A monograph of the genera Crinipellis and Moniliophthora from Southeast Asia including a molecular phylogeny of the nrITS region

Abstract

2009). A monograph of the genera Crinipellis and Moniliophthora from Southeast Asia including a molecular phylogeny of the nrITS region. Fungal Diversity 37: 101-152 Nineteen species of Crinipellis (Basidiomycota, Agaricales) and three species of Moniliophthora (Basidiomycota, Agaricales) are described from Southeast Asia. Thirteen relevant type studies along with excluded and insufficiently known taxa are also included. Five new species and one new form of Crinipellis (C. brunnescens, C. cupreostipes, C. dipterocarpi f. cinnamomea, C. furcata, C. malesiana and C. tabtim), one new species of Moniliophthora (M. marginata) and three formal transfers of species described originally as Crinipellis (Moniliophthora canescens, Moniliophthora nigrilineata and Marasmiellus subochraceus) are proposed here. These are the first distribution reports of Crinipellis and Moniliophthora for Thailand, Malaysia and Indonesia and the geographic distributions were expanded for C. actinophora, C. brunneipurpurea, C. dipterocarpi, C. aff. iopus, C. aff. mirabilis, C. setipes and C. trichialis. Phylogenetic analyses of 12 Southeast Asian Crinipellis, three Moniliophthora and six Chaetocalathus species are included using sequence data from the nrDNA ITS region. Comprehensive descriptions for all 36 taxa along with illustrations when available are included in this study.

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Fungal Diversity
101
A monograph of the genera Crinipellis and Moniliophthora from Southeast Asia
including a molecular phylogeny of the nrITS region
Kerekes, J.F. 1* and Desjardin, D.E. 1
1Department of Biology, San Francisco State Univeristy, 1600 Holloway Avenue, San Francisco, CA 94132, USA.
Kerekes, J.F. and Desjardin, D.E. (2009). A monograph of the genera Crinipellis and Moniliophthora from Southeast
Asia including a molecular phylogeny of the nrITS region. Fungal Diversity 37: 101-152
Nineteen species of Crinipellis (Basidiomycota, Agaricales) and three species of Moniliophthora (Basidiomycota,
Agaricales) are described from Southeast Asia. Thirteen relevant type studies along with excluded and insufficiently
known taxa are also included. Five new species and one new form of Crinipellis (C. brunnescens, C. cupreostipes, C.
dipterocarpi f. cinnamomea, C. furcata, C. malesiana and C. tabtim), one new species of Moniliophthora (M. marginata)
and three formal transfers of species described originally as Crinipellis (Moniliophthora canescens, Moniliophthora
nigrilineata and Marasmiellus subochraceus) are proposed here. These are the first distribution reports of Crinipellis
and Moniliophthora for Thailand, Malaysia and Indonesia and the geographic distributions were expanded for
C. actinophora, C. brunneipurpurea, C. dipterocarpi, C. aff. iopus, C. aff. mirabilis, C. setipes and C. trichialis.
Phylogenetic analyses of 12 Southeast Asian Crinipellis, three Moniliophthora and six Chaetocalathus species are
included using sequence data from the nrDNA ITS region. Comprehensive descriptions for all 36 taxa along with
illustrations when available are included in this study.
Keywords: Marasmiaceae, Agaricomycotina, molecular phylogeny, fungal taxonomy.
Article Information
Received 28 November 2008
Accepted 7 March 2009
Published online 1 August 2009
*Corresponding author: Jennifer Kerekes; e-mail: Kerekesj@berkeley.edu
Introduction
The biodiversity in Southeast Asia is
among the world’s richest. The region is home
to 16 endemic vascular plant families
(Takhtajan et al., 1986) and contains three of
the top ten biodiversity hotspots as described
by Meyers (2000). Although the fungi remain
largely understudied in this region, there have
been recent studies of saprotrophic fungi
(Desjardin and Horak, 1999; Desjardin et al.,
2000; Wilson et al., 2004; Wannathes et al.,
2004, 2007; Tan et al., 2007). These fungi play
very important ecological roles in litter
decomposition and nutrient recycling. The
following is a regional monograph of the
basidiomycete genera Crinipellis Pat. and
Moniliophthora H.C. Evans, Stalpers, Samson
& Benny that includes a phylogenetic analysis
of the nrITS region.
The crinipelloid genera Crinipellis,
Moniliophthora and Chaetocalathus Singer
(Basidiomycota, Agaricales) are distinguished
morphologically from the allied genus
Marasmius Fr. by basidiomes with pileipellis
composed of thick-walled, dextrinoid, hair-like
terminal cells. Chaetocalathus differs from the
other two genera by formation of non-stipitate
basidiomes. Taxonomically significant features
useful in distinguishing amongst crinipelloid
species include pileus coloration, presence or
absence of rhizomorphs, substrate type,
cheilocystidia shape, presence or absence of
pleurocystidia, basidiospore shape and size,
and KOH reaction of pileipellis cells.
The genus Crinipellis was established by
N. Patouillard in 1889 with Agaricus stipitarius
Fr. as the type species. The first worldwide
treatment of the genus was published by Singer
(1943) in which he recognized 33 stipitate
species of Crinipellis placed into two sections,
and he created the allied genus Chaetocalathus
for eleven non-stipitate species. In 1976, Singer
reworked the neotropical Crinipellis, recog-
nizing 41 species belonging to four sections. In
Singer’s "The Agaricales in Modern

102
Taxonomy" (1986), he reported 5 infrageneric
groups, viz., sections Crinipellis, Metuloido-
phorae, Excentricinae, Grisentinae and
Iopodinae.
Currently, over 150 epithets have been
published in the genus with 75 species
recognized as being distinct (Kirk et al., 2001).
Patouillard described a number of Crinipellis
species (a few of which have since been
transferred to the genus Chaetocalathus) from
Southeast Asia, primarily from Vietnam. As
mentioned above, Singer also described new
species in, and made new transfers into, the
genus from the region. Recently, E.J.H. Corner
(1996) reported six species of Crinipellis, five
of which were described as new species from
Singapore and peninsular Malaysia. To date,
fourteen species of crinipelloid fungi have been
reported from Southeast Asia, viz., 12 species
of Crinipellis, and two species of the non-
stipitate genus Chaetocalathus. Another two
species of previously reported Crinipellis are
transferred out of the genus in this study;
therefore, leaving 10 previously reported
species of Crinipellis from Southeast Asia:
C. actinophora (Berk. & Broome) Singer, C.
atrobrunnea Pat., C. brunneipurpurea Corner,
C. carecomoeis var. litseae Singer, C. cervino-
alba Corner, C. dipterocarpi Singer,
C. hepatica Corner, C. pseudostipitaria subsp.
orientalis Singer, C. sepiaria Pat. & Demange,
and C. trichialis (Lév.) Pat. ex Antonín et al.
To date, no monographic treatment or
phylogenetic analysis of Southeast Asian
Crinipellis has been published.
In addition to the reports of Crinipellis
from Southeast Asia, there are a few reports of
Crinipellis from the Old World (excluding
Europe). In 1968 Singer described one new
combination and two new species of Crinipellis
from Africa (Singer in Pegler, 1968).
Stevenson (1964) reported eight species of
Crinipellis from New Zealand, and Pegler
(1977) reported 5 East African species of
Crinipellis, and another three species of
Crinipellis from Sri Lanka (Pegler, 1986). In
2000, Takahashi described three new species of
Crinipellis from Japan. In this monograph, the
taxonomic position of a few of the previously
reported species of Crinipellis mentioned here
has been revised.
Crinipellis has a worldwide distribution
with the greatest diversity in the tropics. Singer
(1943) noted an interesting distribution of
Crinipellis with the greatest concentration in
the Neotropics as compared to the Old World
tropics, as seen in many other genera. This
observation may be a result of the paucity of
saprotrophic fungal collections and research in
the Old World tropics.
The genus has been recognized
historically as containing both saprotrophic and
parasitic species. Recently, the economically
and ecologically important cacao pathogen
Crinipellis perniciosa (Stahel) Singer, also
known as the witches’ broom disease, was
transferred to the genus Moniliophthora (as
Moniliophthora perniciosa (Stahel) Aime and
Phillips-Mora, 2005). The majority of species
are saprotrophic; however, there are a few
reports of parasitic species including C.
siparunae Singer, C. pseudostipitaria and the
type of the genus, C. stipitaria (Fr.) Pat. (= C.
scabella (Alb. & Schwein.) Murrill), which has
been observed to be parasitic on grasses and
cereal plants (Singer, 1943).
Crinipellis species have been included in
a few molecular studies; however, there has not
been a comprehensive molecular study of the
genus. These studies indicate potential sister
relationships and taxonomic placement of
Crinipellis. Recent molecular sequence data by
Moncalvo et al. (2002), Bodensteiner et al.
(2004), Wilson and Desjardin (2005) and
Matheny et al. (2006) indicate that Crinipellis
and Chaetocalathus are sister to Marasmius
sensu stricto. However, an analysis by Aime &
Phillips-Mora (2005) resulted in Crinipellis as
sister to a clade containing Chaetocalathus,
Moniliophthora and Marasmius sensu stricto.
The goal of this research was to docu-
ment the diversity, distribution and ecology of
Crinipellis and Moniliophthora in Southeast
Asia. In addition, a molecular analysis of the
ITS1-5.8S-ITS2 nrDNA region was included in
order to determine species groups and to help
delimit morphologically similar taxa.

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Materials and Methods
Morphological studies
The geographic scope of the project,
Southeast Asia, is circumscribed by
Takhtajan’s et al. (1986) Indomalesian
Botanical Subkingdom with a focus on the
Indo-Chinese Region and the Malesian Region,
including Myanmar, Thailand, Cambodia, Laos,
Vietnam, Malaysia, the Philippines, New
Guinea and Indonesia. Four collecting trips to
Thailand, Malaysia and Indonesia in January
and June/July of 2004 and 2005, yielded 82
specimens of Crinipellis, three specimens of
Chaetocalathus and two specimens of
Moniliophthora. Seventy-eight collections
were borrowed from other herbaria, including
type specimens. Voucher material and
duplicates are deposited in the Harry D. Thiers
Herbarium (SFSU), Herbarium Bogoriense
(BO) in Bogor, Java, Indonesia, the Mycology
Lab, Institute Biology of Science, Faculty of
Science, University of Malaya (KLU-M) in
Malaysia and Chiang Mai University (CMU) in
Chiang Mai, Thailand. In the field, appropriate
macromorphological characters were docu-
mented, including shape, size, color and
surface features of the pileus and stipe, context
and lamellae features, presence or absence of
rhizomorphs, and substrate type. Color terms
and notations in parentheses are from Kornerup
and Wanscher (1978). Micromorphological
characters were described from collected
specimens and specimens borrowed from other
herbaria. These characters include size, shape,
color and reaction to Melzer's reagent of the
basidiospores, cheilocystidia, pleurocystidia,
pileipellis, stipitipellis and trama. Microscopic
terms are those of Largent (1986), Largent et al.
(1977) and Vellinga (1988). The pileipellis of
Crinipellis species is formed from a cutis of
terminal skeletal hyphae, herein called hairs,
arising from chains of thinner-walled
generative hyphae herein called subtending
hyphae. All measurements and colors reported
for microscopic features were made from dried
material rehydrated in 100% ethanol and
followed by distilled water, 3% KOH, Congo
Red or Melzer’s reagent. An Olympus CX41
microscope with a drawing tube was used for
microscopic analysis. Basidiospore statistics
include: xm, the arithmetic mean of the spore
length and the spore width (± standard
deviation) from n spores measured in a single
specimen; xmr, the range of spore means and
xmm, the mean of spore means (± SD) where
more than one specimen is available; Q, the
quotient of spore length and spore width in any
one basidiospore indicated as a range of
variation in n spores measured; Qm, the mean
of Q-values in a single specimen; Qmr, the
range of Q values and Qmm, the mean of Q
values where more than one specimen is
available. Herbarium citations follow
Holmgren et al. (1981) with the exception of
the Mycology Lab, Faculty of Science,
University of Malaya (KLU-M) in Kuala
Lumpur, Malaysia.
In the figures, basidiomes are natural size,
unless stated otherwise. Figures for
basidiospores, basidia, basidioles, cystidia,
pileipellis hairs, and caulocystidia are at 1000x
(scale bar = 10 μm), unless stated otherwise.
Molecular techniques
DNA was extracted from recently
collected dried specimens and herbarium
collections using either the E.Z.N.A. Forensic
DNA Kit (D3591-01) or Fungal Mini Kit
(D3390-01) from Omega Bio-tek, Inc. (P.O.
Box 47310, Doraville, GA 30362). Primers
used for DNA amplification and sequencing
were ITSI, ITS2, ITS3, ITS4 (White et al.,
1990) and ITS1-F, ITS4-B (Gardes and Bruns,
1993). PCR and cycle sequencing reactions
followed standard protocols and were
performed on ABI 9600 thermal cyclers.
Sequencing reactions were run on an ABI 377
and ABI 3100 Genetic Analyzer (Applied
Biosystems, Foster City, CA, USA). Sequences
were edited and contigs assembled using
Sequencher version 3.1.1 (Gene Codes
Corporation, Ann Arbor, Michigan) software.
Phylogenetic analysis
Alignment was performed with ClustalX
(1.81) (Thompson et al., 1997) and by hand
using MacClade 4.03PPC (Maddison and
Maddison, 2001). Maximum parsimony (MP)
and Maximum Liklihood (ML) analyses were
performed with PAUP* v. 4.0b10 (PPC)
(Swofford, 2002) with equal weighting and
gaps treated as missing data. Seventy-one
sequences are included in the data set,

104
including nine previously published sequences
downloaded from GenBank. In addition to the
Crinipellis species, Chaetocalathus, Moni-
liophthora and Marasmius sensu stricto are
also included, with a Marasmius crinis-equi F.
Muell. ex Kalchbr. specimen from Malaysia as
the outgroup. A Maximum parsimony heuristic
search was performed with stepwise addition as
random with 100 replicates and TBR branch
swapping. A bootstrap analysis was performed
with 500 replicates with stepwise addition as
random with 10 replicates, keeping no more
than 100 trees per replicate and TBR branch
swapping. Modeltest 3.06 (Posada and
Crandall, 1998) was used to identify an optimal
model of evolution for the ML analysis. Akaike
Information Criterion (AIC) Model selected
TVM+I+G model, a sub-model of the GTR
model. A ML bootstrap analysis was
performed with 500 replications in GARLI
(Zwickl, 2006). Bayesian analyses were
performed using MrBayes 3.1.2 (Huelsenbeck
and Ronquist, 2001; Ronquist and Hulsenbeck,
2003). MrModeltest 2.2 (Nylander, 2004) was
used to determine the best evolutionary model
(GTR + I + G) following the AIC criterion.
Two independent analyses of 1 000 000
generations were run keeping one tree every
1000 generations. The burn-in value was set to
25% and the remaining trees were used to
calculate a 50% majority rule consensus tree
and to determine the posterior probabilities
(PP).
Sequences generated as part of this
project are listed in Table 1 and were deposited
in GenBank (accession numbers FJ167608-
FJ167669). The alignment was submitted to
TreeBase (SN4057). MycoBank numbers are
provided for all new species (MB 512271 –
MB 512277).
Results
Phylogenetic analyses
The final data set was 907 base pairs in
length. Ambiguously aligned regions in the
alignment (bp 120-200 in ITS1 and 620-800 in
ITS2) were excluded from the analysis. In the
final data set, 657 characters were included
with 196 parsimony informative characters and
338 constant characters. The MP analysis
resulted in 6,600 most equally parsimonious
trees of 752 steps. A phylogram tree produced
by the ML analysis (-lnL = 4516.5117) is
shown in Fig. 1. Both the MP and ML analyses
resulted in the same topology and relationships
for the species groups (except for C.
brunnescens, C. brunneipurpurea, C. aff. iopus
and the clade containing C. piceae and C.
setipes). The 50% majority rule consensus tree
resulting from the Bayesian analysis is also
similar in overall topology although it did not
resolve all of the species groups. There is no
significant bootstrap (BS) or posterior
probability (PP) support for the Crinipellis
clade (excluding C. aff. iopus) in the MP or
Bayesian analyses, however, there was 66% BS
support in the ML analysis. Four
Moniliophthora species formed a clade but
with limited statistical support (53% ML-
BS, .50 PP). Moniliophthora was sister of
Crinipellis aff. iopus but with limited statistical
support (56% ML-BS, .50 PP). Seven species
of Chaetocalathus formed a clade but with
limited statistical support (60% MP-BS, .60 PP)
and were sister to Crinipellis + Moniliophthora
but without statistical support. The maximum
parsimony analysis resulted in Moniliophthora
sister to Crinipellis + Chaetocalathus. The ITS
region resolved species groups with high
statistical support. MP boostrap values of ≥
92%, ML bootstrap values of ≥ 82%, and PP
values of ≥ .97 delimited species groups, with
the exception of C. tabtim (72% MP-BS, 67%
ML-BS, .85 PP). The type species of
Crinipellis, C. scabella, was sister of C.
trichialis with 72% MP-BS, 78% ML-BS
and .99 PP support. Twelve species of
Southeast Asian Crinipellis were resolved in
the molecular analysis although the
relationships amongst them usually received
low statistical support.
Discussion of phylogenetic data
The nrITS region successfully resolved
species groups and aided in distinguishing
morphologically similar taxa. However, due to
the high variability of the region, the ITS
sequence data were not able to resolve the
nodes delimiting crinipelloid genera nor were
they able to resolve the phylogenetic
relationships between species with significant
statistical support. Other analyses using

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Fig. 1. Maximum likelihood tree (-lnL = 4516.5117) inferred from nrITS sequence data. Numbers indicate parsimony
bootstrap values ≥ 50% followed by ML bootstrap values and with posterior probabilities below the branch.

