc
yellow to pale yellow, thick-walled. Pileus and cream-coloured lamellae with yellowish
trama interwoven, dextrinoid. Lamellar trama orange edges. This provisional taxon looks like
hyphae 5-10(-20) µm diam., interwoven, a paler form of M. haematocephalus var.
cylindrical to inflated, smooth, hyaline, leucophyllus Singer, described from Bolivia,
dextrinoid, thin-walled, non-gelatinous. but the latter differs in forming a darker pileus
Stipitipellis hyphae 3-5 µm diam., subparallel, (fawn to pinkish cinnamon), and smaller
cylindrical, greenish brown, smooth, dextrinoid, basidiospores in the range 15-20 × 3.5 µm
thin- to thick-walled, non-gelatinous. Stipe (Singer, 1976).
trama hyphae (4-)7-10 µm diam., parallel,
cylindrical, hyaline, smooth, dextrinoid, thin- 54B. Marasmius haematocephalus “f.
walled, non-gelatinous. Caulocystidia absent. atrobrunneus” Wannathes, Desjardin &
Clamp connections present. Lumyong, nom. prov. (Fig. 64)
Habit, habitat and known distribution: Pileus 1-5 mm diam., obtusely conical to
Scattered to gregarious on dicotyledonous 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, Fig. 64. Marasmius haematocephalus “f. atrobrunneus” (N.
cylindrical to clavate or pyriform, hyaline, Wannathes 117). 1. Basidiomes 2. Basidiospores 3a.
inamyloid, thin-walled; apical setulae 2-4 × 1 Siccus-type cheilocystidia 3b. Non-setulose
cheilocystidia 4. Pleurocystidia 5. Pileipellis, Scale bar 1
µm, cylindrical, often forked, subacute, hyaline = 10 mm, 2-5 = 10 µm
to yellow, thick-walled; b) non-setulose cells
like the pleurocystidia, 35-40 × 7-9 µm, fusoid temple in Padeng Village, 22 August 2003, N.
to clavate, usually attenuated and strangulate at Wannathes 117 (CMU, SFSU).
the apex, often with 2-4 successive apical Discussion: Marasmius haematocephalus “f.
constrictions, hyaline inamyloid, thin-walled. atrobrunneus” differs from all other forms of
Pleurocystidia common, 34-48 × 7-9 µm, this species in forming a smaller (1- 5 mm
clavate to fusoid, usually attenuated and diam.) dark brown pilei without red, violet,
strangulate at the apex, sometimes with 2 purple or yellowish orange tones.
successive apical constrictions, hyaline,
inamyloid, thin-walled. Pileipellis hymeniform, 54C. Marasmius haematocephalus “f.
mottled, composed of Siccus-type broom cells; variabilis” Wannathes, Desjardin & Lumyong,
main body 12-16 × 7-10 µm, clavate to broadly nom. prov. (Figs 65, 69-3)
clavate or pyriform, sometime branched, Pileus 5-12 mm diam., convex, sulcate,
yellow to hyaline, inamyloid, thin- to thick- pruinose, dull, when young olive to greyish
walled; apical setulae 2-5 × 1 µm, cylindrical, blue overall or disc dark purple and margin
subacute, brown to dark brown, thick-walled. brownish purple with dark brownish grey to
Pileus trama interwoven, inamyloid. Lamellar dark olive sulcae, when mature disc dark
trama hyphae 4-6 µm diam., regular to purple and margin fading to reddish brown or
interwoven, cylindrical, smooth, hyaline, purplish brown with greyish blue sulcae.
inamyloid, thin-walled, non-gelatinous. Context brownish grey, thin. Lamellae adnexed,
Stipitipellis hyphae 3-5 µm diam., parallel, subdistant (10-13), narrow to broad (1-2 mm),
cylindrical, brown to greenish brown, smooth, pinkish cream with or without pinkish brown
inamyloid, thin- to thick-walled (up to 0.5 µm), edges, non-intervenose. Stipe 20-32 × 0.5-1
non-gelatinous. Stipe trama hyphae 2-5 µm mm, central, cylindrical, wiry, hollow, glabrous,
diam., parallel, cylindrical, hyaline, smooth, non-insititious, apex purple, base dark
weakly dextrinoid, thin-walled, non-gelatinous. brownish purple. Odor and taste not distinctive.
Caulocystidia absent. Clamp connections Basidiospores (17-)19-22(-24) × 3.5-5(-
present. 5.5) µm [xmr = 20.3-21.6 × 3.9-4.2 µm, xmm =
Habit, habitat and known distribution: 21.0 ± 0.9 × 4.1 ± 0.3 µm, Qmr = 5.1-5.3, Qmm
Scattered to gregarious on grass leaves, = 5.2 ± 0.1, n = 25 spores, s = 2 specimens],
Northern Thailand. clavate to fusoid, often curved in profile,
Material examined: Thailand. Chiang Mai smooth, hyaline, inamyloid, thin-walled.
Province, Mae Taeng District, Street to Pathummigaram Basidia not observed. Basidioles fusoid to
292
Fungal Diversity

body 15-20 × 5-7 µm, cylindrical to clavate,
2-10 × 1(-1.5) µm, cylindrical, subacute to
acute, greenish brown, thick-walled; b) non-
setulose cells like the pleurocystidia, 40-52× 7-
8 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
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
weakly dextrinoid. Lamella trama hyphae 3-9
µm diam., interwoven, cylindrical, smooth,
hyaline, weakly dextriomoid, thin-walled, non-
subparallel, cylindrical, brown, smooth, Fig. 65. Marasmius haematocephalus “f. variabilis” (N.
inamyloid to weakly dextrinoid, thin- to thick- Wannathes 433). 1. Basidiomes 2. Basidiospores 3a.
Siccus-type cheilocystidia 3b. Non-setulose
walled (up to 1 µm), non-gelatinous. Stipe
trama hyphae 3-6 µm diam., parallel, = 10 mm, 2-5 = 10 µm
cylindrical, hyaline, smooth, inamyloid to
weakly dextrinoid, thin-walled, non-gelatinous. haematocephalus.
present. 54D. Marasmius haematocephalus “f.
Habit, habitat and known distribution: robustus” Wannathes, Desjardin & Lumyong,
Scattered to gregarious on dicotyledonous or nom. prov. (Figs 66, 69-2)
bamboo leaves, Northern Thailand. Pileus 10-20(-30) mm diam., obtusely
Material examined: Thailand. Chiang Mai conical when young, convex to plano-convex
Province, Doi Suthep-Pui National Park, Mokfa Waterfall, in age, disc rugulose, sulcate to plicate,
on Hwy 1095, N19º 6.5’ E 98º 46.3’ alt. 1014 m., 25
june 2005, N. Wannathes 336 (CMU, SFSU); Phrae
glabrous, dull, dark reddish brown to dark
Province, Muang District, Cherng Thong Waterfall, 16 violet brown (10-11F6-8) when young, fading
August 2005, N. Wannathes 415 (CMU); Phrae Province, to reddish-greyish brown to greyish purple in
Muang District, Pa Dang, Nong Kam Village, near Nong age. Context greyish cream, thin. Lamellae
Kam reservoir, 18 August 2005, N. Wannathes 430 adnexed, distant to subdistant (9-15), broad (2-
(CMU, SFSU).
4 mm), yellowish grey (2B2) to greyish yellow
Discussion: Marasmius haematocephalus “f. (4B3), non-marginate or brown-marginate,
variabilis” differs from the other forms of M.
non-intervenose. Stipe 13-43(-62) × 0.5-1 mm,
haematocephalus in forming pilei that display
central, cylindrical, wiry, hollow, glabrous,
the full rainbow of colours found in other
non-insititious, apex greyish white, base dark
forms throughout development, including blue,
olive, red, violet, purple and grey. However, “f.
Basidiospores (16-)18-23(-24) × (3.5-)4-
variabilis” still forms pinkish cream lamellae
with or without pinkish brown edges, and a 5(-6) µm [xmr = 18.5-21.6 × 3.9-4.5 µm, xmm =
stipe that has a purple apex and dark brownish 20.0 ± 1.3 × 4.1 ± 0.1 µm, Qmr = 4.2-5.5, Qmm
purple base, colors characteristic of f. = 4.9 ± 0.4, n = 25 spores, s = 9 specimens],
293
clavate to fusoid, often curved in profile,
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) non-
setulose cells like the pleurocystidia, 35-53× 8-
12 µm, fusoid to clavate, usually attenuated
thin-walled. Pleurocystidia common, 38-62 ×
6-15 µm, clavate to fusoid, usually attenuated
thin-walled. Pileipellis hymeniform, not
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
(2-)4-6(-14) µm diam., interwoven, cylindrical
to inflated, smooth, hyaline, weakly dextrinoid, Fig. 66. Marasmius haematocephalus “f. robustus” (N.
thin-walled, non-gelatinous. Stipitipellis Wannathes 433). 1. Basidiomes 2. Basidiospores 3a.
hyphae 3-9(-12) µm diam., subparallel, Siccus-type cheilocystidia 3b. Non-setulose
cylindrical, brown to dark brown, smooth, = 10 mm, 2-5 = 10 µm
weakly dextrinoid, thin- to thick-walled (up to
1 µm), non-gelatinous. Stipe trama hyphae 3- Muang District, Cherng Thong Waterfall, 16 August
12 µm diam., parallel, cylindrical, hyaline, 2005, N. Wannathes 417 (CMU, SFSU); same location,
smooth, dextrinoid, thin-walled, non-gelatinous. 19 August 2005, N. Wannathes 433 (CMU, SFSU).
Caulocystidia absent. Clamp connections Discussion: Marasmius haematocephalus “f.
present. robustus” is unique because of its robust pilei
Habit, habitat and known distribution: (10-30 mm diam.) and range of pileus colours
Scattered to gregarious on dicotyledonous that are similar to both forms haematocephalus
leaves or wood, Northern Thailand. and “violaceus”. This provisional Thai form is
phenetically similar to M. haematocephalus var.
Land on the way to Huai Kok Ma Village, N 18º 48.4’ E macrocephalus Singer, described from Bolivia,
98º 54.6’ alt. 1,146 m., 29 July 2003, N. Wannathes 053 except that the latter grows on monocot leaves
(CMU, SFSU); same location, 14 August 2003, N. (Singer, 1976).
Wannathes 110 and 111, (CMU, SFSU); Chiang Mai
54E. Marasmius haematocephalus “f.
98º 29.6’ alt. 1,703 m., 3 July 2004, N. Wannathes 203 violaceus” Wannathes, Desjardin & Lumyoung,
(CMU, SFSU); Chiang Rai Province, Muang District, nom. prov. (Figs 67, 69-4)
Khun Con Waterfall, 12 June 2005, N. Wannathes 301 Pileus 4-18 mm diam., obtusely conical
(CMU, SFSU); Chiang Mai Province, Doi Suthep-Pui to convex, umbonate, sometimes with a
National Park, Medicinal Plat Garden, 16 June 2005, N.
Wannathes 314 (CMU, SFSU); Chiang Mai Province,
rugulose disc, striate to sulcate, pruinose to
Doi Suthep-Pui National Park, Mokfa Waterfall, on Hwy finely velutinous, dull, dark purple (14F7) to
1095, N19º 6.5’.E 98º 46.3’ alt. 1014 m., 25 June 2005, dark magenta (13F7) overall when young, in
N. Wannathes 330 (CMU, SFSU); Phrae Province, age disc dark violet to greyish violet (14C4)
294
Fungal Diversity
and margin pinkish purple to violet. Context inamyloid, thin-walled. Pleurocystidia