106
Table 1. List of specimens used in the molecular analyses.
Species Origin Collection number
(herbarium)* GenBank accession number
Crinipellis actinophora Malaysia DED7519 (KLU-M) FJ167624
C. actinophora Malaysia JFK78 (KLU-M) FJ167617
C. actinophora Malaysia JFK80 (KLU-M) FJ167618
C. actinophora Malaysia JFK85 (KLU-M) FJ167622
C. actinophora Malaysia JFK86 (KLU-M) FJ167620
C. actinophora Malaysia JFK91 (KLU-M) FJ167625
C. actinophora Malaysia JFK93 (KLU-M) FJ167619
C. actinophora Malaysia JFK99 (KLU-M) FJ167621
C. actinophora Malaysia JFK100 (KLU-M) FJ167623
C. actinophora Malaysia JFK139 (CMU) FJ167626
C. brunneipurpurea Indonesia JFK84 (KLU-M) FJ167646
C. brunneipurpurea Indonesia JFK107 (KLU-M) FJ167645
C. brunnescens Indonesia DED6791 (BO) FJ167627
C cupreostipes Thailand JFK31 (CMU) FJ167641
C cupreostipes Thailand JFK55 (CMU) FJ167640
C cupreostipes Thailand JFK131 (CMU) FJ167642
C. dipterocarpi Thailand DED7570 (SFSU) FJ167656
C. dipterocarpi Thailand DED7602 (SFSU) FJ167651
C. dipterocarpi Thailand JFK126 (CMU) FJ167655
C. dipterocarpi Thailand JFK130 (CMU) FJ167650
C. dipterocarpi Thailand JFK132 (CMU) FJ167647
C. dipterocarpi Indonesia ZT12031 (BO) FJ167654
C. dipterocarpi f.
cinnamomea Malaysia JFK17 (KLU-M) FJ167648
C. dipterocarpi f.
cinnamomea Malaysia JFK20 (KLU-M) FJ167653
C. dipterocarpi f.
cinnamomea Thailand JFK32 (CMU) FJ167649
C. dipterocarpi f.
cinnamomea Indonesia JFK104 (BO) FJ167652
C. furcata Indonesia DED6951 (BO) FJ167658
C. furcata Indonesia JFK103 (BO) FJ167657
C. aff. iopus Papua New Guinea RW829 (GENT) FJ167638
C. aff. iopus Papua New Guinea RW774 (GENT) FJ167636
C. aff. iopus Papua New Guinea AV97/312 (GENT) FJ167637
C. aff. iopus Papua New Guinea AV97/404 (GENT) FJ167639
C. malesiana Indonesia AR491 (BO) FJ167631
C. malesiana Indonesia AR513 (BO) FJ167630
C. malesiana Malaysia KUM084a (KLU-M) FJ167629
C. malesiana Malaysia TYS346 (KLU-M) FJ167628
C. piceae North America DED7622 (SFSU) FJ167632
C. piceae North America DED7758 (SFSU) FJ167633
C. procera New Zealand ZT8490 (ZT) FJ167660
C. setipes Thailand JFK34 (CMU) FJ167634
C. scabella France ZTPAM98 (ZT) FJ167635
C. scabella Germany PB302 AY571033
C. tabtim Thailand JFK129 (CMU) FJ167643
C. tabtim Thailand JFK141 (CMU) FJ167644
C. trichialis Indonesia AWW33 (BO) FJ167608
C. trichialis Indonesia AR216 (BO) FJ167610
C. trichialis Indonesia DED6800 (SFSU) FJ167609
C. trichialis Indonesia DED6924 (SFSU) FJ167613
C. trichialis Indonesia DED7043 (SFSU) FJ167614
C. trichialis Malaysia JFK97 (SFSU) FJ167616
C. trichialis Indonesia JFK108 (BO) FJ167611
C. trichialis Indonesia JFK109 (BO) FJ167612
C. trichialis Indonesia ZT6456 (ZT) FJ167615

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Table 1(continued). List of specimens used in the molecular analyses.
Species Origin Collection number
(herbarium)* GenBank accession number
C. zonata Canada DAOM176761 (SFSU) FJ167659
C. zonata Canada OKM25450 (VPI) AY916692
C. sp. Thailand OKM26890 (VPI) AY916698
C. sp. Guyana MCA1527 (BPI) AY916701
Chaetocalathus craterellus Italy SN223 (ZT) FJ167664
C. fragilis Hawai’i DED6359 (SFSU) FJ167661
C. fragilis Thailand JFK122 (SFSU) FJ167662
C. galeatus Thailand JFK67 (SFSU) FJ167663
C. liliputianus Puerto Rico MCA485 (BPI) AY916682
C. magnus Colombia DED4763(SFSU) FJ167666
C. columellifer Malaysia JFK72 (SFSU) FJ167665
C. sp. Ecuador MCA2538 (BPI) AY916686
Moniliophthora canescens Malaysia DED7518 (SFSU) FJ167668
M. perniciosa Ecuador MCA 2520 (BPI) AY916743
M. roreri var. gileri Ecuador DIS 331 AY230255
M. sp. North America MCA 2500 (BPI) AY916754
Marasmiellus subochraceus Malaysia DED7523 (SFSU) FJ167667
Marasmius crinisequi Malaysia TYS366 (KLU-M) FJ167669
* All available duplicates are deposited at SFSU.
multigene datasets (Aime and Phillips-Mora
2005) successfully delimited Crinipellis,
Moniliophthora and Chaetocalathus with high
statistical support.
Some interesting observations regarding
character evolution can be made. For example,
cheilocystidia shape may be a phylogenetically
informative character. The clade containing
C. trichialis, C. malesiana and the type of the
genus, C. scabella (= C. stipitaria), supported
with 52% MP-BS, 68% ML-BS and .89 PP
values all have simple cheilocystidia. The clade
consisting of C. actinophora, C. cupreostipes,
C. tabtim, C. dipterocarpi, C. dipterocarpi f.
cinnamomea and C. brunnescens, supported
with 68% ML-BS and 1.0 PP values all have
cheilocystidia with numerous knob-like apical
appendages. Pileipellis anatomy is also an
informative feature. Species with few pileus
hairs that arise from chains of short, broad,
pigment-incrusted cells and typically have no
KOH reaction, as represented in the phylogene-
tic tree by C. aff. iopus and Moniliophthora
species, formed a distinct clade in most
analyses, although with limited support.
This study did not attempt to test the
phylogenetic validity of Singer’s (1943, 1976)
infrageneric delimitation of Crinipellis. In
order to do that, one would need to include the
type species of each section in molecular
analyses and use a broad global sampling of
species. However, based on our ITS dataset
analyses of primarily Southeast Asian
Crinipellis, it appears that Singer’s infrageneric
classification is not supported. For example,
C. trichialis is placed in the section Grisentinae
based on the reaction of the pileus hairs turning
grey to green in KOH. Two new species,
C. malesiana and C. brunnescens would belong
in this section based on this KOH reaction
criterion; however, in both the MP and ML
analyses, these three species do not fall into the
same clade. In addition, C. scabella, the type of
section Crinipellis and characterized by a
negative KOH reaction, is nested in the clade
with C. trichialis and C. malesiana. Although
hyphal reactions in KOH are taxonomically
informative and aid in species delimitations,
they do not appear to be particularly useful in
determining phylogenetic relationships.
In summary, the molecular data resolved
12 species of Southeast Asian Crinipellis with
relatively high bootstrap support. Seven species
of Southeast Asian Crinipellis are not
represented in the molecular analyses: C. aff.
mirabilis, C. atrobrunnea, C. carecomoeis var.
litseae, C. cervino-alba, C. hepatica, C. pseu-
dostipitaria subsp. orientalis, and C. sepiaria.
Of the 19 species of Southeast Asian
Crinipellis recognized in this monograph, five
species and one form are new to science (32%).
This study also expanded the geographic
distribution of the following species: C.
actinophora, C. brunneipurpurea, C. diptero-

108
carpi, C. aff. iopus, C. aff. mirabilis, C. setipes
and C. trichialis, along with expanding the
known distribution of the genera Crinipellis
and Moniliophthora. Two species of Crinipellis
are formally transferred to the genus Moni-
liophthora, one new species of Moniliophthora
is described from Malaysia, and one species of
Crinipellis is formally transferred to the genus
Marasmiellus Murrill. In addition, 23 holotype
studies are presented. Future molecular
analyses using multiple genes and a global
sample set will help resolve the phylogenetic
relationships amongst the crinipelloid genera.
Synopsis of Southeast Asian Crinipellis
1. Crinipellis actinophora (Berk. & Broome)
Singer
2. Crinipellis atrobrunnea Pat.
3. Crinipellis brunneipurpurea Corner
4. Crinipellis brunnescens Kerekes,
Desjardin & Retn. sp. nov.
5. Crinipellis carecomoeis var. litseae Singer
6. Crinipellis cervino-alba Corner
7. Crinipellis cupreostipes Kerekes,
Desjardin & Lumyong sp. nov.
8. Crinipellis dipterocarpi Singer
9. Crinipellis dipterocarpi f. cinnamomea
Kerekes, Desjardin & Lumyong f. nov.
10. Crinipellis furcata Kerekes, Desjardin &
Lumyong sp. nov.
11. Crinipellis hepatica Corner
12. Crinipellis aff. iopus Singer
13. Crinipellis malesiana Kerekes, Desjardin
& Vikineswary sp. nov.
14. Crinipellis aff. mirabilis Singer
15. Crinipellis pseudostipitaria subsp.
orientalis Singer
16. Crinipellis sepiaria Pat. et Demange
17. Crinipellis setipes (Peck) Singer
18. Crinipellis tabtim Kerekes, Desjardin &
Lumyong sp. nov.
19. Crinipellis trichialis (Lév.) Pat. ex
Antonín et al.
20. Moniliophthora canescens (Har. Takah.)
Kerekes & Desjardin comb. nov.
21. Moniliophthora marginata Kerekes,
Desjardin & Vikineswary sp. nov.
22. Moniliophthora nigrilineata (Corner)
Kerekes & Desjardin comb. nov.
23. Marasmiellus subochraceus (Corner)
Kerekes & Desjardin comb. nov.
Excluded and Insufficiently Known Taxa,
Including Relevant Type Studies of
Extralimital Species.
24. Agaricus omotrichus Berk.
25. Collybia multicolor Petch
26. Crinipellis carecomoeis var. subelata
Singer
27. Crinipellis elata Pat.
28. Crinipellis filiformis G. Stev.
29. Crinipellis novae-zelandiae G. Stev.
30. Crinipellis patouillardii Singer
31. Crinipellis procera G. Stev.
32. Crinipellis readiae G. Stev.
33. Crinipellis roseola G. Stev.
34. Crinipellis substipitaria G. Stev.
35. Crinipellis velutipes G. Stev.
36. Crinipellis vinacea G. Stev.
Artificial Key to the Crinipellis and
Moniliophthora species from Southeast Asia
1. Pileus brightly pigmented with shades of purple,
violet brown, red to reddish-brown or ruby .......... 2
1.* Pileus not brightly pigmented, with shades of
brown, dull reddish brown, dark brown, black,
tawny or white....................................................... 9
2. Pileipellis hairs turning green, slate blue, grey,
olive-grey or purple in KOH................................. 3
2.* Pileipellis hairs not reacting in KOH .................... 5
3. Pileus ruby colored; pileipellis hairs becoming slate
blue in KOH....................................... 18. C. tabtim
3.* Pileus violet brown, dark purple or dark reddish
brown; pileipellis hairs turning grey, greyish green
or midnight blue in KOH ...................................... 4
4. Pileus violet brown to dark purple; basidiospores
10–14 × 3–5 μm; pileipellis and stipitipellis hairs
turning midnight blue in KOH................................
........................................... 3. C. brunneipurpurea
4.* Pileus dark reddish brown to violet brown;
basidiospores 8–12 × 3–5 μm; pileipellis hairs
initially purple in KOH, becoming grey to olive-
grey in time ............................ 14. C. aff. mirabilis
5. Pileus lilac .................................... 12. C. aff. iopus
5.* Pileus reddish brown............................................. 6
6. Lamellae edge reddish brown marginate; pileus up
to 22 mm diam.; pileipellis hairs subtended by
chains of broad cells ...............................................
.............................. 21. Moniliophthora marginata
6.* Lamellae edge not marginate; pileus less than 10
mm diam; pileipellis hairs subtended by long,
cylindrical tramal hyphae...................................... 7

Fungal Diversity
109
7. Basidiomes associated with dead twigs of Litsea;
basidiospores 12.8–16.5 × 4.5–5.8 μm; pileus
cinnamon buff to tawny olive..................................
................................. 5. C. carecomoeis var. litseae
7.* Basidiomes not associated with Litsea;
basidiospores in the range 6–11 × 3–5.5 µm; pileus
brown rufous to reddish brown or cinnamon ........8
8. Pileus brown rufous to liver-rufous; cheilocystidia
subventricose to subfusiform and without
appendages; stipe pallid to pinkish ochraceous or
bright rufous....................................11. C. hepatica
8. * Pileus reddish brown to cinnamon colored;
cheilocystidia clavate with 1–10 appendages; stipe
reddish brown.... 9. C. dipterocarpi f. cinnamomea
9. (1*) Stipe and abundant rhizomorphs both
brownish orange to copper coloured; stipe 40–550
mm in length ..............................7. C. cupreostipes
9.* Stipe (and rhizomorphs if present) dark brown to
black, never copper-coloured; stipe typically less
than 50 mm long (rarely up to 100 mm)..............10
10. Basidiomes associated with Poaceae................... 11
10.* Basidiomes not associated with Poaceae............. 14
11. Basidiomes associated with bamboo................... 12
11.* Basidiomes associated with other grasses...........13
12. Basidiospores 5.7–7.7 × 4.5–5.7 µm (xm = 6.6 × 5.2
µm); pileus bister to grey, usually azonate with
strigose, radial repent hairs; pileipellis hairs not
turning green in KOH...................... 16. C. sepiaria
12.* Basidiospores 9.6–11.5 × 6–7 µm (xm = 10.3 × 6.5
μm); pileus dark brown with pale brown margin
and distinct concentric zones with erect hairs;
pileipellis hairs turning green in KOH ....................
........................................................19. C. trichialis
13. (11*) Basidiospores 10–12.5 × 5.2–6 μm (xm =
11.1 × 5.6 µm); pileus dark brown to black ............
....................................................2. C. atrobrunnea
13.* Basidiospores 7.7–9.5 × 5–6 μm (xm = 8.3 × 5.5
µm); pileus tan to brown .........................................
................ 15. C. pseudostipitaria subsp. orientalis
14. (10*) Pileipellis hairs or stipitipellis tissues turning
green, olive green, or light brown to green in KOH
............................................................................. 15
14.* Pileipellis or stipe hairs not reacting in KOH...... 18
15. Lamellae staining dark brown to black when
bruised and with age...................4. C. brunnescens
15.* Lamellae not staining dark brown to black ......... 16
16. Rhizomorphs abundant, grey, greyish brown to
dark brown and distinctly coarse, very few black
and wiry rhizomorphs present.....1. C. actinophora
16.* Rhizomorphs absent or consistently thin and wiry,
black, never grey .................................................17
17. Basidiomes robust, with pileus 2–13 mm diam;
pleurocystidia present; pileus hairs and stipe tissue
initially yellowish green to green in KOH,
remaining so in time; rhizomorphs absent ..............
.....................................................13. C. malesiana
17.* Basidiomes not robust, with pileus 1–5 mm diam;
pleurocystidia absent; pileus hairs initially purple in
KOH, becoming grey to olive-greyish in time;
rhizomorphs present................ 14. C. aff. mirabilis
18. (14*) Basidiomes associated with dead twigs of
Litsea; basidiospores 12.8–16.5 × 4.5–5.8 μm (xm =
13.5 × 5.1 µm)......... 5. C. carecomoeis var. litseae
18.* Basidiomes not associated with Litsea;
basidiospores smaller, in the range 5–12.8 × 2–6
µm (xmr = 7.4–11.2× 3.0–4.7 µm)....................... 19
19. Cheilocystidia with more than 3 apical appendages
(if verrucose, see 22); rhizomorphs present, reddish
brown, dark brown or black ................................ 20
19.* Cheilocystidia absent or if present simple (lacking
apical appendages), rarely verrucose, forked, or
with 1–3 apical appendages; rhizomorphs absent or
if present black, thin and wiry............................. 22
20. Basidiomes associated with woody debris;
basidiospores xm = 8.2 × 4.5 µm, Qm = 1.9 .............
...........................................................17. C. setipes
20.* Basidiomes associated with decomposing leaves,
rarely twigs; basidiospores xmmr = 8.6–8.8 × 4.0 µm,
Qmm = 2.2 ............................................................21
21. Pileus light brown to dark brown ............................
....................................................8. C. dipterocarpi
21.* Pileus reddish brown to cinnamon coloured ...........
.......................... 9. C. dipterocarpi f. cinnamomea
22. (19*) Cheilocystidia absent................................. 23
22.* Cheilocystidia simple (lacking apical appendages),
forked or with 1–3 three apical appendages, rarely
verrucose............................................................. 24
23. Pileus white with brown, powdery-fibrous patches
or spots; stipe base pink ..........................................
............................ 23. Marasmiellus subochraceus
23.* Pileus-clay-color to brown; stipe base dark brown
............................ 22. Moniliophthora nigrilineata
24. Basidiomes dark brown to almost black overall;
stipe squamulose, (15–)30–40 mm long;
cheilocystidia typically apically forked...................
..........................................................10. C. furcata
24.* Basidiomes lighter in color; stipe fibrillose to
hispidulous, 4–10 mm long; cheilocystidia not
apically forked .................................................... 25
25. Incrustrations absent on the pileipellis cells; pileus
fawn-buff; species of swamp forests.......................
................................................... 6. C. cervino-alba
25.* Incrustations present on the pileipellis cells; pileus
brown rufous or greyish orange; species of mesic
forests.................................................................. 26
26. Pileus brown rufous to liver-rufous; stipe pallid to
pinkish ochraceous or bright rufous; cheilocystidia

110
without apical appendages ..............11. C. hepatica
26.* Pileus dull greyish orange; stipe brownish grey,
often arising from a thin white subiculum;
cheilocystidia with 0–3 apical appendages..............
................................20. Moniliophthora canescens
Taxonomy
1. Crinipellis actinophora (Berk. & Broome)
Singer, Sydowia 9: 397. 1955.
(Figs 2:1–6, 19)
≡ Marasmius actinophorus Berk. & Broome, J.
Linn. Soc., Bot. 14: 39. 1873.
= Marasmius coronatus Petch, Ann. Roy. Bot.
Gard., Peradeniya 6: 58. 1915.
Type: Sri Lanka, G.H.K. Thwaites 1164
[K #122107].
Analysis of the holotype specimen:
The holotype specimen consists of one
basidiome in poor condition, pressed flat and
glued to a card. As dried: Pileus 4 mm diam,
plano-convex with a circular depression (zone)
around a central papilla, radially shaggy, dark
brown. Lamellae not observable. Stipe 8 × 0.5
mm, base broken off, fibrillose, dark brown.
Few hymenial elements observable;
material revives poorly. Basidioles subfusoid.
No spores, cystidia or basidia observed.
Pileipellis of agglutinated, subparallel, thin- to
thick-walled hairs; hairs > 300 × 3.2–6(–10)
μm, cylindrical, ± equal for most their length,
obtuse, with scattered secondary septa; walls
0.5–2 μm thick, yellowish brown, dextrinoid,
non-incrusted; yellowish olive in KOH.
Description based on fresh material
Pileus (2–)4–10 mm diam., paraboloid
with incurved margins, subumbilicate with a
raised concentric ridge around disc and a
distinct papilla when young, becoming convex
to plano-convex, subumbonate with an acute
papilla and raised concentric zones around the
disc in age, dull, dry, with radially appressed-
fibrillose hairs forming tufts that hang below
margin edge; disc and papilla dark brown
(6F6–7), becoming brown (6E6) to greyish
brown (6D3) or cream at margin in age.
Context < 1 mm thick, white to cream,
unchanging. Lamellae narrowly adnexed to
free, subdistant to close with a few appearing
crowded, with 2–3 series of lamellulae, white.
Stipe 4–20(–35) × < 1 mm, central, terete,
slightly broader at base, insititious, dull, dry,
hirsute, ornamentation dense at base forming
upturned scales; dark brown (6F6) to brown
(6E6), becoming greyish brown (6D3) to cream
with age. Rhizomorphs present and abundant, 1
mm thick, glabrous to pubescent, grey, greyish
brown (6D3) to dark brown (7F5), tough, along
ground binding leaf litter and arising from a
subiculum on decomposing twigs and giving
rise to fruiting bodies. Odor and taste none.
Basidiospores 6–10 × 3–5 μm [xmr =
6.8–8.1 × 3.8–4.0 μm, xmm = 7.5 ± 0.5 × 3.9 ±
0.1 μm, Q = 1.4–2.7, Qmr = 1.7–2.1, Qmm = 1.9
± 0.1; n = 10–25 spores per 20 collections],
ellipsoid, smooth, hyaline, inamyloid, thin-
walled. Basidia 16–22 × 5–6 μm, clavate, 4-
spored. Basidioles clavate to cylindrical with
rounded to subacute apex. Pleurocystidia
absent. Cheilocystidia common, lamellar edge
sterile; main body 14–25 × 3–10 μm,
cylindrical to clavate or irregular, hyaline,
inamyloid, thick-walled; apical appendages (3–)
6 to numerous, 1–4(–15) × 0.5–3 μm, knobby
to finger-like, hyaline, inamyloid, thin walled;
a few collections with hardened crystal
exudates on longer projections. Pileipellis a
cutis, hyphae 24–40 × 4–15 μm, hyaline,
slightly dextrinoid, with dextrinoid spiral
incrustations, clamped, giving rise to terminal
hairs. Hairs 180–750 × 3–6 μm, hyaline to
light brown in water and KOH, dextrinoid,
thick-walled (up to 2 μm), apices bluntly
rounded or tapering, with secondary septations.
Pileus trama subregular; hyphae 3–8 μm diam.,
cylindrical, smooth, hyaline, inamyloid, thin-
walled. Lamellar trama regular; hyphae 3–8
μm diam., cylindrical, smooth, hyaline,
inamyloid, thin-walled. Stipe tissue monomitic,
parallel, tightly packed; stipitipellis composed
of repent cortical hyphae, terminal hairs and
caulocystidia; cortical hyphae 2–2.5 μm diam.,
cylindrical, slightly thick-walled, dextrinoid,
brown in water, becoming green in KOH;
medullary hyphae 1.5–3 μm diam., cylindrical,
slightly thick-walled, dextrinoid, hyaline to
light green in KOH; caulocystidia 50–138 × 10
μm, cylindrical, hyaline, slightly dextrinoid,
slightly thick-walled; terminal hairs 38–460 ×
2.5–8 μm, hyaline to tan in water and KOH,
dextrinoid, thick-walled, with secondary
septations. Clamp connections present.
Habitat: Primarily on decomposing twigs
and few found on decomposing dicot leaves.