purplish white (14A3), thin. Lamellae adnexed, common, 38-73 × 7-13 µm, clavate to fusoid,
distant (7-12), narrow to broad (1-2 mm), usually attenuated and strangulate at the apex,
purplish pink (14A3), non-marginate, non- hyaline, inamyloid, thin-walled. Pileipellis
intervenose. Stipe 13-30 × 0.2-0.5 mm, central, hymeniform, not mottled, composed of Siccus-
cylindrical, wiry, hollow, glabrous, non- type broom cells; main body 10-17 × 4-10 µm,
insititious, apex purplish white, base dark cylindrical to clavate or broadly clavate,
brown. Odor and taste not distinctive. hyaline, inamyloid, thin- to thick-walled; apical
setulae 2-5(-8) × 1 µm, cylindrical, subacute to
acute, brown, thick-walled. Pileus trama
3-8(-12) µm diam., interwoven, cylindrical,
gelatinous. 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)
present.
leaves or on bamboo debris, Northern Thailand.
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 Suthep-
Pui National Park, Mokfa Waterfall, on Hwy 1095, N19º
Fig. 67. Marasmius haematocephalus “f. violaceus” (N. 6.5’.E 98º 46.3’ alt. 1014 m., 25 June 2005, N.
Wannathes 413). 1. Basidiomes 2. Basidiospores 3a. Wannathes 339 (CMU, SFSU); Phrae Province, Muang
Siccus-type cheilocystidia 3b. Non-setulose District, Cherng Thong Waterfall, 16 August 2005, N.
cheilocystidia 4. Pleurocystidia 5. Pileipellis, Scale bar 1 Wannathes 413 (CMU, SFSU).
= 10 mm, 2-5 = 10 µm Discussion: This taxon represents the
dark violet-purple form commonly encountered
Basidiospores (19-)20-24(-25) × 3.5-5 in Southeast Asia and reported only as M.
µm [xmr = 20.6-21.8 × 3.8-4.2 µm, xmm = 21.4 haematocephalus from Papua New Guinea by
± 0.6 × 4.0 ± 0.1 µm, Qmr = 5.3-5.5, Qmm = 5.4 Desjardin and Horak (1997). It differs from f.
± 0.1, n = 25 spores, s = 4 specimens], clavate haematocephalus, as reported by Desjardin et
to fusoid, often curved in profile, smooth, al. (2000) from Indonesia, in that the latter
hyaline, inamyloid, thin-walled. Basidia not form has deep reddish brown to maroon pilei
observed. Basidioles fusoid to clavate. (lacking obvious deep purple and violet tones).
Cheilocystidia composed of 2 types of cells: a) The provisional Thai form may be identical to
Siccus-type broom cells with main body 10-21 M. haematocephalus var. violaceus Singer,
× 5-11 µm, cylindrical to clavate or irregular in described from Bolivia (Singer, 1976),
outline, hyaline, inamyloid, thin-walled; apical although we are hesitant to accept Southeast
Asian populations as representing a New
setulae 2-5(-14) × 1(-2) µm, cylindrical,
World taxon without data to support vicariance
subacute to acute, yellow, thin- to thick-walled;
or recent dispersal.
b) non-setulose cells like the pleurocystidia,
A bipolar mating system was reported for
38-74× 8-10 µm, fusoid to clavate, usually
this form of M. haematocephalus by Tan et al.
attenuated and strangulate at the apex, hyaline,
(2007) based on material from Malaysia.
295
54F. Marasmius haematocephalus (Mont.) Fr. gelatinous. Stipitipellis hyphae 3-7 µm diam.,
f. haematocephalus (Figs 68, 69-5) parallel, cylindrical, brown to greenish brown,
Pileus 3-10(-14) mm diam., obtusely smooth, dextrinoid, thin- to thick-walled (up to
conical when young, convex with or without a 1 µm), non-gelatinous. Stipe trama hyphae 3-
rugulose umbo in age, striate to sulcate, 5(-7) µm diam., parallel, cylindrical, hyaline,
pruinose, dull, disc deep reddish brown to smooth, dextrinoid, thin-walled, non-gelatinous.
maroon, margin greyish red (11C6) to violet Caulocystidia absent. Clamp connections
brown (11D7). Context pinkish cream, thin. present.
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.
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,
Basidia 23-25 × 5-6 µm, cylindrical to clavate,
4-spored. Basidioles fusoid to clavate.
Cheilocystidia composed of 2 types of cells: a)
5-11 µm, cylindrical to clavate, or pyriform,
often branched, hyaline, inamyloid, thin-walled;
apical setulae 1-4(-5) × 1 µm, cylindrical, Fig. 68. Marasmius haematocephalus f. haematocephalus
subacute, yellow to brownish yellow, thin- to (N. Wannathes 428). 1. Basidiomes 2. Basidiospores 3a.
Siccus-type cheilocystidia 3b. Non-setulose
thick-walled; b) non-setulose cells like the
pleurocystidia, 25-41 × 7-12 µm, fusoid to = 10 mm, 2-5 = 10 µm
clavate, usually attenuated and strangulate at
the apex, often with 2-4 successive apical Habit, habitat and known distribution.
constrictions, hyaline inamyloid, thin-walled. Scattered to gregarious on dicotyledonous
Pleurocystidia common, 32-54 × 6-12(-15) µm, leaves, Africa, neotropics, Java, Malaysia,
clavate to fusoid, usually attenuated and Papua New Guinea, Sri Lanka and Thailand.
strangulate at the apex, sometimes with 2 Material examined: Thailand. Chiang Mai
successives apical constrictions, hyaline, Waterfall, on Hwy 1095, N19º 6.5’.E 98º 46.3’ alt. 1014
inamyloid, thin-walled. Pileipellis hymeniform, m., 29 June 2004, Y.S. Tan 277 (CMU, SFSU); Chiang
mottled, composed of Siccus-type broom cells; Rai Province, Muang District, Horticulture Research
main body (6-)8-20 × 5-12 µm, clavate to Center, 12 June 2005, N. Wannathes 296 (CMU, SFSU);
broadly clavate or pyriform, sometimes Chiang Mai Province, Doi Suthep-Pui National Park,
Medicinal Plant Garden, 16 June 2005, N. Wannathes
branched, yellow to hyaline, inamyloid, thin- to 316 (CMU, SFSU); Phrae Province, Muang District,
thick-walled; apical setulae 2-5 × 1(-1.5) µm, Suen Keun, Na Koo Haa Waterfall, 15 August 2005, N.
cylindrical, subacute, brown to dark brown, Wannathes 409 (CMU, SFSU); Phrae Province, Muang
thick-walled. Pileus trama interwoven, weakly District, Cherng Thong Waterfall, 16 August 2005, N.
Wannathes 418 (CMU, SFSU), same location, 19
dextrinoid to dextrinoid. Lamellar trama August2005, N. Wannathes 434 (CMU, SFSU); Phrae
hyphae 3-7(-12) µm diam., interwoven, Province, Muang District, Pa Dang, Nong Kam Village,
cylindrical to inflated, smooth, hyaline, weakly near Nong Kam reservoir, 18 August 2005, N.
dextrinoid to dextrinoid, thin-walled, non- Wannathes 428 (CMU, SFSU).
296
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 pleurocystidia, and an absence of caulocystidia.

commonly reported pantropical species, is Singer (1976) recognized nine distinct varieties
characterized by a small (3-14 mm diam.), of this species occurring in the New World
obtusely conical to convex, striate to sulcate tropics distinguished mainly by pileus
pileus coloured deep reddish brown to maroon, pigmentation. As indicated above, we can
distant (9-14), narrow, pink or purplish cream distinguish six distinct forms in Thailand, some
lamellae, a wiry, non-insititious, glabrous stipe of which are nearly indistinguishable from the
with light violet to purple apex and brown varieties reported by Singer (1976) from the
base, clavate basidiospores with mean range neotropics. Until molecular data from
16.8-21.9 × 4.0-4.3 µm, numerous strangulate neotropical populations are compared with
297
M. haematocephalus f. haematocephalus NW409

99
M. haematocephalus f. haematocephalusTYS277
93 M. haematocephalus f. haematocephalus NW434
M. haematocephalus “f. atrobrunneus” NW117
M. haematocephalus “f. violaceus” NW339

98
M. haematocephalus “f. violaceus” W193
M. haematocephalus “f. violaceus” NW413
83 M. haematocephalus “f.variabilis” W430
52 M. haematocephalus “f. luteocephalus” NW310
M. haematocephalus “f. robustus” NW330
M. haematocephalus “f. robustus” NW433
M. siccus DED 255

5 changes
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 and were sister taxa with 83% BS support;
designate the Thai material with Singer's whereas “f. robustus” was unresolved.
epithets.
In the ML tree, 13 sequences of the 55. Marasmius hypophaeus Berk & M.A.
macromorphologically distinct yet micromor- Curtis, J. Linn. Soc., Bot. 10: 298. 1868.
phologically indistinguishable M. haemato- (Figs 71, 75-1)
cephalus formed a clade (AE) with strong Pileus 2-8 mm diam., paraboloid to
support (1.0 PP, 99% BS) and some internal convex, disc rugulose, striate, pruinose, dull,
phylogenetic structure. A bootstrap branch and brown (7D8) to ferruginous when young,
bound analysis of 2000 replicates of a dataset greyish orange (6B4) in age. Context cream,
including all 13 M. haematocephalus thin. Lamellae free to adnexed, distant (11-13),
sequences with M. siccus as outgroup (Fig. 70), narrow, cream with brownish orange edges,
recognized most of the macromorphological non-intervenose. Stipe 23-35 × < 0.5 mm,
entities as distinct clades. Representative central, cylindrical, wiry, glabrous, insititious
sequences conforming to forma haemato- to non-insititious, apex yellowish white, base
cephalus formed a clade with 99% BS support; dark brown. Odor and taste not distinctive.
those of “f. violaceus” formed a clade with Basidiospores 16-18 × 4 µm [x = 17.2 ±
98% BS support; “f. variabilis” and “f. 0.7 × 4.0 ± 0 µm, Q = 4.0-4.5, Qm = 4.2, n = 25
luteocephalus” showed invariable sequences spores, s = 1 specimen], narrowly ellipsoid to
298
Fungal Diversity
hymeniform, weakly mottled, composed of