Fungal Diversity
111
Known distribution: Malaysia, Thailand,
Sri Lanka, and Singapore.
Material examined: MALAYSIA. Kuala Lumpur,
Gombak Field Station, 12 January 2003, D. E. Desjardin
7519 (KLU-M, SFSU); Selangor, Hulu Langat, Sungai
Chongkak Forest Reserve, N03°12.705', E101°50.472',
elev. 188 m, 7 January 2005, Kerekes 78 (KLU-M,
SFSU); same location, 7 January 2005, Kerekes 79
(KLU-M, SFSU); same location, Kerekes 80 (KLU-M,
SFSU); Selangor, Selayang, Kanching Forest Reserve,
N03°17.954', E101°37.153', elev. ± 110 m, 8 Jan 2005,
Kerekes nos. 85, 86, 87, 88, 89, 90, 91, 92, 93 (KLU-M,
SFSU); same location, 9 Jan 2005, Kerekes nos. 95, 96,
98, 99, 100 (SFSU). SRI LANKA, G.H.K. Thwaites
1164 [Holotype of Marasmius actinophorus: K
#122107]; Hakgala, January 1914, coll. by T. Petch 3923,
on bark [Holotype of Marasmius coronatus: K #122106];
Hakgala, T. Petch 2227, April 1917 [K #122108];
Nuwara Eluya District, Hakgala Botanic Garden, 29
October 1974, D.N. Pegler 2131 [K #122111].
THAILAND, Nakorn Nayok Province, Khao Yai
National Park, Orchid Waterfall trail, 8 July 2005,
Kerekes 139 (CMU, SFSU).
Notes: The Malaysian and Thai
specimens match nicely with both Corner
(1996) and Pegler’s (1986) descriptions and
collections of C. actinophora. The holotype
was examined, however, many taxonomically
important features were not recovered. Petch
(1915) described Marasmius coronatus as a
new species but later recognized it to be the
same as Marasmius actinophorus. The coarse
grey rhizomorphs distinguish this species from
other Crinipellis, along with an obvious papilla
in the disc and cheilocystidia with numerous
short apical appendages. Another characteristic
is the reaction of stipe tissue turning green in
KOH, which was not previously noted. The
rhizomorphs in the collections varied from
being coarse, grey and pubescence to being
thinner, black and glabrous. It may be the case
that the black rhizomorphs are not associated
with C. actinophora, but rather are from a
species of Marasmius. No fruiting bodies were
observed arising from the thin black
rhizomorphs. Occasionally large inflated
terminal hairs were observd in the pileipellis as
seen in Fig. 2:6b. Vizzini et al. (2007) also
observed these types of cells in C.
pedemontana Vizzini, Antonín & Noordel., a
Fig. 2. Crinipellis actinophora (JFK 85). 1. Basidiomes (a. x2. – b. x1). – 2. Basidiospores (JFK 98). – 3. Basidioles. –
4. Cheilocystidia (JFK 78, 85, 98). –5. Caulocystidia. – 6a-b. Pileipellis hairs. – Bar: 2-4 = 10 µm; 5,6a = 25 µm; 6b =
20 µm.

112
species from Italy, and referred to them as
chlamydocytes. Crinipellis actinophora was
generally collected in lowland areas with
elevation ranging from 110–250 m, with the
exception of the single Thailand specimen
collected at 730 m.
2. Crinipellis atrobrunnea Pat., J. Bot.
(Morot) 5: 308. 1891. (Fig. 3: 1–2)
≡ Marasmius atrobrunneus (Pat.) Sacc., Syllog.
Fung. 11: 38. 1895.
Type: TONKIN (VIETNAM), Prov.
Hanoi, Ke So, 31 May 1890, coll. by Bon
#4362 (Holotype and Isotype: FH!).
Macromorphological description adapted from
the protologue and Singer (1943):
Pileus 1–10 mm diam., convex, with
sinuous to crenulate margin, dark brown to
black. Lamellae subfree to adnate, subdistant to
distant, thick, unequal, with at least 1 series of
lamellulae, anastomosing-venose, brown to
grey. Stipe 10–30 × 0.5–1 mm, equal or
subequal, roughly hairy-sulcate, dark brown to
black.
Micromorphological description based on
analysis of the holotype specimen
Basidiospores (9.3–)10–12.5 × 5.2–6 μm
[xm = 11.1 ± 0.1 × 5.6 ± 0.2 μm, Q = 1.8–2.3,
Qm = 2.0 ± 0.2, n = 11 spores per 1 collection],
elongate-ellipsoid to subfusoid, smooth,
hyaline, inamyloid, thick-walled. Basidia not
observed. Pleurocystidia absent. Cheilocystidia
not observed: no intact lamellar edges available
for examination. Pileipellis a cutis of inflated,
thin-walled, clamped, hyphae giving rise to
thick-walled terminal hairs. Hairs 48–160(–
200) × 3.5–5 μm, cylindrical, obtuse, hyaline to
pale yellowish brown, dextrinoid.
Habitat: On grasses.
Known Distribution: Vietnam.
Material examined: Type: TONKIN (VIETNAM),
Prov. Hanoi, Ke So, 31 May 1890, coll. by Bon #4362
(Holotype: FH).
Notes: Our type analysis matches that of
Yang (2000: 446). This represents a species of
Crinipellis, distinctive because of the relatively
long and narrow spores, short pileipellis hairs
and growth on grasses. Singer (1943) reports
the cheilocystidia to be 19–45 × 3.5–6 μm,
cylindrical to subfusoid, with a sinuose-wavy
outline and he did not observe branching.
Fig. 3. Crinipellis atrobrunnea Pat. (Holotype). – 1.
Basidiospores. – 2. Pileipellis hairs. – Bar:1-2 = 10 µm.
Unfortunately, this species was not included in
the molecular analysis because DNA extraction
from the type specimen was not permitted and
no fresh material was encountered.
3. Crinipellis brunneipurpurea Corner, Beih.
Nova Hedwigia 111: 118. 1996.
(Figs 4: 1–6, 19)
Type: Singapore, Gardens Jungle, Corner
s.n. 14 August 1940, herb. Corner (E).
Pileus 3–7 diam. × 2–7 mm tall,
hemispherical to convex with a small umbilicus
and central papilla when young, becoming
convex to broadly plano-convex, subumbilicate
with a central papilla, radially appressed-
fibrillose, with hairs hanging from margin edge,
with raised concentric ridges around disc on
some pilei; violet brown (10–11F7) to dark
ruby (12F4) or dark purple (14F4) with a
lighter colored disc. Context < 1 mm thick,
white to concolorous with pileus (12F4).
Lamellae adnexed to free, close to subdistant,
with 1–3 series of lamellulae, white, staining
purplish red to pinkish purple where bruised.
Stipe 6–33 × 1–2 mm, central, cylindrical, with
slightly twisted, fibrillose hairs forming tufts
near the base, with longitudinal intertwining
furrows above, tough; violet brown (10F7) to
dark ruby (12–14F4) overall. Rhizomorphs
absent but described as pale, thread-like
mycelium on the surface of twigs and leaves by
Corner (1996). Odor and taste none.
Basidiospores 10–14 × 3–5 μm [xmr =
11.6–12.3 × 4.1–4.3 μm, xmm = 12.0 ± 0.5 × 4.2
± 0.1 μm, Q = 2.2–4.7, Qmr = 2.7–3.1, Qmm =
2.9 ± 0.2; n = 25 spores per 2 collections],
subcylindrical to fusiform or narrowly

Fungal Diversity
113
Fig. 4. Crinipellis brunneipurpurea (JFK 107). – 1. Basidiomes (a. x2. – b. x1). – 2. Basidiospores. – 3. Basidia and
Basidioles. – 4. Cheilocystidia. – 5. Pileipellis hair. – 6. Caulocystidia. Bar: 2-4 = 10 µm; 5 = 20 µm; 6 = 25 µm.
amygdaliform with acute apex in side view,
smooth, hyaline, inamyloid, thin- to thick-
walled. Basidia 18–30 × 5–7 μm, clavate, 4-
spored. Basidioles clavate, few with subacute
apex. Pleurocystidia absent. Cheilocystidia
common; main body (12–)20–30(–35) μm,
clavate to irregularly clavate with occasional
small knobs or bifurcating apical appendages,
hyaline, inamyloid, thin- to thick-walled; apical
appendages 3–7 μm long, knobby, hyaline,
inamyloid, thin-walled; repent hyphae along
edge of gill 1–3 μm diam., hyaline, inamyloid,
thin-walled, septate, clamped, in some
instances giving rise to cheilocystidia.
Pileipellis a cutis, with inflated cells 30–40 ×
6–7 μm diam, hyaline, dextrinoid, giving rise
to terminal hairs with basal clamp connections.
Hairs 62–412 × 2.5–5 μm, cylindrical, majority
of apices rounded, secondary septa abundant,
base of hairs often wavy, violet in water,
becoming midnight blue in KOH, dextrinoid,
thick-walled. Pileus trama regular to
subregular; hyphae 3–7 μm diam., cylindrical,
hyaline, dextrinoid, thin-walled. Lamellar
trama regular to subregular; hyphae 3–10 μm
diam., cylindrical, hyaline, inamyloid, thin-
walled. Stipe tissue monomitic, parallel, tightly
packed; stipitipellis composed of repent
cortical hyphae and terminal hairs; cortical
hyphae 3–5 μm diam., cylindrical, slightly
thick-walled, dextrinoid, violet in water, bluish
grey in KOH; medullary hyphae 2–5 μm diam.,
cylindrical, thin-walled, inamyloid to slightly
dextrinoid, hyaline to slightly violet in water,
hyaline in KOH; terminal hairs 52–425 × 5–10
μm, similar to pileipellis hairs, cylindrical,
thick-walled, secondary septa abundant,
dextrinoid, violet in water, becoming midnight
blue in KOH. Clamp connections present.
Habitat: Scattered on decomposing
leaves and a few on twigs, in secondary forest
with Quercus and Dipterocarpus species.
Known Distribution: Indonesia and
Malaysia.
Material examined: INDONESIA, Java, West of
Bogor, Hutan Penelitian, Haurbentes S06°32.646′,
E106°26.211′, elev. 292 m, 14 January 2005, Kerekes
107 (BO, SFSU). MALAYSIA, Selangor, Selayang,
Kanching Forest Reserve N03°17.954′, E101°37.153′,
elev. 50–150 m, 8 June 2005, J.F Kerekes 83 (KLU-M,
SFSU); same location, 8 June 2005, Kerekes 84 (KLU-
M, SFSU).
Notes: Crinipellis brunneipurpurea was
first described from Singapore by Corner (1996)
and is characterized by a brownish purple
pileus with concolorus stipe, free gills that stain
purple with age and large spores (10–14 × 3–5
μm). The type was unavailable for examination,
but the Indonesian and Malaysian collections
match very well Corner’s protologue, except
for smaller basidiomes in the populations we
examined. Corner did not mention a midnight
blue reaction of the pileus and stipe hairs in

114
KOH, which easily distinguishes this species
from other purplish coloured species, such as C.
tabtim and C. iopus. Crinipellis corvina Har.
Takah. differs from C. brunneipurpurea by the
presence of whitish rhizomorphs, smaller
basidiospores (6.5–8 × 3.5–4.5), abruptly
adnexed gills, and growth on bark of Torreya
nucifera Siebold & Zucc. (Takahashi, 2000).
4. Crinipellis brunnescens Kerekes,
Desjardin & Retn. sp. nov.
(Figs 5: 1–6, 19)
MycoBank: 512271
Etymology: brunnescens (Latin) = turning brown;
referring to the brown staining and bruising of the
lamellae.
Pileus 7–15 mm latus, convexus vel late
convexus, atrobrunneus. Lamellae adnexae, albae,
contusae brunnescentes. Stipes 8–12 × 0.4–0.8 mm,
cylindricus, furfuraceus vel tomentosus, atrobrunneus.
Odor saporque nulli. Basidiosporae 6–10 × 4–5 μm,
ellipsoideae, hyalinae. Basidia 15–25 × 4–6 μm, clavata,
hyalinae, tetraspora. Pleurocystidia nulla. Cheilocystidia
12–25 × 4–8 μm, clavata, 2–8 apice surculis. Trama
lamellarum regularis. Caulocystidia 75–327 × 5– 6 μm,
cylindraceis. Holotypus: INDONESIA, Java, Mt.
Halimun National Park, loop trail from Cikiniki, elev. ca
1000 m, 14 January 1998, D.E. Desjardin 6791 (BO).
Pileus 7–15 mm diam., convex to
broadly convex, disc with or without a small,
flattened papilla, or a small central depression
surrounded by a raised ring, margin striate;
surface dull, dry, radially fibrillose; disc dark
brown (7–8F6–8), margin brown (7E7–8) with
darker striae. Context thin, white. Lamellae
horizontal, adnexed, subdistant with 2 series of
lamellulae, convex, broad, white, staining dark
brown to black where bruised and in age. Stipe
8–12 × 0.4–0.8 mm, central, terete, cylindrical,
± equal, curved, tough, pliant, furfuraceous to
tomentose, insititious; dark brown (7–8F6–8)
Fig. 5. Crinipellis brunnescens (DED 6791). – 1. Basidiomes (x1). – 2. Basidiospores. – 3. Basidia and basidioles. – 4.
Cheilocystidia. – 5. Caulocystidia. – 6. Pileipellis hairs. – Bar: 2-4, 5a, 6a = 10 µm; 5b, 6b = 25 µm.

Fungal Diversity
115
overall. Rhizomorphs absent. Odor and taste
none.
Basidiospores 6–10 × 4–5 μm [xm = 8.4
± 0.9 × 4.8 ± 0.4 μm, Q = 1.5–2.3, Qm = 1.7 ±
0.2, n = 25 spores per 1 collection], ellipsoid,
amygdaliform in side view, smooth, hyaline,
inamyloid, thin- to thick-walled. Basidia 15–25
× 4–6 μm, clavate, 4-spored. Basidioles clavate.
Pleurocystidia none. Lamellar edge sterile.
Cheilocystidia common; main body 12–25 × 4–
8 μm, clavate, hyaline, inamyloid, thin-walled;
apical appendages knob-like, 2–8 per cell, 1–6
μm long, hyaline, inamyloid, thin-walled.
Pileipellis a cutis, giving rise to terminal hairs
with basal clamp connections. Hairs 87–340 ×
4–5 μm, cylindrical, most with acute to blunt
apex, lacking obvious secondary septations,
thick-walled, inamyloid to slightly dextrinoid
with dextrinoid incrustations, light tan-brown
in water, becoming light brown to green in
KOH, in KOH incrustations not as obvious.
Pileus trama regular to subregular; hyphae 3–5
μm diam., hyaline, inamyloid, thin-walled.
Lamellar trama subregular to irregular; hyphae
1.5–4 μm diam., hyaline, inamyloid, thin-
walled. Stipe tissue monomitic, parallel, tightly
packed; stipitipellis composed of repent
cortical hyphae and terminal hairs; cortical
hyphae 3–5 μm diam., cylindrical, slightly
thick-walled, slightly dextrinoid, light tan-
brown in water, tan-brown to green in KOH;
medullary hyphae 2–5 μm diam., cylindrical,
parallel, tightly packed, hyaline, strongly
dextrinoid, thin-walled; terminal hairs 75–327
× 5– 6 μm, cylindrical, thick-walled, inamyloid
to slightly dextrinoid, with dextrinoid
incrustations, light tan-brown in water and
KOH. Clamp connections present.
Habitat: Solitary on undetermined
hardwood sticks under Castanopsis sp. in
montane primary forest.
Known distribution: Indonesia.
Material examined: INDONESIA, Java, Mt.
Halimun National Park, loop trail from Cikiniki, elev. ca
1000 m, 14 January 1998, D.E. Desjardin 6791
(Holotype: BO; Isotype: SFSU).
Notes: This species is characterized by
dark brown to black staining of the broad
lamellae when bruised and with age, a short
stipe, and dextrinoid incrustations on the pileus
hairs. Crinipellis brunnescens is somewhat
similar to C. herrerae Singer (1976) described
from Mexico, but the latter differs in forming a
deep rusty brown pileus, lamellae that do not
stain brown, a pallid stipe, and pileus hairs that
lack incrustations and do not turn green in
KOH. The numerous apical appendages on the
cheilocystidia of C. brunnescens differentiate
this species from C. mirabilis (no. 14 below).
Microscopically, C. stupparia (Berk. & M.A.
Curtis) Pat., from the New World tropics
resembles C. brunnescens, however C.
stupparia forms a light brown pileus under 5
mm diam., a much longer stipe (up to 25 mm),
and the lamellae do not stain brown (Singer,
1943).
5. Crinipellis carecomoeis var. litseae
Singer, Lilloa 8: 489. 1943. (Fig. 6: 1–2)
Type: Philippines, Laguna, Mt. Makiling,
5 Sept. 1917, coll. by N. Catalan, det. by
Patouillard as Crinipellis stipitaria, on dead
twigs of Litsea perrottetii (Holotype: FH!).
Macromorphological description adapted
from protologue
Pileus 3–4 mm diam. in dried condition,
mostly campanulate with a more or less
prominent papilla, becoming convex to plano-
convex in age with either a flat disk, an
umbilicus or more frequently with a small
papilla, hairy all over; cinnamon buff to tawny
olive in dried condition. Lamellae narrowly
adnexed or free, close, rather broad, probably
white when fresh. Stipe 7–21 × 0.2–0.5 mm
when dried, equal, more or less hirsute and
somewhat grooved below, concolorous with
the pileus. No mention of rhizomorphs, odor or
taste.
Micromorphological description based on
analysis of the holotype specimen
Basidiospores 12.8–16.5 × 4.5–5.8 μm
[xm = 13.5 ± 1.5 × 5.1 ± 0.5 μm, Q = 2.4–2.9,
Qm = 2.7 ± 0.5; n = 11 spores per 1 collection],
subfusoid, smooth, hyaline, inamyloid, thin-
walled. Basidia not observed. Basidioles
subfusoid. Pleurocystidia absent. Lamellar
edge badly colonised with a deuteromycete;
fertile; no cheilocystidia observed. The cystidia
described by Singer (1943) as “most commonly
with conidia-like appendices which are
confined to the very top or the upper half of the
cheilocystidium and are subglobose with

116
Fig. 6. Crinipellis carecomoeis var. litseae (Holotype). –
1. Basidiospores. – 2. Pileipellis hairs. – Bar: 1-2 = 10
µm.
attenuate connection with the main body” are
undoubtedly chains of conidia from the
deuteromycete colonisation. Pileipellis a cutis
giving rise to thick-walled hairs. Hairs
immeasurable in length (> 200 μm) × 3.5–6.5
μm, evenly cylindrical, not appreciably
attenuated towards the apex, walls smooth,
yellowish to yellowish brown, inamyloid to
dextrinoid; thick-walled 1.5–2.5 μm, apex of
hairs with or without ladder-like secondary
septations; base of hairs sometimes with dark
brown plaque-like incrustations.
Habitat: On dead twigs of Litsea
perrottetii.
Known distribution: Philippine Islands.
Material examined: PHILIPPINES, Laguna, Mt.
Makiling, 5 September 1917, coll. by N. Catalan, det. by
Patouillard as Crinipellis stipitaria, on dead twigs of
Litsea perrottetii (Holotype: FH).
Notes: This taxon is apparently known
only from the holotype specimen. It is
tentatively accepted here as representing a
distinct species of Crinipellis and we have no
comment on whether it represents a variety of
C. carecomoeis (described originally from
Cuba) or a synonym of another taxon.
Crinipellis carecomoeis var. subelata Singer
(1943), described from material collected in
New Caledonia, is accepted as an insufficiently
known taxon in this monograph (see no. 22
below).
6. Crinipellis cervino-alba Corner, Beih.
Nova Hedwigia 111: 119. 1996.
Type: Malaysia, Johore, Mawai, Corner
s.n. 28 September 1934 (herb. Corner, E).
Description adapted from the protologue:
Pileus 3–6.5 mm diam., convex to nearly
plane, with a minute acute umbo, appressed-
fibrillose to felted in radial ridges, fawn buff
with a fuscous disc. Context thin, pale brown.
Lamellae free, subdistant (12–14) with 2–3
series of lamellulae, narrow, white. Stipe 4–7 ×
0.3–0.5 mm, slightly thickened at base,
appressed fibrillose-felted, pale fuscous fawn.
Rhizomorphs absent. Odor and taste none.
Basidiospores 9–11 × 4.5–5.5 μm,
fusiform, smooth, hyaline. Basidioles sub-
acerose. Pleurocystidia absent. Lamellar edge
sterile. Cheilocystidia with main body –20 × 5–
7 μm, irregularly clavate, verrucose with
obtuse warts 1–3 × 1.0 μm. Pileipellis hairs 3–
6 μm diam., cylindrical and tapering to a
subacute apex 2 µm diam., smooth (not
incrusted), thick-walled, brown.
Habitat: On dead wood in swamp forest.
Known distribution: Malaysia.
Material examined: Type was not available for
study. Notes: Corner noted how this species is
similar to C. herrerae Singer, described from
Mexico (Singer, 1976), although C. cervino-
alba is lighter in color, has a shorter stipe and
has cheilocystidia with shorter apical
appendages. This species has an unusual
habitat in swamp forests and was not
recollected during our field studies.
7. Crinipellis cupreostipes Kerekes,
Desjardin & Lumyong sp. nov.
(Figs 7: 1–6, 19)
MycoBank: 512272
Etymology: cupreus (Latin) – copper-colored;
stipes (Latin) -stipe; referring to the copper colored stipe.
Pileus 1–9 mm latus, hemisphaericus dein
convexus, brunneus. Lamellae adnexae. Stipes 40–550 ×
0.5–1 mm, cylindricus, pubescentes, cupreus. Odor
saporque nulli. Basidiosporae 9–11 × 4–4.5 μm,
fusiformis, hyalinae. Basidia 19–20 x 4–5 μm, clavata,
hyalinae, tetraspora. Pleurocystidia nulla. Cheilocystidia
14–17 x 4–6 μm, clavata, 1–7 apice surculis. Trama
lamellarum regularis. Caulocystidia 72–160 x 3–6 μm,