Siccus-type broom cells; main body 11-19 × 7-
9 µm, cylindrical to clavate, pale yellow,
inamyloid, thin-walled; apical setulae 2-5 × 1-
1.5 µm, crowded, cylindrical, often wavy,
subacute, brownish yellow to light brown, thin-
to thick-walled. Pileus trama interwoven,
dextrinoid. Lamellar trama hyphae (3-)4-7(-10)
µm diam., interwoven, cylindrical to inflated,
subparallel, cylindrical, yellowish brown to
light brown, smooth, weakly dextrinoid, thin-
walled, non-gelatinous. Stipe trama hyphae 3-7
weak dextrinoid, thin-walled, non-gelatinous.
Scattered to gregarious on monocotyledonous
or dicotyledonus leaves, Cuba, Java and
Northern Thailand.
Province, Khun Chae National Park, N 19º 4.4' E 99º
23.5' alt. 963 m., 10 June 2005, N. Wannathes 285
(CMU, SFSU)
Discussion: Marasmius hypophaeus is
characterized by a small, striate pileus coloured
ferruginous to greyish orange, distant lamellae,
colored cream lamellae with brownish orange
Fig. 71. Marasmius hypophaeus (N. Wannathes 285). 1. edges, basidiospores with mean 17.2 × 4.0 µm,
Basidiomes 2. Basidiospores 3. Cheilocystidia 4a. Non- and growth on both moncotyledonous and
setulose cheilocystidia 4b. Siccus-type pleurocystidia 5. dicotyledonous leaves. Described originally
from Cuba, this is the second report of the
species from the Old World tropics. Desjardin
fusoid, smooth, hyaline, inamyloid, thin-walled.
et al. (2000) reported the species from Java.
Basidia not observed. Basidioles cylindrical to
clavate. Cheilocystidia common, of Siccus-type
56. Marasmius aff. pallescens Murrill, N.
broom cells; main body 10-15 × 6-8 µm, Amer. Fl. 9(4): 261. 1915. (Figs 72, 75-2)
cylindrical to clavate, hyaline, inamyloid, thin- Pileus 4-15 mm diam., obtusely conical
walled; apical setulae 5-11 × 1(-3) µm, to convex, sometimes with a small papilla
cylindrical, wavy, often forked, conical to when young, broadly conical with rugulose
subacute, yellow to brownish yellow, thin- to disc in age, sulcate, pruinose, dull, light brown
thick-walled. Pleurocystidia composed of 2 (7D5-6) to pale greyish brown (7D3-4) when
type of cells: a) rarely present, Siccus broom young, grey to greyish orange (6B-C3) in age.
cells with main body 13-15 × 4-6 µm, Context light brownish cream, thin. Lamellae
cylindrical, hyaline, inamyloid, thin-walled; adnexed, subdistant (12-16), broad (1-3 mm),
apical setulae 3-5 × 1 µm, cylindrical, conical light brownish cream, non-marginate, non-
to subacute, hyaline, thick-walled; b) non- intervenose. Stipe 20-50 × 0.5-0.8 mm, central,
setulose cells, common, 41-48 × 7-9 µm, cylindrical, wiry, glabrous, non-insititious,
cylindrical to clavate, strangulate at the apex, apex yellowish white, base dark brown. Odor
hyaline, inamyloid, thin-walled. Pileipellis and taste not distinctive.
299
µm diam., interwoven, cylindrical to inflated,
parallel, cylindrical, brown, smooth, dextrinoid,
thick-walled (up to 1µm), non-gelatinous. Stipe
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.
57. Marasmius bondoi Wannathes, Desjardin
Pleurocystidia 5. Pileipellis, Scale bar 1 = 20 mm, 2-5 = & Lumyong, sp. nov. (Figs 73, 75-3)
10 µm MycoBank: MB512430
Basidiospores (16-)17-18(-19) × 4-5 µm Etymology: ‘bondoi’ (Thai) = on the mountain;
referring to the habitat where the holotype specimen was
[x = 17.5 ± 0.7 × 4.1 ± 0.3 µm, Q = 3.6-4.8, Qm found.
= 4.3, n = 25 spores per 1 specimen], narrowly Pileus 12-32 mm diametro, convexus usque
ellipsoid to fusoid, smooth, hyaline, inamyloid, plano-convexus, umbonatus vel subumbonatus, striatus
thin-walled. Basidia not observed. Basidioles usque sulcatus, pruinosus, hebetatus, disco atrobrunneo
cylindrical to clavate. Cheilocystidia common, of usque brunneo, margine flavo-brunneo, hygrophanus,
ubique pallide brunneo-flavo post arescendum.
Siccus-type broom cells; main body 6-18 × 5- Contextus flavo-brunneus, tenuis. Lamellae annexae
10 µm, cylindrical to clavate, hyaline, usque liberae, distantes usque subdistantes (10-20) cum
inamyloid, thin- to thick-walled; apical setulae 2-4 seriebus lamellularum, latae (2-4 mm), cinereo-
3-6 × 1 µm, cylindrical, conical to subacute, aurantiacae, haud marginatae, haud intervenosae. Stipes
yellow to pale yellow, thick-walled. Pleuro- 32-92 × 1-2 mm, centralis, cylindratus, filo metallico
similis, cavus, glaber, haud insititius, apice cinereo-flavo,
cystidia common, 46-53 × 10-15 µm, basi rubro-brunnea usque violaceo-brunnea. Odor suco
cylindrical to fusoid, constricted at the apex, cocois similis. Sapor non proprius. Basidiosporae 13-
hyaline, inamyloid, thin-walled. Pileipellis 18(-20) × 3-5 µm, anguste ellipsoideae usque fusoideae,
hymeniform, mottled, composed of Siccus-type oblique curvae, laeves, hyalinae, inamyloideae,
tenuitunicatae Basidia 25-29 × 7-9 µm, cylindrata usque
broom cells; main body 9-17 × 6-8 µm,
clavata, 4-spora. Basidiolae fusoideae usque clavatae.
cylindrical to clavate, hyaline, inamyloid, thin- Cheilocystidia abundantia, typi Sicci; (7-)12-21 × (3-)5-
walled; apical setulae 3-5 × 1 µm, crowded, 10 µm, cylindrata usque clavata, hyalina, inamyloidea,
cylindrical, subacute, brownish yellow to light tenuitunicata; setulis apicalibus 3-15(-21) × 1-2 µm,
brown, thick-walled. Pileus trama interwoven, cylindratis usque conicis, interdum lobatis, subacutis,
dextrinoid. Lamellar trama hyphae 3-8(-10) 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,