Fungal Diversity
117
Fig. 7. Crinipellis cupreostipes (JFK 31). – 1. Basidiomes (JFK 31, 55, 131) (a. 0.5x. b. 0.5x. c. 2x. d. 0.5x). – 2.
Basidiospores. – 3. Basidioles. – 4. Cheilocystidia. – 5. Caulocystidia. – 6. Pileipellis hairs. – Bar: 2-4 = 10 µm; 5a, 6a
= 10 µm; 5b, 6b = 25 µm.
cylindraceis. Holotypus: THAILAND, Chiang Mai
Province, Doi Inthanon Naitonal Park, jct of Highway
1009 and road to Mae Chem, N18 31.58', E098 29.64',
1703 m, 27 June 2005, Kerekes 131 (CMU).
Pileus 1–9 mm diam., truncately
hemispherical to convex, becoming plano-
convex with a circular depression surrounding
a central papilla, a few with a raised disc and
concentric raised zones around papilla at
maturity; many stipes topped with tiny,
unopened primoridal pilei, these convex to
conical with inrolled margins and a distinct
papilla; radially appressed-fibrillose; papilla
black to dark brown; disc dark brown (6F7)
becoming brown (6D7) to cream at margin
with brown (6E7) fibrils, becoming paler
overall with age. Context < 1 mm thick, cream
to light brown (6D6), unchanging. Lamellae
adnexed to free, subdistant to close with 2–3
series of lamellulae, white to cream. Stipe 40–
550 x 0.5–1 mm, central, cylindrical, wiry,
rhizomorphic stipes often terminating in
primordial caps, few branching; with white to
cream cobweb-like pubescence along stipe;
white at extreme apex becoming brownish
orange (6C7) to brown (6D7) and resembling
copper wire. Rhizomorphs abundant and
similar to stipes, resembling copper wire, most
arise from decomposing leaf litter and stand
erect, some terminating in primordial pilei.
Odor and taste none.
Basidiospores 9–11 × 4–4.5 μm [xm = 9.6
± 0.6 × 4.0 ± 0.1 μm, Q = 2.2–2.8, Qm = 2.4 ±
0.2, n = 25 spores per 1 collection], fusiform,
smooth, hyaline, inamyloid, thin- to thick-
walled. Basidia 19–20 × 4–5 μm, clavate, 4-
spored. Basidioles clavate to cylindrical with
rounded to subacute apex. Pleurocystidia
absent. Cheilocystidia common; main body 14–
17 × 4–6 μm, clavate, hyaline, inamyloid, thin-
walled; apical appendages 1–7 μm long,

118
cylindrical or finger-like, hyaline, inamyloid,
thin walled. Pileipellis a cutis of hyaline,
slightly dextrinoid hyphae giving rise to
terminal hairs with basal clamp connections.
Hairs 260–520 × 4–5 μm diam., cylindrical,
apices bluntly rounded or tapering, with
secondary septations, thick-walled, slightly
dextrinoid, tan to hyaline in water, light brown
to tan in KOH. Lamellar and pileus trama
regular; hyphae 2–7 μm diam., hyaline,
inamyloid. Stipe tissue monomitic, parallel,
tightly packed; stipitipellis composed of repent
cortical hyphae and terminal hairs; cortical
hyphae 4–6 μm diam., cylindrical, dextrinoid,
copper yellow in water, yellowish tan in KOH;
medullary hyphae 4–7 μm diam., cylindrical,
inamyloid, hyaline in water, hyaline in KOH;
terminal hairs 72–160 × 3–6 μm, cylindrical,
thick-walled (1–2 μm), dextrinoid, hyaline to
brownish yellow in water and KOH, similar to
hairs in pileipellis but more with an acute and
tapering apex and less secondary septa. Clamp
connections present.
Habitat: Gregarious on decomposing
dicot leaves and small twigs, in montane
primary cloud forest containing Quercus,
Magnolia, and Ficus species.
Known distribution: Thailand.
Material examined: THAILAND. Chiang Mai
Province, Doi Inthanon National Park, jct of Highway
1009 and road to Mae Chem, N18 31.58', E098 29.64',
1703 m, 25 June 2004, Kerekes 31 (CMU, SFSU); same
location, 3 July 2004, Kerekes 55 (CMU, SFSU); same
location, 27 June 2005, Kerekes 131 (Holotype: CMU;
Isotype: SFSU).
Notes: The most distinctive and
diagnostic feature of this new species is the
very long copper coloured stipes (up to 550
mm), and abundant copper coloured
rhizomorphs, many of which terminate in
primordial pilei. These are unique characters
within the genus Crinipellis. Crinipellis elata
Pat., accepted here as an insufficiently known
taxon, was described as having a long stipe (up
to 170 mm), but the stipe is greyish brown not
copper colored, and unfortunately the holotype
specimen is represented by immature, sterile
basidiomes (see no. 27 below).
8. Crinipellis dipterocarpi Singer, Lilloa 8:
496. 1943. (Fig. 8: 1–5, 19)
Type: VIETMAN (TONKIN), Eberhardt
LBA #6 det. by Patouillard as Crinipellis
stipitarius, on leaves and fruits of Diptero-
carpus costatus (Holotype: F!).
Analysis of the holotype specimen
The holotype specimen consists of five
pilei and eight stipes (two as intact basidiomes)
loose in the packet or with stipe attached to
dipterocarp leaf fragments. As dried: Pileus 3–
4 mm diam., convex-umbilicate, hirsute,
evenly brown around a darker umbilicus.
Lamellae close, broad, brown with white
crystalline edges. Stipe up to 15 × 0.5 mm, pale
greyish brown, pubescent, arising from the
substrate, lacking obvious rhizomorphs.
Basidiospores (7–)7.5–10 × 3.2–3.8 μm
[xm = 8.3 ± 1.0 × 3.5 ± 0.2 μm, Q = 2.1–2.6,
Qm = 2.4 ± 0.2, n = 10 spores per 1 collection],
elongate-ellipsoid to subfusoid, smooth,
hyaline, inamyloid, thin-walled or a few thick-
walled. Basidia clavate, 4-spored, clamped.
Basidioles subfusoid to fusoid. Pleurocystidia
absent. Cheilocystidia common, lamellar edge
sterile; main body 8–16 × 3.2–8 μm,
subcylindrical to clavate, with 4–8 apical
appendages; apical appendages 1.5–6.5 × 1–1.8
μm finger-like, thick-walled, hyaline,
inamyloid. Pileipellis of thick-walled hairs.
Hairs immeasurable in length, > 300 × 2–3.2
μm, cylindrical and often sinuous below but
gradually tapering to an acute apex, hyaline to
yellow or ochraceous, inamyloid to dextrinoid
(with dextrinoid contents), with scattered
secondary septation; walls up to 1 μm thick,
smooth, non-incrusted; subtending hyphae
non–incrusted, thinner-walled, inamyloid.
Description based on fresh material
Pileus 1–10 mm diam., hemispherical to
convex with inrolled margins and no obvious
papilla, becoming convex to plano-convex in
age with a shallow umbilicus, no ridges or
furrows of concentric circles around disc,
hirsute to radially appressed-fibrillose with
hairs often hanging over the margin; disc and
hairs light brown (6D6) to dark brown (6–7F7),
with age margin becoming pale greyish orange
(5B3–4) to cream. Context < 1 mm thick, white.
Lamellae adnate to shallowly adnexed,
subdistant to close with 1–3 series of lamellu-
lae, white. Stipe 11–50 × <1 mm, central, terete,
cylindrical, equal, tough, longitudinally striate,
twisted, insititious, hirsute overall,

Fungal Diversity
119
Fig. 8. Crinipellis dipterocarpi (JFK 130). – 1. Basidiomes (JFK 136) (x2). – 2. Basidiospores. – 3. Basidia and
basidioles. – 4. Cheilocystidia. – 5a-b. Pileipellis hair. – Bar: 2-4, 5b = 10 µm; 5a = 50 µm.
ornamentation dense at base; extreme apex
white to cream, brown (6E7) to dark brown
(6F8) below. Rhizomorphs very thin,
uncommon, arising from decomposing leaf
litter, reddish brown to black, not present in all
collections. Odor and taste none.
Basidiospores 5–10 × 3–5.5 μm [xmr =
8.0–8.8 × 3.8–4.1 μm, xmm = 8.6 ± 0.7 × 4.0 ±
0.1 μm, Q = 1.3–3.0, Qmr = 2.0–2.3, Qmm = 2.2
± 0.1; n = 10–25 spores per 9 collections],
elongate-ellipsoid to subfusoid, smooth,
hyaline, inamyloid, thin-walled or a few thick-
walled. Basidia 18–22 × 5–6 μm, clavate, 4-
spored. Basidioles clavate to subfusoid or
fusoid. Pleurocystidia absent. Cheilocystidia
common, lamellar edge sterile; main body 8–
19 × 3.2–10 μm, subcylindrical to broadly
clavate, hyaline, inamyloid, thin-walled, with
4–9 apical appendages; apical appendages 1.5–
6.5 × 1–1.8 μm, finger-like, hyaline, inamyloid,
thick-walled. Pileipellis a cutis with large
inflated cells, some with reddish brown and
dextrinoid incrustations, giving rise to terminal
hairs with basal clamp connections. Hairs 400–
537 × 2–6 μm, cylindrical and often sinuous
below but gradually tapering to an acute or
rounded apex, secondary septations scattered to
abundant, thick-walled (up to 2 μm), inamyloid
to dextrinoid, light brown to tan with some
incrustations in water, hyaline to yellow or
ochraceous in KOH. Lamellar and pileus
trama regular; hyphae 3–5 μm diam.,
cylindrical, smooth, hyaline, inamyloid, thin-
walled. Stipe tissue monomitic, parallel, tightly
packed; stipitipellis composed of repent
cortical hyphae and terminal hairs and
caulocystidia; cortical hyphae 3–5 μm diam.,
cylindrical, greenish brown in KOH; medullary
hyphae up to 5 μm diam., cylindrical, hyaline
in KOH; caulocystidia 25–40 × 5–6 μm,
cylindrical; terminal hairs 85–300 × 6–8 μm,
thick-walled (2–3 μm), inamyloid to dextrinoid,
hyaline to brown in KOH, similar to hairs in
pileipellis but more with an acute apex and less
secondary septa. Clamp connections present.
Habitat: Scattered to gregarious on
leaves and small sticks of undetermined dicot
trees in montane primary forests with Pinus,
Castanopsis and Dipterocarpaceae species, and
in secondary forests with Dipterocarpus
obtusifolius Teijsm. ex Miq. and Pinus kesiya
Royle ex Gordon.

120
Known Distribution: Indonesia, Malaysia,
Thailand, and Vietnam.
Material examined: INDONESIA. Borneo, East
Kalimantan, Kayan Mentarang National Park, 5 April
2003, A. Retnowati 410 (same as ZT 12031) (SFSU).
MALAYSIA. Pahang, Fraser’s Hill, Bishop trail, 26
September 2003, TYS 111 (KLU-M, SFSU).
THAILAND. Chiang Mai Province, Huai Nam Dang
National Park, Hwy 1095, 24 June 2003, D. E. Desjardin
7570 (SFSU); Chiang Mai Province, 22 km marker on
Hwy 1095 on road to Mae Hong Son, elev. ca 725 m, 2
July 2003, D. E. Desjardin 7602 (SFSU); Chiang Mai
Province, Doi Inthanon National Park, Hwy 1009 at 25
km marker, N18˚32.56', E98˚33.51', elev. 1075 m, 6
June 2005, Kerekes 130 (CMU, SFSU); Chiang Mai
Province, Doi Inthanon National Park, jct of Highway
1009 and road to Mae Chem, N18˚31.58', E98˚29.64',
elev 1700 m, 25 June 2004, Kerekes 33 (CMU, SFSU);
same location, 3 July 2004, Kerekes nos. 51, 52, 53, 54
(CMU, SFSU); same location, 27 June 2005, Kerekes
nos. 130, 132 (CMU, SFSU); Chiang Mai Province, Mae
Sae Village near 50 km marker on Hwy 1095,
N19˚14.599', E98˚38.456', elev. 962 m, 26 June 2005,
Kerekes 126 (CMU, SFSU); same location, 26 June
2005, Kerekes 127 (CMU, SFSU); same location, 3 July
2005, Kerekes 136 (SFSU). VIETNAM (TONKIN).
Eberhardt LBA #6, det. by Patouillard as Crinipellis
stipitarius, on leaves and fruits of Dipterocarpus
costatus (Holotype: FH).
Notes: Our collections of C. dipterocarpi
generally have longer stipes than reported in
the protologue by Singer, and the
cheilocystidia measured in our type study and
from recently collected material were shorter
than those reported in the protologue, 8–19 µm
versus 15–27 μm (respectively). There was
also no report in the protologue of incrustations
on the pileipellis cells or terminal hairs. The
incrustations are apparent in the recent
collections, especially when mounted in water
or Melzer's reagent (where they are dextrinoid).
Our recent collections were collected at high
elevations in forests with Dipterocarpaceae;
unfortunately, the leaves and substrate were not
always identifiable. Singer (1976) reported C.
dipterocarpi to be close to C. foliicola Singer
(from Florida and Venezuela), C. phyllophila
Singer (from Mexico), and C. piceae Singer
(from temperate Asia and North America). In
addition to disparate geographic location,
habitat and substrate, C. dipterocarpi can be
distinguished from all of these species in
cheilocystidia and pileus hair morphologies.
For a detailed comparison, refer to Singer
(1976: 31).
9. Crinipellis dipterocarpi f. cinnamomea
Kerekes, Desjardin & Lumyong, forma nov.
(Figs 9: 1–6, 19)
MycoBank: 512273
Etymology: cinnamomeus (Latin) – referring to
the cinnamon brown coloured pileus and stipe.
Pileus 2–7 mm latus, hemisphaericus,
cinnamomeus. Lamellae liber ad adnexae. Stipes 15–45
× < 1 mm, cylindricus, fibrillosus, rubrobrunneus. Odor
saporque nulli. Basidiosporae 6–11 × 3–5 μm,
ellipsoideae, hyalinae. Basidia 15–21 × 4–5 μm, clavata,
hyalinae, tetraspora. Pleurocystidia nulla. Cheilocystidia
9–20 × 5–7 μm, clavata, 1–10 apice surculis.
Caulocystidia 67–150 × 5–7.5 μm, cylindraceis.
Holotypus: THAILAND. Chiang Mai Province, Doi
Inthanon National Park, jct of Highway 1009 and road to
Mae Chem, N18˚31.58', E98˚29.64', elev. 1703 m, 3 July
2004, Kerekes 50 (CMU).
Pileus 2–7 mm diam., when young
truncately hemispherical and papillate to
hemispherical-subumbilicate with a central
papilla, margin incurved, in age becoming
convex, subumbilicate with a less obvious
papilla, radially appressed-fibrillose with hairs
hanging over edge of margin; reddish brown
(8E–F7) to cinnamon-coloured gradually
becoming light brown (6D6) near the margin,
in age margin becoming cream beneath the
hairs. Context < 1 mm thick, white, unchanging.
Lamellae free to adnate, subdistant to close
with 2 series of lamellulae, white. Stipe 15–45
× < 1 mm, central, cylindrical, ± equal, tough,
insititious, fibrillose to squamulose overall,
ornamentation denser when young and near the
base, with longitudinal furrows; reddish brown
(8E7), becoming cream with reddish brown
hairs but often appearing light brown (6D6–7)
overall. Rhizomorphs not present in all
collections, if present, hair-like, short (5–30
mm), reddish brown (8F7) to brown (7E6), on
decomposing debris; no fruiting bodies arising
from the rhizomorhps. Odor and taste none.
Basidiospores 6–11 × 3–5 μm [xmr =
8.5–9.3 × 3.8–4.0 μm, xmm = 8.8 ± 0.3 × 4.0 ±
0.1 μm, Q = 1.4–3.0, Qmr = 2.2–2.3, Qmm = 2.2
± 0.1; n = 20–25 spores per 6 collections],
ellipsoid to oblong or subcylindrical, smooth,
hyaline, inamyloid, thin-walled. Basidia 15–21
× 4–5 μm, clavate, 4-spored. Basidioles clavate
to cylindrical with rounded to subacute apex.
Pleurocystidia absent. Cheilocystidia crowded
along gill edge; main body 9–20 × 5–7 μm,

Fungal Diversity
121
Fig. 9. Crinipellis dipterocarpi f. cinnamomea (JFK 32). – 1. Basidiomes (JFK 32, 104) (x2). – 2. Basidiospores. – 3.
Basidia and basidioles. – 4. Cheilocystidia. –5a. Pileipellis cell. 5b. Pileipellis hair. 6. Caulocystidia. – Bar: 2-5a = 10
µm; 5b, 6 = 25 µm.
clavate to broadly clavate, hyaline, inamyloid,
thin-walled to slightly thick-walled, with 1–10
apical appendages; apical appendages 2–6 μm
long, finger-like, hyaline, inamyloid, thin
walled. Pileipellis a cutis of inflated cells, 6–11
μm diam., hyaline, with peg-shaped to irregular,
reddish brown incrustations, dextrinoid,
hyaline to tan in KOH, giving rise to terminal
hairs with basal clamp connections. Hairs 62–
475 × 4–5 μm, cylindrical, apices rounded to
acute, few with blunt apex, with widespread
secondary septations, thick-walled (1.5–3.5
μm), dextrinoid, golden brown to yellowish
brown in KOH, with reddish brown
incrustations in water and Melzer’s reagent
(few). Pileus trama regular; hyphae 3–7 μm
diam., hyaline, inamyloid. Lamellar trama
regular; hyphae 3–5 μm diam., hyaline,
inamyloid. Stipe tissue monomitic, parallel,
tightly packed; stipitipellis composed of repent
cortical hyphae and terminal hairs; cortical
hyphae 2–8 μm diam., cylindrical, thick-walled,
dextrinoid, light golden brown; medullary
hyphae 2–6 μm diam., cylindrical, hyaline,
thin-walled, strongly dextrinoid; terminal hairs
67–150 × 5–7.5 μm, cylindrical, thick-walled,
with distant or absent secondary septations,
dextrinoid, light brown to yellow brown in
water and KOH. Clamp connections present.
Habitat: Scattered on decaying dicot
leaves and twigs in primary montane cloud
forest and in mixed forests with Magnoliaceae,
Castanopsis, Quercus, and Ficus species.
Known Distribution: Indonesia, Malaysia,
and Thailand.
Material examined: INDONESIA. Java, Cibodas
Botanical Garden, trail to Mt. Gede, S06˚44.675',
E107˚00.423', elev. 1406 m, 22 January 1999, A.
Retnowati 151 (SFSU); same location, 13 January 2005,
Kerekes 104 (BO, SFSU). MALAYSIA. Pahang
Province, Fraser’s Hill, Kindersley Trail, 9 September
2003, T. Shin 193, (SFSU); Pahang Province, Fraser’s
Hill, Bishop’s trail, N03˚42.949', E101˚44.430', elev.
1212 m, 15 January 2004, Kerekes 17 (KLU-M, SFSU);
Pahang Province, Fraser’s Hill, Pine Tree Trail,
N03˚42.700', E101˚43.650', elev. 1320–1140 m, 16
January 2004, Kerekes 20 (KLU-M, SFSU).
THAILAND, Chiang Mai Province, Doi Inthanon
National Park, jct of Highway 1009 and road to Mae
Chem, N18˚31.58', E98˚29.64', elev. 1703 m, 25 June
2004, Kerekes 32 (CMU, SFSU); same location, 3 July

122
2004, Kerekes 50 (Holotype: CMU; Isotype: SFSU);
same location, 27 June 2005, Kerekes 133 (CMU,
SFSU).
Notes: Crinipellis dipterocarpi f.
cinnamomea has a characteristic reddish brown
to cinnamon coloured pileus and stipe. In
addition to this distinct colouring, the
incrustations on the pileipellis cells subtending
the terminal hairs have a more distinct reddish
brown colour and the spores are slightly longer
than C. dipterocarpi f. dipterocarpi. Both
forms were often collected in the same location.
Crinipellis dipterocarpi f. cinnamomea is
phenetically similar to C. tabtim, but the latter
species has pileus hairs that turn grey to slate
blue in KOH and have distinct burgundy
coloured incrustations in water. Crinipellis
hepatica also has a reddish brown (brown
rufous) pileus and stipe, however, the
cheilocystidia of this species lack apical
appendages (Corner, 1996). There is 92% MP-
BS, 82% ML-BS and 1.0 PP support for
inclusion of C. dipterocarpi f. cinnamomea
within the C. dipterocarpi f. dipterocarpi clade,
and all four included f. cinnamomea sequences
form an internal clade with 88% BS and 1.0 PP
support (Fig. 1). Interestingly, DED 7570 a
collection with purple pileus, is also included
in the C. dipterocarpi clade suggesting that
there may be additional color forms within the
species.
10. Crinipellis furcata Kerekes, Desjardin &
Lumyong sp. nov. (Figs 10: 1–6, 19)
MycoBank: 512274
Etymology: furcata (Latin) – forked; referring to
the forked cheilocystidia.
Pileus 6–11 mm latus, convexus vel late
convexus, atrobrunneus. Lamellae liber ad adnexae.
Stipes (15–) 30–40 × 1 mm, cylindricus, squamulosus,
atrobrunneus. Odor saporque nulli. Basidiosporae 6.5–9
× 4–5 μm, ellipsoideae, hyalinae. Basidia 23–30 × 6–8
μm, clavata, hyalinae, tetraspora. Pleurocystidia nulla.
Cheilocystidia (15–) 22–42 × 4–15 μm, clavata,
bifurcates, 0–2 apice surculis. Trama lamellarum
regularis. Caulocystidia -75 × 6–7 μm, cylindraceis.
Holotypus: INDONESIA. Bali, south ridge of Mt. Catur,
east side of Lake Bratan, elev. 1200–1800 m, 17 January
1999, D. E. Desjardin 6951 (BO).
Pileus 6–11 mm diam., convex to plano-
convex, umbilicate, papillate, disc scaly,
margin radially-fibrillose, extreme margin with
long hairs that form a complete fibrillose
partial veil and remain as an appendiculate
margin on expanded pilei; surface dull, dry;
dark brown (7F6–8) to nearly black overall.
Context < 1 mm thick, white to cream.
Lamellae horizontal, free to adnexed, close to
crowded with 2–3 series of lamellulae, narrow
(up to 1.5 mm), white to cream, non-marginate.
Fig. 10. Crinipellis furcata (DED 6951). – 1. Basidiomes (x1). – 2. Basidiospores. – 3. Basidia and basidioles. – 4.
Cheilocystidia. – 5. Pileipellis hair. – 6. Caulocystidia. – Bar: 2-6 = 10 µm.