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,
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).
Fig. 73. Marasmius bondoi (N. Wannathes 237). 1.
convex, umbonate or subumbonate, striate to Basidiomes 2. Basidia 3. Basidiospores 4. Cheilocystidia 5.
sulcate, pruinose, dull, disc dark brown to Pleurocystidia 6. Pileipellis, Scale bar 1 = 20 mm, 2-6 =
brown, margin yellowish brown, hygropha- 10 µm
nous, fading to pale brownish yellow overall
with moisture loss. Context yellowish brown µm, cylindrical to fusoid, constricted and
(5D5), thin. Lamellae adnexed to free, distant sometimes lobed at the apex, hyaline,
to subdistant (10-20) with 2-4 series of inamyloid, thin-walled. Pileipellis hymeniform,
lamellulae, broad (2-4 mm), greyish orange, mottled, composed of Siccus-type broom cells;
non-marginate, non-intervenose. Stipe 32-92 × main body (8-) 13-20(-25) × 6-10 µm,
1-2 mm, central, cylindrical, wiry, hollow, cylindrical to clavate or pyriform, hyaline to
glabrous, non-insititious, apex greyish yellow pale yellow, inamyloid, thin- to thick-walled;
(4B4), base reddish brown (8E8) to violet apical setulae 4-10(-15) × 1-1.5 µm, crowded,
brown (10F8). Odor of coconut juice. Taste not cylindrical, subacute, yellowish brown to
distinctive. brown, thick-walled. Pileus trama hyphae 3-8
Basidiospores 13-18(-20) × 3-5 µm [xmr µm diam., interwoven, cylindrical to inflated,
= 14.2-18.3 × 3.5-4.6 µm, xmm = 16.4 ± 1.3 × smooth, hyaline, strongly dextrinoid, thin-
3.9 ± 0.3 µm, Qmr = 3.5-4.9, Qmm = 4.2 ± 0.5, n = walled. Lamellar trama hyphae 3-8(-13) µm
25 spores, s = 10 specimens], narrowly ellipsoid diam., interwoven to regular, cylindrical to
to fusoid, curved in profile, smooth, hyaline, inflated, smooth, hyaline, dextrinoid, thin-
inamyloid, thin-walled. Basidia 25-29 × 7-9 µm, walled, non-gelatinous. Stipitipellis hyphae 3-
cylindrical to clavate, 4-spored. Basidioles fusoid 6(-9) µm diam., subparallel, cylindrical,
to clavate. Cheilocystidia abundant, of Siccus- yellowish brown to brown, smooth, weakly
type broom cells; main body (7-)12-21 × (3-)5- dextrinoid, thick-walled (up to 2 µm), non-
10 µm, cylindrical to clavate, hyaline, gelatinous. Stipe trama hyphae 4-13(-19) µm
inamyloid, thin-walled; apical setulae 3-15(-21) diam., parallel, cylindrical, hyaline, smooth,
× 1-2 µm, cylindrical to conical, sometimes dextrinoid, thin-walled, non-gelatinous.
lobed, subacute, yellow to pale yellow, thin- Caulocystidia absent. Clamp connections
walled. Pleurocystidia common, 40-58 × 7-15
301
M. bondoi NW237
98 M. bondoi NW390
M. bondoi NW386
63
M. bondoi NW011
55 M. bondoi NW320
M. bondoi NW384
74
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. were recognized that differed in lamellar
Gregarious to densely gregarious, typically in spacing, albeit with low BS support, viz.,
cespitose clusters on wood or on specimens with distant (10-15) lamellae
dicotyledonous leaves, Northern Thailand. [NW011, 237, 386 and 390] and specimens
Material examined: Thailand. Chiang Mai with subdistant (13-20) lamellae [NW320, 384
Province, Doi Suthep-Pui National Park, Sangasabhasri and 399]. All other taxonomically significant
Land on the way to Huai Kok Ma Village, N 18º 48.4’ E
micro- and macromorphological features of all
(CMU, SFSU); same location, 6 June 2004, Y.S. Tan M. bondoi specimens are overlapping and we
318 (CMU, SFSU); same location, 24 July 2004, N. recognize them as representing a single species
Wannathes 237 (CMU, SFSU); same location, 24 June with variable lamellar spacing.
2005,N. Wannathes 320 (CMU, SFSU); Chiang Mai Marasmius bondoi is most phenetically
Province, Doi Suthep-Pui National Park, trial opposite
with the way to Huai Kok Ma Village, 2 July 2005, N.
similar to two African species, M. grandisetulosus
Wannathes 384 (CMU, SFSU); same location, 2 July Singer and M. tenuisetulosus (Singer) Singer.
2005, N. Wannathes 386 (CMU: holotype; SFSU: Both of the latter species are reported to have
isotype), N. Wannathes 388, 389 and 390(CMU, SFSU); distinctly striped pilei with darker sulcae
Chiang Mai Province, Doi Suthep-Pui National Park, (brown or rusty-tawny) and paler ridges
Medicinal Plant Garden, 3 August 2005, N. Wannathes
399 (CMU, SFSU).
(yellowish grey to orange or tawny). The Thai
Discussion: Marasmius bondoi is M. bondoi has an evenly pigmented pileus that
characterized by a convex, subumbonate, is never striped.
sulcate pileus with brown and yellowish brown In the ML tree, clade AB contained seven
tones, distant to subdistant (10-20) lamellae sequences of M. bondoi with M. graminipes on
with or without brown edges, basidiospores a relatively long branch embedded within the
with means in the range 14.2-18.3 × 3.5-4.6 µm, clade as sister to one sample (NW 390) but
with low support (0.52 PP, 53% BS). In a
Siccus-type broom cells with setulae 2-15 × 1-2
branch and bound analysis with 2000 replicates
µm, fusoid-constricted pleurocystidia, and
and M. plicatulus as outgroup (Fig. 74), the
growth on wood or dicot leaf substrates. In a
seven M. bondoi sequences cluster with low BS
separate branch and bound analysis involving
support (55%) and the morphologically distinct
only M. bondoi and M. graminipes with M.
M. graminipes is sister to the M. bondoi clade.
plicatulus as outgroup (Fig. 74), several clades
302
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 (1976, 1986) and Antonín and Noordeloos