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123
Stipe (15–) 30–40 × 1 mm, central, cylindrical,
terete, equal, tough, solid, insititious, dull, dry,
squamulose and dark brown (7F6–8) overall.
Rhizomorphs absent. Odor and taste none.
Basidiospores 6.5–9 × 4–5 μm [xmr =
7.2–7.6 × 4.3–5.0 μm, xmm = 7.4 ± 0.3 × 4.7 ±
0.4 μm, Q = 1.3–2.3, Qmr = 1.4–1.8, Qmm = 1.6
± 0.2; n = 10–25 spores per 3 collections],
ellipsoid, smooth, hyaline, inamyloid, thin- to
thick-walled. Basidia 23–30 × 6–8 μm, clavate,
4-spored, sterigmata 5–6 μm long. Basidioles
clavate to cylindrical with rounded to subacute
apex. Pleurocystidia absent. Cheilocystidia
crowded along gill edge; main body (15–) 22–
42 × 4–15 μm, clavate to lageniform or
irregular, majority bifurcated, hyaline, thin-
walled, inamyloid; apical appendages (2–)15–
28 × 2–4 μm, 0–2 per cell (rarely more than
two), hyaline, inamyloid, thin-walled.
Pileipellis a cutis of inflated cells 5–8 μm
diam., hyaline to light brown, slightly
dextrinoid, with granular or spiral incrustations,
giving rise to terminal hairs with basal clamp
connections. Hairs > 70 × 4–6 μm, cylindrical,
apices rounded, with abundant, widely spaced
secondary septations, dextrinoid, brown in
water and KOH. Lamellar and pileus trama
subregular; hyphae 3–9 μm diam., hyaline,
inamyloid. Stipe tissue monomitic, parallel,
tightly packed; stipitipellis composed of repent
cortical hyphae and terminal hairs; cortical
hyphae 3–5 μm diam., cylindrical, slightly
thick-walled, dextrinoid, hyaline to yellowish
brown in KOH; medullary hyphae 1–4 μm
diam., cylindrical, strongly dextrinoid; terminal
hairs < 75 × 6–7 μm, cylindrical, similar to
pileipellis hairs, thick-walled, weakly
dextrinoid, hyaline to yellow or brown in KOH.
Clamp connections present.
Habitat: Solitary on woody debris in
montane primary and secondary forests and
botanical garden.
Known Distribution: Indonesia and
Thailand.
Material examined: INDONESIA, Bali, south
ridge of Mt. Catur, east side of Lake Bratan, elev. 1200–
1800 m, 17 Jan 1999, D. E. Desjardin 6951 (Holotype:
BO; Isotype: SFSU); Java, Cibodas Botanical Garden,
trail to Mt. Gede, S06° 44.675′, E107° 00.423′, elev.
1406 m, 13 Jan 2005, Kerekes 103 (BO, SFSU).
THAILAND, Chiang Mai Province, Huai Nam Dang
National Park, N19°18.29′, E98°35.88′, elev. 1538 m, 28
June 2004, Kerekes 38 (CMU, SFSU).
Notes: Crinipellis furcata is distinctive
because of the fibrillose to squamulose, dark
brown to black pileus and stipe, the distinctly
bifurcated cheilocystidia with long apical
projections, and a negative reaction of pileus
hairs or stipe tissue in KOH. This species is
similar to C. corvina Har. Takah., described
recently from Japan, but the latter species has a
purplish black pileus, and consistently thick-
walled cheilocystidia that rarely bifurcate
(Takahashi, 2000). Crinipellis mirabilis differs
from C. furcata by forming a reddish brown to
violet brown pileus with hairs that turn green in
KOH, and has simple cheilocystidia.
Crinipellis furcata is also phenetically similar
to C. atrobrunnea and C. pseudostipitaria
subsp. orientalis, however, both of the latter
species have larger basidiospores, and grow on
grasses.
11. Crinipellis hepatica Corner, Beih. Nova
Hedwigia 111: 119. 1996.
Type: SINGAPORE, Bukit Timah,
Corner s.n., 27 March 1943 (herb. Corner, E;
alc. form.)
Description adapted from the protologue
Pileus 7–9 mm diam., convex to plane,
often acutely umbonate at first with umbo
disappearing in age, faintly sulcate, opaque,
villous in the center, hoary with decumbent
white fibrils toward the margin, brown rufous
or liver-rufous. Context thin, paler than pileus
surface. Lamellae adnexed to sinuate,
subventricose, scarcely crowded (14–17) with
3 series of lamellulae, pale flesh coloured
becoming deeper pinkish ochraceous, paler
towards the white edge. Stipe 7–10 × 0.7–1 mm,
cylindrical, subpruinose at the apex to
fibrillose-puberulous at the base; apex pallid
becoming pinkish ochraceous to bright rufous
near the base. Odor and taste none.
Basidiospores 9–11 × 4.5–5.5 μm,
subcylindrical, smooth, hyaline. Basidia 4-
spored. Basidioles subacerose. Gill edge sterile.
Cheilocystidia with main body 25–45 × 8–11
μm, subventricose to subfusiform, obtuse to
acute, without appendages. Pileipellis cells of
repent hyphae with rufous brown incrustations,
giving rise to terminal hairs. Hairs –500 × 5–8
μm, or longer, scarcely tapered to the obtuse or
subcapitate apices, lacking secondary
septations, thick-walled, hyaline to pale yellow.

124
Habitat: On dead sticks in the forest.
Known distribution: Singapore.
Material examined: Type was not available for
study. Notes: We have not examined the type
specimen stored at E in alcohol-formalin
solution, nor have we encountered the species
in fresh condition; therefore our concept of the
species is based entirely on the protologue.
Corner indicated that C. hepatica was similar
to C. omotricha (Berk.) D.A. Reid, first
described from South Africa, but the latter
species grows on decayed grasses and has a
greyish to pinkish brown pileus (Pegler, 1986).
12. Crinipellis aff. iopus Singer, Rev. Mycol.
(Paris) 4: 63. 1939. (Figs 11: 1–4, 19)
Type: Collected by Nevodovski in the
Botanical Garden of Alma Ata, Kazakhstan,
U.S.S.R., May 1937 (LE).
Macromorphological description adapted
from the protologue (Singer 1942), and from
photographs of recently collected material
Pileus 6–12 mm diam., convex-
umbilicate, margin initially incurved, sulcate to
the disc, bright lilac when young, becoming
white with age with pale fulvous-brown striae.
Context thin, white. Lamellae narrowly
adnexed, distant with 2 series of lamellulae,
white. Stipe 30–35 × 1 mm, cylindrical, equal,
base scarcely thickened, insititious,
pulverulent-velutinous; bright lilac when fresh,
white at the apex and brownish grey
downwards when dried. Rhizomorphs not
mentioned. Odor and taste none.
Micromorphological description based on
fresh material
Basidiospores 9–12 × 4–5 μm [xmr =
10.2–11.1 × 4.0–4.4 μm, xmm = 10.5 ± 0.5 × 4.3
± 0.2 μm, Q = 2.0–3.0, Qmr = 2.3–2.4, Qmm =
2.5 ± 0.1; n = 25 spores per 3 collections],
ellipsoid to subcylindrical, smooth, hyaline,
inamyloid, thin-walled. Basidia 25–32 × 5–7
μm, clavate, 4-spored, with sterigmata 3–5 μm
long. Basidioles clavate to cylindrical.
Pleurocystidia absent. Gill edge hetero-
morphous. Cheilocystidia common; main body
12–26 × 5–12 μm, clavate to broadly clavate,
hyaline, inamyloid, thick-walled with 0–6
apical appendages; apical appendages 1–11 μm
long, hyaline, inamyloid, thin walled.
Pileipellis a cutis with chains of inflated cells
25–68 × 9–11 μm, clamped, with distinct
brown incrustations in water, incrustations
hyaline in KOH, incrustations and inflated cells
dextrinoid, giving rise to terminal hairs with
basal clamp connections. Hairs few, 125–275 ×
3–6.5 μm, cylindrical, apices generally
tapering and rounded, lacking secondary
septations, thick-walled, inamyloid to weakly
dextrinoid, hyaline to light brown in water,
hyaline in KOH. Pileus trama regular; hyphae
3–5 μm diam., hyaline, inamyloid, thin-walled.
Lamellar trama regular; hyphae 3–6 μm diam.,
cylindrical, hyaline, inamyloid, thin-walled.
Stipe tissue monomitic, parallel, tightly packed;
stipitipellis composed of repent cortical hyphae
and terminal hairs; cortical and medullary
hyphae indistinguishable, 2–5 μm diam.,
cylindrical, hyaline to light brown, cortical
hyphae strongly dextrinoid; terminal hairs
similar to pileipellis hairs, 100–188 × 5–7.5 μm,
thick-walled, inamyloid to weakly dextrinoid,
hyaline to light brown in water and KOH.
Clamp connections present.
Habitat: On twigs in disturbed primary
and secondary rainforests.
Known distribution: Botanical Garden of
Alma Ata, Kazakhstan, and Papua New Guinea.
Material examined: PAPUA NEW GUINEA.
Madang, Baitabag, Kau Wildlife Area, S5˚09', E145˚06',
elev. ± 170 m, 2 February 1997, R. Walleyn 829
(GENT); Madang, Silibob, S5˚12’, E145˚45’, elev. ±
100 m, 29 January 1997, R. Walleyn 774 (GENT);
Madang, Surinamvalley, Boroki Lake, S5˚52', E145˚51',
elev. 600–750 m, A. Verbeken 97–312 (GENT); Madang,
left of bridge, Binek River, S5˚12', E145˚45' ,elev. 100
m, 24 February 1997, A. Verbeken 97–404 (GENT).
Notes: We are tentatively identifying the
Papua New Guinea specimens as representing
Crinipellis iopus, described originally from a
Botanical Garden in Kazakhstan (Singer, 1939,
1942). Whether the species is indigenous to
Kazakhstan or was introduced into the
botanical garden with plants from somewhere
else is not known. Singer (1976) designated C.
iopus the type species of the section Iopodinae,
an infrageneric group established to
accommodate brightly pigmented species (red,
crimson, purple, lilac, violet) with pileus hairs
that do not turn green or grey in KOH.
Unfortunately, the protologue does not provide
details regarding the cheilocystidia or the

Fungal Diversity
125
Fig. 11. Crinipellis aff. iopus (RW 829). – 1. Basidiospores. – 2. Basidia and basidioles. – 3. Cheilocystidia (RW 829,
AV 97-312). –4-5. Pileipellis hairs. – Bar: 1-4 = 10 µm; 5 = 25 µm.
hyphae subtending the pileipellis hairs, both
taxonomically important characters. Until
further material of this intriguing species is
located in Kazakhstan and compared with the
Papua New Guinea material, our determination
will remain tentative.
In the ML tree, C. aff. iopus was sister of
Moniliophthora but with low statistical support,
and its placement in the phylogenetic analyses
made Crinipellis non-monophyletic. Interes-
tingly, Singer (1976) included species currently
accepted in Moniliophthora (e.g., M. perni-
ciosa) in his sect. Iopodinae and our molecular
data support this contention (Fig. 1). In C. aff.
iopus, the chains of inflated, incrusted hyphae
subtending the uncommon pileipellis hairs are
very similar to the pileipellis anatomy of
Moniliophthora species, as is the KOH reaction,
and suggest transfer to Moniliophthora may be
warrented. A positive outcome of such an
approach would be a monophyletic Crinipellis
based on ITS data. Until further genes are
sequenced, more taxa are added to the dataset,
and the Papua New Guinea taxon is positively
identified, formal transfer of C. iopus is not
justified.
13. Crinipellis malesiana Kerekes, Desjardin
and Vikineswary sp. nov.
(Figs 12: 1–7, 19)
MycoBank: 512275
Etymology: Malesia – the region including the
Malay Peninsula and Indonesia; malesiana refers to the
distribution of this species in Malesia.
Pileus 2–13 mm latus, convexus, brunneus.
Lamellae adnexae. Stipes 4–22 × 1–2 mm, cylindricus,
fibrillosus, brunneus. Odor saporque nulli.
Basidiosporae 8–12.5 × 4–6.5 μm, oblongatus, hyalinae.
Pleurocystidia 30–50 × 5–9.5 μm, clavata, hyalinae.
Cheilocystidia (11–) 21–40 × 4–9 μm, clavata, 1–3 apice
surculis. Trama lamellarum regularis. Caulocystidia 22–
45 × 5–8 μm, cylindraceis. Holotypus: INDONESIA.

126
Fig. 12. Crinipellis malesiana (AR 491). – 1. Basidiomes (x2). – 2. Basidiospores. – 3. Basidioles. – 4. Cheilocystidia.
–5. Caulocystidia.– 6. Pleurocystidia. – 7. Pileipellis hair. – Bar: 2-7= 10 µm.
Central Java, Kabupaten Jepara, Keeawatan
Karimunjawa, Desa Kanimunjawa, Kawpung alang-
alang, Legon Goprak, 13 June 2004, A. Retnowati 491
(BO). Pileus 2–13 mm diam., convex and
shallowly umbilicate or papillate when young,
becoming broadly convex and papillate to
umbonate with age, often with one or two
raised concentric ridges surrounding central
papilla, striate to sulcate, dull, dry, radially
fibrillose-hairy to hispid, hairs extending
beyond margin; brown overall when young,
becoming dark brown (7F8) at the disc and
brown to brownish orange at the margin in age.
Context < 1 mm thick, pale brown to yellowish
white. Lamellae adnexed to adnate, close to
crowded with 3–4 series of lamellulae,
yellowish white to white. Stipe 4–22 × 1–2 mm,
central, cylindrical, thickened near base, dry,
dull, fibrillose-hairy to hispid overall,
insititious, brown overall, slightly paler than
the pileus. Rhizomorphs absent. Odor and taste
none.
Basidiospores 8–12.5 × 4–6.5 μm [xmr =
11.1–11.4 × 4.9–5.2 μm, xmm = 11.3 ± 0.1 × 5.0
± 0.2 μm, Q = 1.5–2.7, Qmr = 2.2–2.3, Qmm =
2.2 ± 0.1; n = 25 spores per 3 collections],
phaseoliform in side view, oblong in frontal
view, smooth, hyaline, inamyloid, thin- to
thick-walled. Basidia none observed.
Basidioles 15–34 × 5.5–8 μm, clavate to
cylindrical with rounded to subacute apex.
Pleurocystidia 30–50 × 5–9.5 μm, projecting
up to 15 μm beyond basidioles, clavate to
fusoid, hyaline, inamyloid, thick-walled, not
refractive. Cheilocystidia abundant; main body
(11–) 21–40 × 4–9 μm, clavate to broadly
clavate or cylindrical, hyaline, inamyloid, thin-
to thick-walled, majority entire, few with 1–3

Fungal Diversity
127
apical appendages 1–3 μm long. Pileipellis a
cutis of hyphae 5–6 μm diam., slightly inflated,
hyaline to light brown, inamyloid to dextrinoid,
slightly thick-walled, giving rise to terminal
hairs with basal clamp connections. Hairs 175–
555 × 2.5–5 μm, cylindrical, apices mostly
rounded, few with acute apices, with secondary
septations, thick-walled (2–3 μm), weakly
dextrinoid, becoming yellowish green to green
in KOH. Pileus trama subregular; hyphae 2–8
μm diam., hyaline, inamyloid to weakly
dextrinoid near cutis, slightly thick-walled.
Lamellar trama subregular; hyphae 2–8 μm
diam., hyaline, inamyloid, slightly thick-walled.
Stipe tissue monomitic, parallel, tightly packed;
stipitipellis composed of repent cortical hyphae,
caulocystidia and terminal hairs; cortical
hyphae 2.5–5 μm diam., cylindrical, thick-
walled, dextrinoid, becoming light pea green in
KOH; medullary hyphae 2–5 μm diam.,
cylindrical, thin-walled, inamyloid to weakly
dextrinoid, hyaline; caulocystidia 22–45 × 5–8
μm, thick-walled cylindrical; terminal hairs
72–440 × 5–10 μm, cylindrical, apices
narrowly rounded, thick-walled, inamyloid to
weakly dextrinoid, hyaline to light brown in
water, hyaline to light yellowish green in KOH.
Clamp connections present.
Habitat: Scattered to gregarious on
woody debris.
Known distribution: Indonesia and
Malaysia, Thailand.
Material examined: INDONESIA. Central Java,
Kabupaten Jepara, Keeawatan Karimunjawa, Desa
Kanimunjawa, Kawpung alang-alang, Legon Goprak, 13
June 2004, A. Retnowati 491 (Holotype: BO; Isotype:
SFSU); Central Java, Kabupatau Jepara, Keeamatan
Karimunjawa, Desa Karimunjawa, Legan Lele, 15 June
2004, A. Retnowati 513 (BO; SFSU). MALAYSIA.
Kedah, Langkawi Island, Gunung Raya, 13 April 2003,
KUM 084a (KLU-M, SFSU); Keha, Langkawi Island,
Matchinchang, 1 September 2004, TYS 346 (KLU-M,
SFSU).
Notes: Crinipellis malesiana is
characterized by the following combination of
features: a dark brown pileus and brown robust
stipe; simple, entire cheilocystidia; clavate to
fusoid pleurocystidia; pileus hairs and stipe
tissue that turn yellowish green in KOH; and
elongate-ellipsoid basidiospores with Qmm =
2.2. The greening reaction of the tissues in
KOH suggests that this species belongs in
section Grisentinae as established by Singer
(1942, 1976). Crinipellis malesiana is similar
to C. atrobrunnea, another species with a dark
brown pileus and relatively long and narrow
spores (Q = 2.1), described from Vietnam.
Cheilocystidia were not recovered in our type
analysis of C. atrobrunnea (FH!) nor were they
reported in Yang’s (2000) type study; however,
cheilocystidia were reported by Singer (1942)
as cylindric or subfusoid with no branching, or
branching up to five or more times (Singer,
1976). Crinipellis malesiana is distinguished
from C. atrobrunnea by the presence of
pleurocystidia, and the greening reaction of
pileus hairs and stipe tissues. In addition, C.
atrobrunnea is reported only to grow on
Gramineae, whereas C. malesiana was found
only on wood and dicot twigs. Crinipellis
mirabilis Singer, described from New
Caledonia, also has a brown pileus and hairs
that turn green in KOH. It is distinguished from
C. malesiana by forming slightly smaller
basidiospores (9.5 × 4.4 µm), by lacking
distinctive pleurocystidia, and has hairs that are
initially purple in KOH but fade to greyish
green in time. Crinipellis hepatica Corner from
Singapore is also similar to C. malesiana, but
the former differs by forming reddish brown
pilei, lacks pleurocystidia, and there is no
mention of a green KOH reaction to pileus
hairs. The four collections of C. malesiana
studied by us and included in the molecular
analyses form a clade with 100% MP-BS,
100% ML-BS and 1.00 PP support, and sister
of C. scabella and C. trichialis in all analyses.
One sequence of an unidentified Crinipellis
collected in Thailand was downloaded from
GenBank (AY916698). The latter sequence
also nested in the C. malesiana clade with
100% BS and PP support, suggesting strongly
that it represents C. malesiana. We have not
studied voucher material associated with this
undetermined GenBank entry. However, these
data suggest that the species may range
northward from Indonesia to Thailand.
14. Crinipellis aff. mirabilis Singer, Lilloa 8:
497. 1943. (Fig. 13: 1–7)
Type: NEW CALEDONIA. Between Col
d’ Amieu and Mégropo, collected by Le Rat
#101, 1907, on twigs, det. by Patouillard as
Crinipellis stipitaria var. castaneus (Holotype:
FH!).

128
Fig. 13. Crinipellis aff. mirabilis (JFK 137). – 1. Basidiomes (a. x3. – b-c. x2.). – 2. Basidiospores. – 3. Basidioles. – 4.
Cheilocystidia. – 5. Caulocystidia. – 6. Pleurocystidia. – 7. Pileipellis hairs. – Bar: 2-7 = 10 µm.
Analysis of the holotype specimen
The holotype specimen consists of three
intact basidiomes, one loose in the packet and
two attached to woody sticks. As dried: Pileus
4–5 mm diam., conical-papillate, with long
shaggy hairs overall, dark brown or reddish
brown. Lamellae subdistant, seemingly narrow
but most are missing, pallid. Stipe 25–30 × 0.5
mm, pubescent, dark brown, arising directly
from the woody substrate, associated with very
narrow, wiry, branched, finely pubescent, dark
brown rhizomorphs.
Basidiospores (adhered to pileus hairs
and stipe apex) 8.3–10.5 (–12) × 4.2–4.8 μm,
ellipsoid to subfusoid, hyaline, smooth,
inamyloid, thin-walled or in age thick-walled.
All hymenial cells were collapsed and
degraded; no data on hymenial elements
obtainable. Pileipellis a cutis giving rise to
hairs. Hairs 3.2–6.5 μm diam., cylindrical with
an acute or obtuse apex, thick-walled (0.5–1.5
μm), sinuous at the base, with regular, ladder-
like secondary septa; walls dark brown to
reddish brown in water but immediately
greyish olive in 3% KOH, dextrinoid.
Description based on fresh material
Pileus 1–5 mm diam., when young
convex with inrolled margin, papillate, in age
becoming convex to plano-convex with papilla
in a depressed disc, with 1–3 raised concentric
ridges around disc, papilla and ridges less
obvious in older specimens; surface radially
appressed-fibrillose, margin with appendiculate
fibrils; dark reddish brown (7–8F6–8) to violet
brown (11F7) when young, in age becoming
lighter in colour in zone around the disc to
cream at the margin with violet brown (11F7)
fibrillose hairs. Context < 1 mm thick.
Lamellae free to adnexed or narrowly adnate,
close to subdistant with 1–2 series of
lamellulae, narrow, white, non-marginate. Stipe
8–22 × 0.5–1 mm, central, terete, cylindrical, ±
equal, tough, pliant, longitudinally striate,
fibrillose, subinsititious; dark reddish brown
(7–8F6–8) to violet brown (11F7). Rhizo-
0morphs thin, hair-like, glabrous, black. Odor
and taste none.
Basidiospores 8–12 × 3–5 μm [xmr = 9–
10 × 3.8–4.8 μm, xmm = 9.5 ± 0.5 × 4.4 ± 0.5
μm, Q = 1.8–2.9, Qmr = 2.0–2.4, Qmm = 2.2 ±

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129
0.2; n = 11–25 spores per 3 collections],
subcylindrical to amygdaliform in side view,
few oblong to ellipsoid, smooth, hyaline,
inamyloid, thin-walled. Basidia 19–26 × 5–7.5
μm, clavate, 4-spored. Basidioles clavate to
cylindrical with a few fusiform to subfusiform.
Pleurocystidia absent. Lamellar edge sterile.
Cheilocystidia common; main body 26–30 × 5–
10 μm, clavate to narrowly ventricose, entire,
simple, hyaline, inamyloid, thin-walled or with
walls up to 0.5 µm thick. Pileipellis a cutis
25–50 × 6–7.5 μm, clavate, slightly thick-
walled, weakly dextrinoid, hyaline, giving rise
to terminal hairs with basal clamp connections.
Hairs 87–337 × 2.5–5 μm, cylindrical, very
rarely branched, apices rounded to acute, with
secondary septations, thick-walled, dextrinoid,
purple-violet in water, initially dark purple in
KOH, colour fading with time and becoming
grey to olive-grey in KOH. Pileus trama
subregular to regular; hyphae 3–6 μm diam.,
thin-walled, hyaline in water, distinctly olive in
KOH. Lamellar trama regular; hyphae 2–6 μm
diam, thin-walled. Stipe tissue monomitic,
parallel, tightly packed; stipitipellis composed
of repent cortical hyphae, caulocystidia and
terminal hairs; cortical hyphae 1.5–3 μm diam.,
cylindrical, thick-walled, dextrinoid, hyaline to
light yellow with greenish tints in KOH;
medullary hyphae 1.5–2 μm diam., cylindrical,
thick-walled, strongly dextrinoid, hyaline to
light yellow with greenish tints in KOH;
caulocystidia 57–82 × 5–8.75 μm, cylindrical,
some branched, base flexuous, thick-walled,
dextrinoid, hyaline to greyish purple in water;
terminal hairs 162–327 × 5–6 μm, cylindrical,
apices rounded or acute, with secondary
septations, thick-walled, dextrinoid, greyish
purple in KOH, fading to greyish green in time.
Clamp connections present.
Habitat: Scattered on twigs of
undetermined dicots in primary forest, and on
decomposing dicot leaves (and possibly woody
roots) in termite mounds in a bamboo
secondary forest.
Known distribution: Malaysia, New
Caledonia, and Thailand.
Material examined: MALAYSIA. Kuala Lumpur,
Gombak Field Station, 12 January 2003, Desjardin 7521
(SFSU). NEW CALEDONIA. Between Col d’ Amieu
and Mégropo, collected by Le Rat # 101, 1907, on twigs,
det. by Patouillard as Crinipellis stipitaria var. castaneus
(Holotype: FH). THAILAND. Chiang Mai Province,
Mae Taeng Dist., Si Lanna National Park, 4 July 2005,
JFK 137 (SFSU).
Notes: Crinipellis mirabilis, described
originally from woody debris in New
Caledonia, is a poorly known species
represented by a holotype specimen consisting
of three basidiomes in rather poor condition
and missing most of their lamellae (FH!). We
tentatively recognize the Thailand and
Malaysian specimens as C. aff. mirabilis. Our
collections differ from the protologue in
forming slightly smaller basidiomes with more
violet-brown pigmentation instead of "chestnut
brown" or "Mars brown", and growth on leaves
instead of twigs. Micromorphologically, the
Thai and Malaysian specimens match quite
closely the holotype specimen of C. mirabilis.
Singer reported the basidiospores as 8.5–9 ×
3.8–4.2 µm in the protologue, whereas our
analysis of the holotype specimen yielded
basidiospores in the range 8.3–12 × 4.2–5.0 µm.
Crinipellis mirabilis is the type species of
section Grisentinae Singer (1976), a group
characterized by hairs that turn green in KOH.
15. Crinipellis pseudostipitaria subsp.
orientalis Singer, Lilloa 8: 472. 1943.
(Fig. 14: 1–2)
Type: VIETNAM (TONKIN), La Pho, 23
July 1909, coll. by Demange #379, on grass
debris, det. by Patouillard as Crinipellis
stipitaria (Holotype: FH!).
Macromorphological description adapted
from the protologue:
Pileus 3–13 mm diam., umbonate, with
or without a depression around the papilla,
hairy, tan to brown. Lamellae attenuated-
adnexed to mostly free, moderately crowded to
subdistant, medium broad, probably white
when fresh. Stipe about three times as long as
the diameter of the pileus, equal, solid, hairy,
concolorous with the pileus.
Analysis of the holotype specimen
The holotype specimen consists of four
fragmented basidiomes in poor condition,
attached to grass stems; separated by Singer
from another different species that was part of
the original Demange collection. The 4
basidiomes are in a separate packet labeled
"type." As dried: Pileus 1–3 mm diam,
umbonate, silky-shaggy, pallid beige. Lamellae

130
Fig. 14. Crinipellis pseudostipitaria subsp. orientalis
(Holotype). – 1. Basidiospores. – 2. Pileipellis hairs. –
Bar: 1-2 = 10 µm.
absent; all removed. Stipe 5–8 × 0.5 mm,
pubescent, cream-beige, arising directly from
the substrate. Rhizomorphs absent.
Basidiospores 7.7–9.5 × 5–6 μm [xm =
8.3 ± 0.5 × 5.5 ± 0.3 μm, Q = 1.5–1.3, Qm = 1.7
± 0.3; n = 15 spores per 1 collection], broadly
ellipsoid, smooth, hyaline, inamyloid, thin-
walled, rarely thick-walled. No hymenial
elements observable; all lamellae absent.
Pileipellis a cutis giving rise to hairs. Hairs 4–
7 μm diam., cylindrical, thick-walled (1.3–3.2
μm) to nearly solid with only a tiny or
occluded lumen slightly sinuous near the base,
narrowing slightly and abruptly near the apex,
broadly obtuse; secondary septa not seen,
hyaline to yellowish brown, inamyloid to
dextrinoid; base of some hairs with granular to
plaque-like, thin incrustations, mostly non-
incrusted. Stipitipellis similar to the pileipellis
with similar hairs.
Habitat: On grass debris.
Material examined: Type: VIETNAM
( TONKIN), La Pho, 23 July 1909, coll. by Demange
#379, det. by Patouillard as Crinipellis stipitaria (FH;
holotype).
Notes: Crinipellis pseudostipitaria subsp.
orientalis is distinctive because of the grass
substrate, broadly ellipsoid spores 5–6 μm in
diam, and cylindrical, nearly solid hairs.
Because no lamellae remained in the holotype
specimen, data on cystidia so important in
Crinipellis taxonomy were unobtainable.
Singer (1942) reported the cheilocystidia as
rarely simple, with the majority “more or less
forked or branched or even almost echinate,”
22–30 × 6–7.5 μm. We have not encountered
fresh material of this taxon so we can provide
no data on hymenium cell morphology.
According to Singer, C. pseudostipitaria subsp.
orientalis is the Eastern Hemisphere’s broad-
spored form of C. pseudostipitaria subsp.
pseudostipitaria (ut subsp. occidentalis Singer,
nom. invalid) from the Western Hemisphere,
which has narrower spores. Singer (1942)
described C. pseudostipitaria var. mesites
Singer as a form transitional between the two
subspecies.
16. Crinipellis sepiaria Pat. & Demange [ut
saepiarius], Bull. Soc. Mycol. Fr. 26: 36. 1910.
(Fig. 15: 1–3)
≡ Marasmius sepiarius (Pat. & Demange) Sacc.
& Trotter. [ut saepiarius], Syllog. Fung. 21: 112. 1912.
Type: VIETNAM (TONKIN), Hanoi,
Sept. 1908, coll. by M. Demange #290, on
herbaceous debris and bamboo (Holotype:
FH!). Macromorphological description
adapted from the protologue and from Singer
(1942):
Pileus 10–21 mm diam., convex to
plano-convex, umbilicate or depressed at the
disc, zonate, radially fibrillose-hairy, margin
ciliate; bister, not darker than "snuff brown" or
"sayal Brown" when dried (greyish to grey
according to Yang, 2000). Lamellae narrowly
adnexed to subfree, subdistant to distant, thin,
moderately broad, white. Stipe 20–30 × 1–2
mm, rigid, cylindrical, fibrillose-squamulose to
hirsute, brown at the apex becoming brown to
black at the base. Rhizomorphs absent.
Fig. 15. Crinipellis sepiaria (Holotype). – 1.
Basidiospores. – 2. Basidia and basidioles. – 3.
Pileipellis hairs. – Bar: 1-3 = 10 µm.
Analysis of the holotype specimen
The holotype specimen consists of 6
fragmented basidiomes, pressed flat, in fair
condition. As dried: Pileus 8–18 mm diam,
campanulate with two concentric ridges and
depressions surrounding a small umbo, shaggy,

Fungal Diversity
131
brown to almost dark brown. Lamellae
subdistant, broad, pale brown. Stipe 12–18 ×
0.75–1.5 mm, appressed-fibrillose, dark brown.
Basidiospores 5.7–7.7 × 4.5–5.7 μm [xm
= 6.6 ± 0.6 × 5.2 ± 0.4 μm, Q = 1.2–1.5, Qm =
1.3 ± 0.1; n = 20 spores per 1 collection],
broadly ellipsoid to ovoid, smooth, hyaline,
inamyloid, thin-walled. Basidia 4-spored,
clavate, clamped. Basidioles subfusoid.
Pleurocystidia absent. Lamellar edges
colonised by a deuteromycete; cheilocystidia
not observable, none recovered. Pileipellis a
cutis giving rise to hairs. Hairs > 300 × 3–6.5
μm, cylindrical, apex not appreciably narrowed,
obtuse; thick-walled (0.5–2.3 μm), hyaline to
yellowish brown in water, yellowish olive in
3% KOH, dextrinoid; apex of hairs non-
incrusted, base of hairs and some subtending
hyphae with brown, granular incrustations, a
few with secondary septations.
Habitat: On herbaceous debris and
bamboo in bamboo groves.
Known distribution: Vietnam
Material examined: VIETNAM (TONKIN),
Hanoi, Sept. 1908, coll. by M. Demange #290, on
herbaceous debris and bamboo (Holotype: FH).
Notes: We have not encountered fresh
material of this species so our concept is based
on examination of the holotype specimen and
data provided in the protologue and in Singer
(1942). Crinipellis sepiaria is distinctive
because of the small ovoid spores, large pilei
(up to 20 mm when fresh), and growth on
herbaceous debris and bamboo. Our type study
matches the data provided by Yang (2000).
Singer (1942) mentioned that there is a
painting of the type collection by Nguyen
Manh Hoan preserved in the Farlow Reference
Library, Harvard University. This illustration is
reproduced in color microfiche in
"Champignons du Tonkin" with an introduction
by Don Pfister (1985).
17. Crinipellis setipes (Peck) Singer, Lilloa 8:
493. 1943. (Fig. 16: 1–6)
= Agaricus stipitarius var. setipes Peck, N.Y. St.
Mus. Rep. 38: 109. 1885.
Pileus 2–9 diam. × 1–2 mm tall,
hemispherical to convex and subumbilicate
when young, becoming convex to plano-
convex, subumbilicate with a small papilla that
becomes less obvious with age, radially
fibrillose, margin distinctly serrulate; disc dark
brown (7F7), margin brown (7E7) to cream
(4A3), becoming paler with age. Context < 1
mm thick, light brown (7D6). Lamellae free to
adnexed, subdistant to close with 2 series of
lamellulae, white. Stipe 16–100 × < 1 mm,
central, cylindrical, fibrillose, more densely
fibrillose near base, light brown with cream-
colored (4A3) fibrillose hairs at the base.
Rhizomorphs thin, black to dark brown (7F8).
Odor and taste none. Basidiospores 7–9 × 4–5
μm [xm = 8.2 ± 0.6 × 4.5 ± 0.5 μm, Q = 1.4–2.3,
Qm = 1.9 ± 0.2; n = 25 spores per 1 collection],
ellipsoid, smooth, hyaline, inamyloid, thin-
walled. Basidia 16–24 × 4–6 μm, clavate, 4-
spored. Basidioles clavate. Pleurocystidia
absent. Gill edge sterile. Cheilocystidia
common; main body 12–15 × 4–5 μm, clavate,
hyaline, inamyloid, thin-walled, with 4–7
apical appendages; apical appendages 1–3 μm
long, hyaline, inamyloid, thin walled.
Pileipellis a cutis of inflated cells 9–12 μm
diam, light tan-brown, giving rise to terminal
hairs with basal clamp connections. Hairs 50–
437 × 4–7.5 μm diam, cylindrical, tapered to
acute apices, with few secondary septations,
thick-walled (1–2 μm), weakly dextrinoid, light
tan-brown in KOH. Pileus trama subregular;
hyphae 3–5 μm diam., hyaline, inamyloid,
thin-walled. Lamellar trama regular; hyphae
2–5 μm diam., hyaline, inamyloid, thin-walled.
Stipe tissue monomitic, parallel, tightly packed;
stipitipellis composed of repent cortical hyphae
and terminal hairs; cortical hyphae 2–3 μm
diam., cylindrical, thick-walled, strongly
dextrinoid, light brownish-green in KOH;
medullary hyphae 3–5 μm diam., cylindrical,
thin-walled, dextrinoid to strongly dextrinoid,
hyaline in KOH; terminal hairs 77–450 × 5–
7.5 μm, cylindrical, thick-walled, strongly
dextrinoid, hyaline to light tan-brown in water
and KOH. Clamp connections present.
Habitat: On woody debris in montane
cloud forest.
Known distribution: China, Thailand,
Northeastern North America.
Material examined: THAILAND. Chiang Mai
Province, Doi Inthanon National Park, Summit of Doi
Inthanon, Nature Trail, N18˚35.35', E98˚29.22', elev.
2542 m, 25 June 2004, Kerekes 34 (CMU, SFSU).
Notes: Crinipellis setipes, described
from northeastern North America and reported
from China (Singer 1942) was collected at the

132
Fig. 16. Crinipellis setipes (JFK 34). – 1. Basidiomes (x2). – 2. Basidiospores. – 3. Basidia and basidioles. – 4.
Cheilocystidia. –5. Caulocystidia.– 6. Pileipellis hair. – Bar: 2-4, 6 = 10 µm; 6 = 25 µm.
summit of Doi Inthanon, the tallest point in
Thailand (at 2542 meters), among vegetation
similar to eastern Himalayan vegetation. This
species is nearly indistinguishable from C.
piceae Singer, and we are following Singer
(1942, 1976) and Redhead (1986, 1989) in
recognizing the two entities as distinct species.
Crinipellis setipes differs from C. piceae by
lacking an obvious basal disc, by growth on
dicotyledonous leaf litter and woody debris
instead of on spruce needles, and by forming a
longer stipe. According to Redhead (1989) C.
piceae has a circumboreal-bicoastal distri-
bution, whereas C. setipes has an amphi-
Pacific/asian-Appalachian distribution. Our
Thai report of C. setipes expands the known
geographic distribution of C. setipes and
confirms an asian-Appalachian distribution.
Crinipellis procera (Stevenson 1964) is also
similar to C. setipes in regards to basidiome
size, spore size and cheilocystidia morphology,
but differs in forming a conical pileus with a
conical papilla compared to an umbilicate
pileus in C. setipes, and lacks rhizomorphs (see
type study #31 below). DNA extractions and
sequencing were unsuccessful from two
specimens of C. setipes from North America
[(DED 4644 and DED 4318 from Tennessee
(both SFSU)]. The Thai specimen of C. setipes,
collected on undetermined dicot twigs falls in a
clade with North American specimens of C.
piceae from spruce needles with 100 % MP-BS,
98% ML-BS and 1.0 PP support (Fig. 1), and
was sister to one of the C. piceae specimens
(DED 7758) in the parsimony analysis (data
not shown), suggesting possibly that the two
species are conspecific. Sequences of
northeastern North American C. setipes
populations need to be added to the analyses to
clarify their taxonomic status. We have
identified our Thai material as C. setipes
mainly because of the dicotyledonous wood
substrate.
18. Crinipellis tabtim Kerekes, Desjardin &
Lumyong sp. nov. (Figs 17: 1–6, 19)
MycoBank: 512276
Etymology: tabtim (Thai) = ruby-colored,
referring to the pileus color.
Pileus 1–11 mm latus, primo hemisphaericus,
dein convexus, rubineus. Lamellae adnexae. Stipes 15–
30 × < 1 mm, cylindricus, fibrillosus, rubrobrunneus.
Odor saporque nulli. Basidiosporae 8–11 × 4–5.5 μm,
ellipsoideae, hyalinae. Basidia clavata, hyalinae,
tetraspora. Pleurocystidia nulla. Cheilocystidia 10–20 ×
5–9 μm, clavata, 3–12 apice surculis. Trama lamellarum
regularis. Caulocystidia 88–228 × 3–6 μm, cylindraceis.
Holotypus: THAILAND. Chiang Mai Province, Doi
Inthanon National Park, 27 June 2005, JFK 129
(Holotype: CMU).
Pileus 1–11 mm diam., when young
hemispherical to convex with inrolled margin,
shallowly umbilicate, becoming plano-convex
with concentric ridges around the umbilicus in