(1993) subdivided sect. Sicci into four series
In most cases, the ITS sequence data (viz., Atrorubentes, Haematocephali, Leonini,
supported recognition of morphospecies with Spinulosi) based on the presence or absence of
high statistical support. For example, three pleurocystidia, setae or simple cylindrical
specimens of the new species M. jasminodorus caulocystidia. None of these series are
(in clade AA, Fig. 1) that were collected in monophyletic in our analyses. There was very
diverse geographical sites (different mountains at limited statistical support for the deeper nodes
different elevations and forest types in different in the ML or MP analyses (Fig. 1) suggesting
provinces) formed a clade with 0.98 PP and that the ITS region may be of limited value in
100% BS support and showed almost no clarifying the infrageneric delimitation of
genetic variability. Marasmius. Before any formal reorganization
Singer (1976) recognized Marasmius as of taxa within Marasmius is proposed, further
representing 11 sections. Based on nLSU rDNA phylogenetic analyses based on a global sample
sequences data, Wilson and Desjardin (2005) set and additional genes (LSU, rpb2, ef1-α)
restricted the genus Marasmius to only seven must be conducted.
sections (viz. Globulares, Hygrometrici, Leveil- Although the ITS data alone are of
leani, Marasmius, Neosessiles, Scotophysini, limited use in elucidating infrageneric
Sicci) and recognized the other four Singerian phylogenetic structure, they are very useful in
sections (viz., Alliacei, Androsacei, Epiphylli, aiding taxonomic delimitations at the species
Fusicystides) as belonging to other genera. rank. In most cases, species delimitations based
Only five of the sections in Marasmius sensu on morphological characters are concordant
stricto were encountered in northern Thailand with those suggested by ITS sequence
(Hygrometrici and Scotophysini excluded) and similarity. And this pattern holds true for
of these, only sect. Marasmius subsect. geographically distant and ecologically distinct
Marasmius was monophyletic based on the ITS populations [cf., M. jasminodorus (.98 PP,
dataset (Fig. 1; clade D). In addition, Singer 100% BS), M. brunneoolivascens (.99 PP, 92%
303
BS), M. bambusiniformis (1.0 PP, 100% BS), The presence or absence of pleuro-
M. laticlavatus (1.0 PP, 100% BS), M. cafeyen cystidia, setae or simple caulocystidia have
(1.0 PP, 100% BS)]. been used to distinguish series with sect. Sicci,
Historically, morphological characters although these infrageneric groups are not
have been used in combination to delimit monophyletic based on the ITS data
sections (collariate lamellae, insititious stipe, Likewise, basidiospore size is a
pileipellis anatomy), series (pleurocystidia, taxonomically significant feature to aid in
setae or simple caulocystidia), and species delimiting species, but is of limited use at
(basidiospore size, broom cell setulae, cystidia higher taxonomic levels. For example, M.
shape). The ITS data support the use of these pellucidus, with xmm = 7.2 × 3.3 µm, belongs to
morphological characters to aid in delimiting a subclade within clade AA (with .77 PP, 78%
taxa at various taxonomic ranks, albeit not BS) and is distinct from the other members (M.
necessarily as circumscribed by Singer (1976, araucariae, M. jasminodorus, M. inthanonensis)
1986), Antonín and Noordeloos (1993), who have basidiospore means in the range
Desjardin and Horak (1997), and Desjardin et 10.1-10.4 × 3.6-3.9 µm. Alternatively, shared
al. (2000). For example, all taxa that lack basidiospore size may be a clade-defining
collariate lamellae belong in clades A and F feature. Clade AC, with two setoid (SS) species
(Fig. 1) and are phylogenetically distinct from (M. nummularius, M. trichotus) and two non-
taxa with collariate lamellae (all other clades). setoid species with simple caulocystidia (SA;
However, all collariate species (sect. M. iras, M. xestocephalus) all share narrowly
Marasmius) do not form a monophyletic group. fusoid basidiospores with means in the range
In addition, all species with non-insititious 12.1-13.2 × 3.7-4.1 µm.
stipes belong to the well-supported clade A Combinations of these micro-characters
(sects. Sicci and Globulares), whereas those and ITS data can be used to define clades. For
with insititious stipes belong in the other clades. example, four species within clade AD with 1.0
Moreover, all tested species with a unifactorial PP and 100% BS are circumscribed by
(bipolar) mating system belong in clade A, Globulares-type pileipellis cells, long-clavate
whereas those with a bifactorial (tetrapolar) basidiospores with means in the range 22.8-
mating system belong to the clades with 30.0 × 4.4-5.6 µm, and an absence of
collariate lamellae and insititious stipes. Based pleurocystidia and caulocystidia. The ITS data
on these data, clade A, representing all are a tremendous help in delimiting the
included species of sects. Sicci and Globulares pleurocystidiate species M. haematocephalus
can be distinguished from all other Marasmius (clade AE with 1.0 PP, 99% BS) from other
by the presence of a non-insititious stipe, taxa, and in distinguishing its various forms
bipolar mating system and absence of collariate whose pilei range in color from yellowish
lamellae. A pileipellis formed from Siccus-type orange to brown, olive, greyish blue, red and
broom cells versus Globulares-type cells (non- violet. Micromorphologically all isolates of this
setulose) that currently differentiates sect. Sicci clade are indistinguishable; however, the
from sect. Globulares is not a phylogenetically branch and bound analysis (Fig. 70) clearly
significant feature according to the ITS data. distinguishes five of the six forms into well-
Of the eight species with Globulares-type supported lineages.
pileipellis elements included in the analyses,
five belong to clade AD, one in clade AA, and Conclusions
two are in clade AF. Desjardin and Horak
(1997) suggested that the separation of species The ITS sequence data generally support
into sections Sicci and Globulares based on the recognition of species circumscribed by
pileipellis anatomy was untenable when they macro- and micromorphological characters, and
described M. nexus Desjardin & Horak, a in many cases can be used to delimit clades of
species from Papua New Guinea with a taxa with shared morphological features.
pileipellis formed from equal numbers of However, ITS sequences are of limited use in
Siccus-type cells and Globulares-type cells. recognizing currently delimited infrageneric
304
Fungal Diversity
taxa at the series rank and above. Additional Desjardin, D.E., Retnowati, A. and Horak, E. (2000).
species and genes need to be sequenced and Agaricales of Indonesia: 2. A preliminary
monograph of Marasmius from Java and Bali.
analyzed with multiple algorithms in order to Sydowia 52, 92-193.
clarify the infrageneric delimitation of the Desjardin, D.E., Flegel, T.W. and Boonpratuang, T.
genus Marasmius (2004). Basidiomycetes, in: Jones EBG,
Tanticharoen M, Hyde KD, Thai Fungal
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and Yee Shin Tan for help with specimen collection and Herbarium Specimens, Vilgalys Lab [Online].
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A monograph of Marasmius (Basidiomycota) from Northern Thailand based