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133
Fig. 17. Crinipellis tabtim (JFK 129). – 1. Basidiomes (a. x3. – b-c. x2). – 2. Basidiospores. – 3. Basidium and
basidioles. – 4. Cheilocystidia. – 5. Caulocystidium. – 6. Pileipellis hair. – Bar: 2-4, 6, 7a = 10 µm; 5 = 50 µm.
age, dull, dry, radially appressed-fibrillose; disc
dark violet brown to dark ruby (10–12F7),
margin violet brown to ruby (10D–E4), fading
to almost cream-coloured at the margin in age.
Context < 1 mm thick, white, unchanging.
Lamellae narrowly adnexed to free, close with
2–3 series of lamellulae, white. Stipe 15–30 ×
< 1 mm, central, terete, cylindrical, equal
above, subbulbous at base, longitudinally
striate, twisted-fibrous, insititious, dull, dry,
fibrillose to hirsute; reddish brown (9E5) to
light brown (6D7). Rhizomorphs No obvious
rhizomorphs, however, some collections with
the presence of a white mycelial pad on the
leaves. Odor and taste none.
Basidiospores 8–11 × 4–5.5 μm [xmr =
9.1–9.3 × 4.0–4.1 μm, xmm = 9.2 ± 0.1 × 4.1 ±
0.1 μm, Q = 1.8–2.8, Qmr = 2.2–2.3, Qmm = 2.3
± 0.9; n = 25 spores per 4 collections],
lacrymoid to elongate ellipsoid, smooth,
hyaline, inamyloid, thin-walled. Basidia
clavate, 4-spored, hyaline, inamyloid, thin-
walled. Basidioles clavate with rounded to
subacute apex. Pleurocystidia, absent.
Cheilocystidia common; main body 10–20 × 5–
9 μm, clavate, hyaline, inamyloid, thin-walled,
with 3–12 finger-like apical appendages; apical
appendages 1–13 μm long, hyaline, inamyloid,
thin-walled. Pileipellis a cutis of inflated cells
4–10 μm diam., giving rise to terminal hairs
with basal clamp connections. Hairs 70–140 ×
2.5–8 μm, cylindrical, mostly with acute apices,
with secondary septations, thick-walled,
dextrinoid, burgundy in water, purplish-grey in
KOH and becoming slate blue with time, with
golden reddish brown to burgundy
incrustations on pileipellis cells and hairs.
Pileus trama subregular; hyphae 3–7(–18) μm
diam., hyaline, inamyloid, thin-walled.
Lamellar trama interwoven; hyphae 3–5.5 μm
diam., cylindrical, smooth, hyaline, inamyloid,
thin-walled. Stipe tissue monomitic, parallel,
tightly packed; stipitipellis composed of repent
cortical hyphae and terminal hairs; cortical and
medullary hyphae 2–5 μm diam., cylindrical,
hyaline to yellow-green, dextrinoid; terminal
hairs 88–228 × 3–6 μm, cylindrical, thin- to
thick-walled, weakly dextrinoid, light brown to
reddish brown with burgundy tints in water,
hyaline to light green in KOH. Clamp
connections present.
Habitat: Scattered on undetermined
decomposing dicot leaves.
Known distribution: Thailand.
Material examined: THAILAND, Nakorn Nayok
Province, Khao Yai National Park, 8 July 2005, Kerekes
141 (CMU, SFSU); Chiang Mai Province, Doi Inthanon
National Park, 27 June 2005, Kerekes 129 (Holotype:
CMU; Isotype: SFSU); same location, 27 June 2005,
Kerekes 134 (CMU, SFSU); Chiang Mai Province, Mae
Taeng District, Tumg Joaw Willage, 30 June 2004,
Kerekes 44 (CMU, SFSU).
Notes: Diagnostic features of Crinipellis
tabtim include: a bright burgundy to violet
brown or dark ruby-colored pileus with a
brown to reddish brown stipe; pileus hairs that
turn slate blue in KOH with time; pileipellis
cells and hairs with burgundy-colored

134
incrustations when mounted in water; and
cheilocystidia with numerous finger-like
projections up to 13 μm in length. The slate
blue reaction of the pileus hairs in KOH
suggests that this species belongs in section
Iopodinae subsection Iopodinae as
circumscribed by Singer (1976). Crinipellis
tabtim is similar to C. purpurea Singer,
described from Colombia, but the latter species
differs in forming pileus hairs that do not
becoming slate blue in KOH, by forming a
deeper purple to purplish black pileus and stipe,
slightly shorter basidiospores (mostly 8.3 ×
3.8–4 µm), and lacks incrustations on
pileipellis hyphae (Singer 1976). Crinipellis
iopus (see above) differs from C. tabtim by
forming a bright lilac pileus that bleaches white,
and by developing larger basidiospores. In
addition, in the ITS analyses (Fig. 1), C. tabtim
and C. aff. iopus are in distant clades; rather, C.
tabtim and C. cupreostipes, two species with
distinctly different morphologies, are sister
species but with low statistical support (59%
MP-BS, 64% ML-BS, .88 PP).
19. Crinipellis trichialis (Lév.) Pat. ex
Antonín, R. Ryoo & H.D. Shin, Mycotaxon
108: 432. 2009. (Figs 18: 1–7, 19)
= Agaricus trichialis Lév., Ann. Sci. Nat., Bot.
sér. 3, 5: 113. 1846.
≡ Naucoria trichialis (Lév.) Sacc., Syllog. Fung.
5: 853. 1887.
Type: JAVA, Zollinger #2078, on stems
of bamboo (Holotype: FH!).
Analysis of the holotype specimen:
The holotype specimen consists of 3
basidiomes in fair condition, one loose in the
packet and two attached to a fragmented
bamboo stem, plus 3 stipes attached to the
same stem. As dried: Pileus ca 3 mm diam.,
conical to campanulate, shaggy, dark brown.
Lamellae close, relatively narrow, dark brown.
Stipe 6–8 × 0.5 mm, non-insititious, tomentose,
dark brown, arising directly from the substrate.
Rhizomorphs absent.
Basidiospores (8.5–) 9.6–11.5 × (5.5–)
6–7 (–7.4) μm [xm = 10.3 ± 0.7 × 6.5 ± 0.5 μm,
Q = 1.3–1.8, Qm = 1.6 ± 0.1; n = 22 spores per
1 collection], broadly ellipsoid, smooth,
hyaline, inamyloid, thin- to thick-walled.
Basidia 27–32 × 6.2–7 μm, clavate, 4–spored,
clamped. Basidioles subfusoid. Pleurocystidia
scattered, clavate to cylindrical, projecting
above basidia, thin- to thick-walled, often with
amorphous yellowish brown to brown apical
incrustations. Cheilocystidia not seen; material
too poor and lacking intact lamellar edges.
Pileipellis cutis giving rise to thick-walled
hairs. Hairs 3.2–7 μm diam. (avg. 5.5 μm),
cylindrical for most their length, apex
gradually narrowed to an obtuse or acute tip,
smooth or roughened near apex, base with
roughened-granular incrustations, brown in
water, olive-brown in 3% KOH, dextrinoid;
walls 0.5–2.8 μm thick; secondary septa not
seen. Stipitipellis similar to pileipellis.
Description based on fresh material
Pileus 5–18 mm diam., hemispherical to
convex-cylindrical when young, becoming
convex to plano-convex or applanate with age,
disc with a tuft of scales forming a small
papilla, with concentric zones of erect fibrils or
scales around the disc, margin fibrillose;
papilla and central zone dark brown (6–7F6–8),
middle zone brown (6–7E6–8), margin
brownish yellow (5C5–8) or paler. Context < 1
mm thick, white. Lamellae adnexed to free,
close to crowded with 2–4 series of lamellulae,
white to pale yellowish white (3A1–2). Stipe
6–27 × 0.5–1.5 mm, central, cylindrical, equal
with an enlarged base, tough, pliant, fibrillose
to hairy overall, insititious, brown to dark
brown. Rhizomorphs absent. Odor and taste
none.
Basidiospores 6–11 × 5–8 μm [xmr =
8.8–9.8 × 5.8–6.3 μm, xmm = 9.3 ± 0.9 × 6.0 ±
0.2 μm, Q = 1.0–2.0, Qmr = 1.3–1.6, Qmm = 1.5
± 0.1; n = 10–25 spores per 12 collections],
ellipsoid, smooth, hyaline, inamyloid, thin-
walled. Basidia clavate, 4-spored. Basidioles
18–35 (–40) × 6–8 μm, clavate to cylindrical
with rounded to subacute apex. Pleurocystidia
31–60 × 6–9 μm, clavate, hyaline, inamyloid,
thick-walled. Cheilocystidia common; main
body 19–32 × 4–10 μm, clavate, simple or a
majority with 1–4 apical appendages, hyaline,
inamyloid, thin-walled. Pileipellis a cutis, 5.5–
6 μm diam., hyaline, inamyloid, with spiral
encrustations, giving rise to terminal hairs with
basal clamp connections. Hairs 45–460 × 2.5–5
μm diam., cylindrical, apex rounded to acute,
with secondary septations, thick-walled,
dextrinoid, yellowish-brown to greenish brown
in KOH. Pileus and lamellar trama subregular;

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135
Fig. 18. Crinipellis trichialis (JFK 109). – 1. Basidiomes (x2). – 2. Basidiospores (JFK 108). – 3. Basidia and basidioles.
– 4. Cheilocystidia. – 5. Caulocystidia. – 6. Pleurocystidia. – 7a-b. Pileipellis hairs. – Bar: 2-4, 7a = 10 µm; 5, 7b = 25
µm.
hyphae 3–6 μm diam., hyaline, inamyloid,
slightly thick-walled. Stipe tissue monomitic,
parallel, tightly packed; stipitipellis composed
of repent cortical hyphae and terminal hairs;
cortical hyphae 1–3 μm diam., cylindrical,
dextrinoid, light yellow brown to green in
KOH; medullary hyphae 3–5 μm diam.,
cylindrical, strongly dextrinoid, hyaline;
terminal hairs 100–325 × 7.5–10 μm,
cylindrical, apices acute, with secondary
septations, dextrinoid, yellowish brown to
greenish brown in KOH. Clamp connections
present.
Habitat: Solitary to gregarious,
primarily on bamboo or rarely on decomposing
dicot sticks.
Known distribution: Brazil, Indonesia,
Malaysia, Venezuela.
Material examined: INDONESIA. Java, Bogor,
Bogor Botanical Gardens, S06˚35.955', E106˚47.720',
elev. 266 m, 7 January 1998, D.E. Desjardin 6716 (BO,
SFSU); same location, 15 January 1998, D.E. Desjardin
6800 (SFSU); same location, 12 January 1999, D.E.
Desjardin 6924 (SFSU); same location, 6 January 2000,
D.E. Desjardin 7043 (SFSU), same location, 12 January
2000, A.W. Wilson 33 (SFSU); same location, 24
February 2000, A. Retnowati 216 (BO, SFSU); same
location, 15 January 2005, Kerekes 111 (BO, SFSU);
Java, Cibodas, 29 December, 1998, E. Horak (BO; SFSU)
(same as AR 045); Java, West of Bogor, Hutan
Penelitian, Haurbentes, S06˚32.646', E106˚26.261', elev.
292 m, 14 January 2005, Kerekes 108 (BO, SFSU); same
location, 14 January 2005, Kerekes 109 (BO, SFSU);
Java, Buitenzorg, coll. by F. von Hoehnel, on bamboo,
det. by Rolf Singer (FH). Java, Zollinger #2078, on
stems of bamboo (Holotype: FH). MALAYSIA.
Selangor, Selayang, Kanching Forest Reserve,
N03˚17.954' E101˚37.153', elev. 110 m, 9 January 2005,
Kerekes 97 (SFSU).
Notes: Our recently collected specimens,
along with the holotype, were from Java,
Indonesia with only one collection from
Malaysia, where this species appears to be

136
Fig. 19. Crinipellis Basidiomes. – 1, 2. Crinipellis actinophora (JFK 100) rhizomorphs in the field and basidiomes. – 3.
Crinipellis brunneipurpurea (JFK 84). – 4. Crinipellis brunnescens (DED 6791). – 5. Crinipellis furcata (DED 6951). –
6. Crinipellis trichialis (JFK 108). – 7. Crinipellis malesiana (KUM 084a). – 8. Crinipellis dipterocarpi f. cinnamomea
(JFK 50). – 9. Crinipellis tabtim (JFK 44). – 10. Crinipellis cupreostipes (JFK 31). –11. Crinipellis cupreostipes (JFK
131). – 12. Crinipellis aff. iopus. – 13. Crinipellis dipterocarpi (JFK 51).
1
2
4
3
5 6
7
8
9
10
11
12
13

Fungal Diversity
137
restricted to dead Bambuseae. Distinguishing
characteristics are the concentric zones of color
on the pileus with long, erect, hispid hairs
around the disc, the presence of pleurocystidia,
and the green KOH reaction of the pileus hairs
and stipe tissues. Singer (1976, 1986) placed C.
trichialis in section Grisentinae because of the
green KOH reaction. Our molecular data,
however, indicate that C. trichialis is sister of
C. scabella (type species of the genus), a
species that lacks a green KOH reaction, one of
the defining features of sect. Crinipellis.
The holotype specimen from Java (Zollinger
#2078, FH) is incorrectly labeled as “Agaricus
trichophorus Lév.”, which indicates a different
species collected by Zippelius from Java on
wood. The latter (Ag. trichophorus Zipp. ex
Lév., Ann. Sci. Nat., Bot. sér. 3, 2: 171. 1844)
is probably a species of Mycena (Pers.) Roussel
or Hemimycena Singer with a white pileus
bearing setae and a glabrous stipe.
20. Moniliophthora canescens (Har. Takah.)
Kerekes & Desjardin, comb. nov.
(Fig. 20: 1–5, 24)
MycoBank: 513037
Basionym: Crinipellis canescens Har.
Takah., Mycoscience 41: 171. 2000.
Type: JAPAN, Iriomote Island, Okinawa
Pref., along the Urauchi River, 2 June 1999,
solitary to scattered on a dead fallen twig of
broad-leaved tree, KPM-NC-0005014
(Holotype: CBM). Type not examined.
Description based on fresh material:
Pileus 9–15 mm diam., broadly convex
to plano-convex, umbilicate, sulcate,
appressed-radially-fibrillose, dull, dry; dull
greyish orange (5B5–6) overall. Context thin,
pliant. Lamellae adnate, distant with 1–2 series
of lamellulae, broad (1–2 mm), bright organte
(5A6–8). Stipe 5–10 × 0.75–1 mm, central,
terete, cylindrical, ± equal, curved, hispidulous
overall, insititious, often arising from a thin
white subiculum: brownish grey (6C3) overall.
Rhizomorphs thin, wiry, black, glabrous
rhizomorphs; rhizomorphs adhere masses of
twigs and debris together, arboreal. Odor and
taste none Basidia 22–28 × 5–7 μm, clavate, 4-
spored. Basidioles clavate. Pleurocystidia
absent. Gill edge heteromorphous with few
cheilocystidia. Cheilocystidia with main body
20–27 × 4–6 μm, clavate to irregular, simple or
with up to 3 apical appendages 2–5 μm long,
hyaline, inamyloid, thin-walled. Pileipellis a
cutis of inflated cells 27–50 × 6–10 μm with
dextrinoid incrustations, hyaline to light brown
in KOH, giving rise to terminal hairs with basal
clamp connections. Hairs few and realatively
short, 87.5–115 × 6–8 μm, thick-walled (2–3
μm), lacking secondary septations, hyaline to
yellow with orange tints in water, hyaline in
KOH. Clamp connections present.
Basidiospores 7–10 x 4–5 μm [xm = 8.5 ±
0.7 × 4.1 ± 0.3 μm, Q = 1.6–2.3, Qm = 2.1 ± 0.3;
n = 25 spores per 1 collection], ellipsoid,
smooth, hyaline, inamyloid, thin-walled.
Habitat: Basidiomes rare, rhizomorphs
common, on undetermined dicot plants in
primary forest. Known distribution: Iriomote
Island (southwestern Japan), Malaysia.
Material examined: MALAYSIA. Kuala Lumpur,
Gombak Field Station, 12 Jan. 2003, D.E. Desjardin
7518 (SFSU).
Notes: We requested the type collection
on loan from Japan, however it was not
available for examination. Our material from
Malaysia matches nicely with the protologue of
C. canescens. The ITS sequence data (Fig.1)
for DED 7518 indicate that the species belongs
in the Moniliophthora clade with 53% ML-BS
and .50 PP support. Based on the micro- and
macromorphological characteristics and ITS
sequence data, we recognize Crinipellis
canescens as belonging to the genus
Moniliophthora and make the formal transfer
herein.
21. Moniliophthora marginata Kerekes,
Desjardin & Vikineswary, sp. nov.
(Fig. 21: 1–6, 24)
MycoBank: 512277
Etymology: marginatus (Latin) – margined,
referring to the brownish-red margins of the lamellae.
Pileus 22 mm latus, flabelliformis, rubrobrunneus.
Lamellae liberae, marginatus rubrobrunneus. Stipes 6–8
× 1 mm, cylindricus, fibrillosus, rubrobrunneus. Odor
saporque nulli. Basidiosporae 5–10 × 3–5 μm,
oblongatae, hyalinae. Basidia 35–36 × 6 μm, clavata,
hyalinae, tetraspora. Pleurocystidia nulla. Cheilocystidia
19–40 × 4–7, clavata. Trama lamellarum regularis.
Caulocystidia 60–150 × 5–6.25 μm, cylindraceis.
Holotypus: MALAYSIA. Pahang Province, Fraser’s Hill,
Bishop’s trail, N03°42.949', E101°44.430', elev. 1212 m,
15 January 2005, Kerekes 15 (Holotype: KLU-M).
Pileus 22 mm diam., circular to
rounded flabelliform, centrally depressed,

138
Fig. 20. Moniliophthora canescens (DED 7518). – 1. Basidiomes. – 2. Basidiospores. – 3. Basidium and basidioles. – 4.
Cheilocystidia. – 5a-b. Pileipellis hairs. – 5c. Pileipellis hair subtending cell. – Bar: 2-4, 5b, 5c = 10 µm; 5a = 25 µm.
sulcate, with scaly or wart-like tufts of fibrils
concentrated near the disc, interwoven
appressed-fibrillose elsewhere; reddish brown
(9D–F6), darkest near disc (9F6) and lighter
towards margin, interwoven hairs range from
reddish brown (9F6) to light reddish brown
(9D6). Context < 1 mm thick, brownish violet
(11D6). Lamellae free, close with 3 series of
lamellulae, narrow (1–2 mm), off white with
brownish red (10D6) edges. Stipe 6–8 × 1 mm,
central, cylindrical, fibrillose to squamulose,
insititious; apex white, becoming brownish red
(10D6) below. Odor and taste none.
Basidiospores 5–10 × 3–5 μm [xm = 7.0 ±
1.0 × 4.0 ± 0.6 μm, Q = 1.3–2.3, Qm = 1.8 ± 0.3;
n = 25 spores per 1 collection], oblong with
some tapering towards the apex to
amygdaliform in side view, smooth, hyaline,
inamyloid, thin-walled. Basidia 35–36 × 6 μm,
clavate, 4-spored, thin-walled to slightly thick-
walled. Basidioles clavate. Pleurocystidia
absent. Cheilocystidia with main body 19–40 ×
4–7 μm, narrowly cylindrical to clavate,
lacking obvious apical appendages although
some with small knobs near the apex, hyaline
to brown, weakly dextrinoid, partially thick-
walled, most basally clamped. Pileipellis a
cutis to a trichoderm with inflated chains of
hyaline cells 28–60 × 6.5–11 µm, coarsely
incrusted, giving rise to terminal hairs with
basal clamp connections. Hairs 50–337 × 5–10
μm, cylindrical, apices generally rounded,
some with secondary septations, thick-walled,
strongly dextrinoid, purple in KOH. Pileus
trama regular; hyphae 3–7 μm diam., hyaline,
weakly dextrinoid near hairs, thin-walled.
Lamellar trama regular; hyphae 2.5–9 μm
diam., hyaline, inamyloid, thin-walled. Stipe
tissue stipitipellis composed of repent cortical
hyphae and terminal hairs; cortical hyphae 1.5–
4 μm diam., cylindrical, pale yellowish brown
to hyaline, dextrinoid, thin-walled; medullary
hyphae 1–3 μm diam., cylindrical, hyaline,
dextrinoid, thin-walled; terminal hairs 60–150
× 5–6 μm, cylindrical, apices rounded, some
with secondary septations, thick-walled,
dextrinoid, purple to hyaline in KOH. Clamp
connections present.
Habitat: In a primary montane cloud
forest on undetermined decaying woody stem.
Known distribution: Malaysia.
Material examined: MALAYSIA. Pahang
Province, Fraser’s Hill, Bishop’s trail, N03°42.949 ',
E101°44.430' ,elev. 1212 m, 15 January 2005, Kerekes
15 (Holotype: KLU-M; Isotype: SFSU).