Article in Fungal diversity · July 2009

Dennis E Desjardin Nopparat Wannathes

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Kevin David Hyde Brian A. Perry

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A monograph of Marasmius (Basidiomycota) from Northern Thailand based on

Key words: Agaricales, fungi, phylogeny, taxonomy

Introduction present here a monograph of Marasmius from

Materials and methods DNA extraction, Polymerase Chain Reaction

0.96/ 87 M. pellucidus NW321 -G

M. cafeyen NW130 (type) -MS

Taxonomy 10. Stipe arising directly from rhizomorphs.............. 11

brownish yellow, smooth, weakly dextrinoid to

diam.), conical pilei with a dark brown papilla,

differs in forming darker brown pilei, fewer

Section Marasmius, subsect. Sicciformes

5. Marasmius berambutanus Desjardin, Retn.

6. Marasmius purpureisetosus Corner, Nova

Material examined: Thailand. Chiang Mai

7. Marasmius brevicollus Corner, Beih. Nova

basidiospores with mean 14.2 × 3.2 µm, and

cylindrical, wiry, glabrous, insititious, black

Tan 275 (CMU, SFSU); same location, 7 July 2004, N.

14. Marasmius straminiceps Wannathes, broadly clavate or sometimes irregular in

(9-10), non-marginate lamellae; a lack of rhizomorphs, has thick-walled cheilocystidia,

Habit, habitat and known distribution.

Japan, Malaysia, Thailand and Papua New

cylindrical, obtuse to subacute, hyaline, thin-

Thai material matches nicely with Indonesian

31. Marasmius coklatus Desjardin, Retnowati

32. Marasmius araneocephalus Wannathes,

[= Marasmius atrorubens (Berk.) Berk., lamellulae, narrow, cream, non-marginate, non-

35. Marasmius ochroleucus Desjardin & E.

= 12.2 × 4.1 µm vs 9.7 × 3.9 µm µm), and

36. Marasmius luteolus Berk. & M.A. Curtis,

Pleurocystidia absent. Pileipellis hymeniform,

Material examined: Thailand. Chiang Rai

39. Marasmius xestocephalus Singer, Bull.

Pileus 3-14 mm diam., obtusely conical cylindrical to cylindrical or irregular in outline,

47. Marasmius imitarius Wannathes, Desjardin

Koo Haa Waterfall, 15 August 2005, N. Wannathes 410

50. Marasmius ganyao Wannathes, Desjardin

Pileus 6-23 mm diametro, convexus cum umbone

Trama pilei intertexta, hyphis (4-)6-13(-17) µm diametro,

Pileus 4-21 mm diam., convex to plano-

clavate. Cheilocystidia composed of 2 types of

and margin pinkish purple to violet. Context inamyloid, thin-walled. Pleurocystidia

Fig. 69. Basidiomata of forms of Marasmius haematocephalus. 1. M. haematocephalus “f. luteocephalus”

Discussion: Marasmius haematocephalus, a pleurocystidia, and an absence of caulocystidia.

M. haematocephalus f. haematocephalus NW296

93 M. haematocephalus f. haematocephalus NW434

M. haematocephalus f. haematocephalus NW428

M. haematocephalus “f. atrobrunneus” NW117

M. haematocephalus “f. violaceus” NW339

M. haematocephalus “f. violaceus” NW413

83 M. haematocephalus “f.variabilis” W430

52 M. haematocephalus “f. luteocephalus” NW310

M. haematocephalus “f. robustus” NW330

M. haematocephalus “f. robustus” NW433