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139
Fig. 21. Moniliophthora marginata (JFK 15). – 1. Basidiomes (a. x2). – 2. Basidiospores. – 3. Basidia. – 4.
Cheilocystidia. – 5. Caulocystidia. – 6a. Pileipellis hair segment. – 6b. Pileipellis hair. – Bar: 2-4, 6a = 10 µm; 5, 6b =
25 µm.
Notes: DNA was extracted from
specimen JFK 15, however, sequencing was
not successful; therefore placement of M.
marginata in the phylogenetic tree is unknown.
Features of M. marginata suggest a close
affinity with members of Crinipellis sect.
Iopodinae of which many members are
currently accepted in Moniliophthora. This
species is close to Crinipellis eggersii Pat., but
the latter differs by forming larger
basidiospores (11–13 × 5.5–6.3 µm) and forms
purple to violet purple basidiomes (Singer
1976). We suspect that C. eggersii belongs in
the genus Moniliophthora as originally
suggested by Aime and Phillips-Mora (2005),
but a formal transfer will not be made until
material from Ecuador (type locality) is studied.
Moniliophthora marginata is also similar to C.
trinitatis Dennis (sect. Iopodinae), described
from Trinidad, but the latter species differs in
forming forked cheilocystidia, and grows on
living Vitex trees (Singer, 1976).
22. Moniliophthora nigrilineata (Corner)
Desjardin & Kerekes, comb. nov.
(Fig. 22: 1–2)
MycoBank: 513038
Basionym: Crinipellis nigrilineata
Corner, Beih. Nova Hedwigia 111 120. 1996.
Type: SINGAPORE, Mandai Road, 15
Oct. 1934, coll. by E.J.H. Corner, record
#192520 (Holotype: E!).
Analysis of the holotype specimen
The holotype specimen consists of > 10
badly fragmented basidiomes, pressed flat, in
poor condition. As dried: Pileus 4–8 mm diam.
(10–35 mm diam in protologue when fresh),
ranging from clay-color to brown, hirsute.
Lamellae adnate to subdecurrent, close to

140
Fig. 22. Moniliophthora nigrilineata (E.J.H. Corner,
record #192520). – 1. Basidiospores. – 2a. Pileipellis
hairs. – 2b. Pileipellis cell subtending a terminal hair. –
Bar: 1-2 = 10 µm.
distant, narrow to moderately broad, pallid to
brown. Stipe 8–16 × 0.5 mm, pruinose, dark
brown at base, arising directly from the
substrate; rhizomorphs absent.
Basidiospores 9–12.8 × 3.5–4.5 μm [xm =
11 ± 1.4 × 4.0 ± 0.4 μm, Q = 2.3–3.0, Qm = 2.7
± 0.4; n = 10 spores per 1 collection], elongate-
ellipsoid to subfusoid, smooth, hyaline,
inamyloid, thin-walled. Basidia clavate, 4-
spored. Basidioles subfusoid. Pleurocystidia
absent. Cheilocystidia absent; lamellar edge
fertile. Pileipellis 8–19 μm diam., inflated,
short-celled, smooth or brown-incrusted,
hyaline or often with brown plasmatic
pigments, inamyloid, unchanged in KOH, thin-
walled giving rise to hairs. Hairs 60–110 ×
5.5–6.5 μm, few, scattered, irregularly
cylindrical, obtuse, evenly orange-brown or
with paler base; thick-walled (1–3 μm), smooth,
hyaline, inamyloid to weakly dextrinoid,
unchanged in KOH.
Notes: The pileipellis anatomy is unlike
that of Crinipellis, in that it lacks numerous
long, narrow, dextrinoid skeletal hairs. In
addition, the lamellae were reported as
subdecurrent, furcated and connected by
shallow veins. These features in combination
with the elongated, subfusoid basidiospores
and an absence of cheilocystidia indicate that
the species is better placed in the genus
Moniliophthora, and a formal transfer is made
herein. This species is not included in the
phylogenetic analysis because extraction of
DNA from the type specimen was not
permitted.
23. Marasmiellus subochraceus (Corner),
Kerekes & Desjardin comb. nov.
(Fig. 23: 1–5, 24)
MycoBank: 513039
Basionym: Crinipellis subochracea
Corner, Beih. Nova Hedwigia 111: 121. 1996.
Type: SINGAPORE, Reservoir Jungle, 1
Aug. 1940, coll. by E.J.H. Corner, record
#192521 (Holotype: E!).
Analysis of the holotype specimen
The holotype specimen consists of one
half of one pileus attached to a stipe, plus one
stipe lacking a pileus, both attached to a woody
stick; in poor condition. As dried: Pileus 3 mm
diam, pallid, hirsute with white to golden hairs.
Lamellae not observable. Stipe 5 × < 0.5 mm,
fibrillose, brown, insititious, arising directly
from the substrate. Rhizomorphs absent.
Basidiospores 9.6–12 × 3.5–4.2 µm
(n=5), subfusoid to clavate, smooth, hyaline,
inamyloid, thin-walled. Hymenial elements not
recovered; material revived poorly. No basidia,
basidioles or cystidia observed. Pileipellis a
cutis of inflated cells giving rise to terminal
hairs. Hairs ranging from cylindrical to
ventricose, 60–180 µm long, base 4–10 µm
diam, apex 2–3.2 µm diam, gradually narrowed
to a subacute tip; walls hyaline, inamyloid,
smooth, 0.5–3 µm thick. Subtending hyphae
inflated, 9–16 (–25) µm, hyaline, inamyloid,
thin-walled; some terminal cells like the
subtending hyphae, fusoid to clavate, hyaline,
thin-walled.
Macromorphological description based on
fresh material:
Pileus 4–10 mm diam, convex, disc with
dark brown, powdery-fibrous patches or spots,
margin with scattered dark brown, short hairs,
ground color white and appressed-silky under
ornamentation. Context < 1 mm thick.
Lamellae narrowly adnate, subdistant to distant
with 1–2 series of lamellulae, moderately broad,
white. Stipe 5–10 × 1 mm, central, terete,
cylindrical, ± equal, curved, tough, dull, dry,
silky, subinsititious, white with an incarnate
(pink) base. Odor and taste none.

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141
Fig. 23. Marasmiellus subochraceus (DED 7523). – 1. Basidiomes (x1). – 2. Basidiospores. – 3. Basidioles. – 4.
Pileipellis terminal cells. – 5. Pileipellis hairs and Pileipellis subtending cell. – Bar: 2-4 = 10 µm; 5 = 25 µm.
Micromorphological description based on
data from the holotype specimen and fresh
material
Basidiospores 10.0–15 × 4–6 μm [xmr =
10.0–12.5 × 3.8–4.5 μm, xmm = 11.2 ± 1.8 × 4.2
± 0.5 μm, Q = 1.9–3.8, Qmr = 2.6–2.8, Qmm =
2.7 ± 0.1; n = 5–25 spores per 2 collections],
subfusoid to clavate, smooth, hyaline,
inamyloid, thin-walled. Basidia and basidioles
18–24 × 6–8 μm; basidia clavate, 4-spored;
basidioles clavate. Pleurocystidia absent.
Cheilocystidia absent, gill edge fertile.
Pileipellis a cutis of inflated cells, 45–85 × 12–
13.5 μm clavate to allantoid, with encrustations,
dextrinoid, light tan to yellowish tan in KOH,
giving rise to terminal hairs with basal clamp
connections. Hairs sparse, 60–205 × 4–10 μm,
cylindrical to ventricose, generally rounded at
apex or gradually narrowing to a subacute apex,
lacking secondary septations, thick-walled
(0.5–3 μm), inamyloid, light brown in KOH.
Subtending hyphae inflated, 9–16 (–25) μm
diam., hyaline, inamyloid, thin-walled; some
terminal cells like the subtending hyphae,
fusoid to clavate, hyaline, thin-walled. Pileus
trama hyphae 3–9 μm diam., cylindrical,
hyaline, inamyloid. Lamellar trama hyphae 2–
6 μm diam., cylindrical, hyaline, inamyloid.
Stipe tissue monomitic, parallel. No obvious
difference between the cortical and medullary
hyphae, 2–5 μm diam., cylindrical, hyaline,
inamyloid, thin-walled; no terminal hairs
observed. Clamp connections present.
Notes: The pileipellis anatomy of scattered
inamyloid hairs in combinations with broadly
clavate, thin-walled, inamyloid cells is not
characteristic of Crinipellis. These features, in
combination with a white pileus, forked
lamellae, the absence of hymenial cystidia, and
subfusoid basidiospores indicate that the
species is better placed in the genus
Marasmiellus, sect. Stenophylloides Singer. In
preliminary parsimony analyses, M.
subochraceus was sister to a clade containing
Crinipellis and Moniliophthora species with
low bootstrap support, no PP support, and on a
very long branch, whereas in preliminary
maximum likelihood analyses M. subochraceus
nested within the Crinipellis clade without
statistical support and on a very long branch.
Because of its divergent sequence and
consistent placement on a long branch in all
analyses, M. subochraceus was not included in
the final ML analyses. Based on overall
morphological similarity we recognize C.
subochracea as belonging to the genus
Marasmiellus and a formal transfer is made
herein.
Habitat: Scattered on dead twigs of
undetermined dicot tree in primar forest.
Known Distribution: Malaysia, Singapore.
Material examined: MALAYSIA. Kuala Lumpur,
Gombak Field Station, 12 January 2003, D.E. Desjardin
7523 (SFSU). SINGAPORE. Reservoir Jungle, 1 August
1940, coll. by E.J.H. Corner, record #192521 (Holotype:
E).

142
Fig. 24. Moniliophthora basidiomes. – 1. Moniliophthora marginata (JFK 15). – 2. Moniliophthora canescens (DED
7518). – 3. Marasmiellus subochraceus (DED 7523).
Excluded and Insufficiently Known Taxa,
Including Relevant Type Studies of
Extralimital Species
24. Agaricus omotrichus Berk., London J. Bot.
2: 510. 1843. (Fig. 25: 1–3)
≡ Collybia omotricha (Berk.) Sacc., [ut
homotricha] Syllog. Fung. 5: 217. 1887.
≡ Crinipellis omotricha (Berk.) D.A. Reid [ut
homotricha], Contr. Bolus Herb. 7: 111. 1975.
Type: SOUTH AFRICA, Uitenhage,
December, H. Zeyher #111, on decayed grasses
(Holotype: K!).
Analysis of the holotype specimen
The holotype specimen consists of 4
basidiomes in fair condition, attached to grass
stems; one basidiome cut in half lengthwise
and only half remains. As Dried: Pileus 6–8
mm diam., convex, short-shaggy, dirty brown.
Lamellae distant, broad, dark brown. Stipe 10–
15 × 1–1.5 mm, shaggy, brown, arising directly
from substrate; rhizomorphs absent.
Basidiospores 9–10.2 × 4.2–5.5 μm, [xm
= 9.6 ± 0.5 × 4.9 ± 0.4 μm, Q = 1.7–2.4, Qm =
2.0 ± 0.2; n = 10 spores per 1 collection],
ellipsoid, smooth, hyaline, inamyloid, thin-
walled. Basidia clavate, 4-spored. Basidioles
clavate to subfusoid. Pleurocystidia absent.
Cheilocystidia scattered, lamellar edge fertile;
30–37 × 6.5–8.5 μm, clavate to lageniform,
obtuse, simple (non-diverticulate), hyaline,
thin-walled. Pileipellis of tangled to subparallel,
thick-walled hairs. Hairs 3.2–5.5 μm diam.,
rather short (< 200 μm), cylindrical to sinuous
below, gradually tapered to a narrow, subacute
tip, apically thin-walled, base 0.5–2.2 μm thick,
hyaline to yellow, inamyloid to weakly
dextrinoid, smooth.
Habitat: On decayed grasses.
Known distribution: South Africa;
reported from Sri Lanka.
Notes: This is considered a good species
of Crinipellis, distinct because of the
moderately large spores, short pileipellis hairs,
simple cheilocystidia and grass habit.
Additional material examined: SRI LANKA.
Peradeniya, May 1844, Gardner #10, on the ground (K);
Colombo, Victoria Park, 7 May 1981, coll. by C.
Dharma-Wardana, (DAOM 189041). Cited by Berkeley
(1847) and Pegler (1986), respectively, as contaxic with
Agaricus omotrichus.
The material from Peradeniya (Gardner
#10), did not provide much taxonomically
important data, however the pileipellis hairs of
this specimen are distinctly different from
those of the holotype of A. omotrichus (K!),
1
2
3

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143
Fig. 25. Agaricus omotrichus (Holotype). – 1.
Basidiospores. – 2. Cheilocystidia. – 3. Pileipellis hairs.
– Bar: 1-3 = 10 µm.
such that this specimen should not be
considered as belonging to that species. The
hairs of Gardner #10 are more similar to the
grass inhabiting Crinipellis pseudostipitaria
subsp. orientalis Singer, described from
Vietnam. The specimen collected in Colombo
(DAOM 189041) is also not contaxic with the
holotype specimen of A. omotrichus. The latter
has clavate, simple cheilocystidia, thinner-
walled pileipellis hairs, and larger
basidiospores. The Colombo material at
DAOM probably represents an eastern form of
Crinipellis pseudostipitaria, as it was originally
annotated by Redhead. However, it does not
represent C. pseudostipitaria subsp. orientalis,
which has larger basidiospores (Holotype: FH!).
Status: Based on the data on hand, C.
omotricha does not occur in Southeast Asia.
25. Collybia multicolor Petch, Ann. Roy. Bot.
Gard., Peradeniya 6: 197. 1917. (Fig. 26: 1)
≡ Crinipellis multicolor (Petch) Pegler, Kew Bull.
Add. Ser. 12: 139. 1986.
Type: SRI LANKA, Hakgala, May 1910,
T. Petch # 3053, ex alcohol (Holotype: K!).
Analysis of the holotype specimen:
The holotype specimen consists of 2
fragmented basidiomes, pressed flat, in poor
condition. As dried: Pileus 9–11 mm diam,
plano-convex with a depressed disc, hirsute
with erect, minutely fascicled hairs, dark brown.
Lamellae subdistant, very broad, some with
white-crystalline edges, dark brown. Stipe 15 ×
2 mm, densely pubescent, olive-brown, arising
directly from the substrate; rhizomorphs absent.
All hymenial elements golden brown in
3%KOH; not reviving well. Hyphal pegs
absent. Basidiospores, basidia, basidioles and
cystidia not recovered. Pileipellis of erect,
agglutinated-fasciculate, < 100 × 4–8 μm,
cylindrical below, gradually tapered to a
narrower, obtuse apex, thin- to thick-walled,
hyaline to brown, inamyloid. Pileus and
lamellar trama of gelatinized hyphae 4–16 μm
diam., with walls 2–6.5 μm thick, irregularly
cylindrical, hyaline, inamyloid.
Habitat: On dead vegetable matter and
dead wood.
Known distribution: Sri Lanka.
Notes: This is not a Crinipellis species as
established by Pegler (1986). The non-
dextrinoid and very short pileipellis hairs, in
combination with the gelatinized skeletal
tramal hyphae up to 6.5 μm thick indicate that
the species is closer to Lentinus Fr. or Panus Fr
Status: Without better material to analyze, C.
multicolor will have to remain as an
insufficiently known species.
26. Crinipellis carecomoeis var. subelata
Singer, Lilloa 8: 488. 1943. (Fig. 26: 2)
Type: NEW CALEDONIA, Col d’Amieu,
1907, coll. by Le Rat, det. by Patouillard as
Crinipellis stipitaria var. castaneus, on fallen
leaves (Holotype: FH!).
Analysis of the holotype specimen:
The holotype specimen consists of one
intact basidiome plus two stipes attached to
leaf fragments, in poor condition. As dried:
Pileus 2 mm diam, convex, hairy, pale brown
overall. Lamellae all removed. Stipe 15 × 0.5
mm, pubescent, pallid greyish brown,
associated with fine, brown, wiry, glabrous
rhizomorphs.
No lamellae are present on the extant
basidiome. No basidiospores nor other.
hymenial cells observed; all degraded.
Pileipellis of thick-walled hairs. Hairs 2.2–
5.5(–6) μm diam., nearly all evenly sinuous,
diam. equal, thick-walled (1–1.5 μm), apices
cylindrical and obtuse, lacking secondary
septations near apices, these scattered near base
of hairs; hyaline to pale yellow, inamyloid to
very weakly dextrinoid, non-incrusted.
Stipitipellis setoid; acute-lanceolate, up to 11.5
µm diam at broadest with walls up to 3.5 μm
thick, hyaline.

144
Fig. 26. 1. Collybia multicolor – a. Pileipellis hair. – 2. Crinipellis carecomoeis var. subelata – a. Pileipellis hairs b.
Caulocystidia. – 3. Crinipellis elata – a. Pileipellis hairs b. Caulocystidia. – 4. Crinipellis filiformis – a. Basidiospores b.
Pileipellis hair. – Bar: 1-4 = 10 µm.
Habitat: On fallen leaves.
Known distribution: New Caledonia.
Notes: This epithet represents an
insufficiently known species of Crinipellis. The
protologue was based on dried material that
was presumably nearly all used up during
description. No intact lamellae are present so
data on hymenial elements are unavailable.
The species forms distinctive entirely sinuous
pileus hairs and setoid caulocystidia.
Status: Insufficiently known.
27. Crinipellis elata Pat., Bull. Soc. Mycol.
France 24: 167. 1908. (Fig. 26: 3)
≡ Marasmius elatus (Pat.) Sacc. & Trotter, Syllog.
Fung. 21: 112. 1912.
= Crinipellis carecomoeis f. (monstr.) elata (Pat.)
Singer, Lilloa 8: 489. 1942.
Type: NEW CALEDONIA, La Foa et la
Col d’ Amieu, 1907, coll. by Le Rat #96, on
leaves (Holotype: FH!).
4
a
b

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145
Analysis of the holotype specimen
The holotype specimen consists of two
intact immature basidiomes in fair condition,
plus two stipes, attached to a single dicot leaf.
As dried: Pileus immature, < 1 mm diam,
campanulate, silky-hairy, brown. Lamellae not
observable. Stipe 45–170 × ≤ 0.5 mm, greyish
brown, appressed-pubescent; rhizomorphs
absent.
No hymenial elements were observable;
pilei immature. Pileipellis of radially arranged,
subparallel hairs. Hairs 2.5–4 μm diam.,
average about 3.2 μm diam., tips cylindrical,
obtuse, thin-walled (≤ 0.5 μm), below the tip
evenly sinuous for the rest of the length, thick-
walled to solid (1–1.5 μm), hyaline to pale
yellow, dextrinoid, smooth, non-incrusted.
Stipitipellis of repent hairs; hairs cylindrical to
sinuous, 2.2–4 μm diam. with walls up to 1.5
μm thick, similar to pileipellis hairs; lacking
erect setoid lanceolate hairs.
Habitat: On leaves.
Known distribution: New Caledonia.
Notes: Singer (1942) considered C. elata
as an abnormal sterile form of C. carecomoeis
var. subelata, with a long stipe. He reported C.
elata as growing on wood, but the holotype is
clearly on a dicot leaf. Crinipellis elata is not
contaxic with C. carecomoeis var. subelata as
accepted by Singer. The latter taxon has erect,
setoid, lanceolate hairs on the stipe that are
lacking in C. elata.
Status: Crinipellis elata will remain
insufficiently known until fertile specimens of
this long-stiped taxon are found.
28. Crinipellis filiformis G. Stev., Kew Bull.
19: 42. 1964. (Fig. 26: 4)
Type: NEW ZEALAND, Buller Bush, 27
Oct. 1947, coll. by G. Stevenson Cone #213,
under tawa (Holotype: K!).
Analysis of the holotype specimen:
The holotype specimen consists of one
packet containing a leaf fragment with a few
reddish brown, glabrous, wiry rhizomorphs;
and a second packet containing one broken
stipe plus one quarter of one pileus. As dried:
Pileus 2 mm diam, disc with a small brown
papilla, shaggy-hairy, cream-colored on the
margin. Lamellae close, narrow, cream-colored.
Stipe 8 × 0.5 mm, greyish brown, with cream-
colored pubescence.
Basidiospores 8.3–10.5 × 4.2–5.2 μm [xm
= 9.1 ± 0.7 × 4.7 ± 0.3 μm, Q = 1.7–2.2, Qm =
1.9 ± 0; n = 12 spores per 1 collection],
ellipsoid, smooth, hyaline, inamyloid, thin-
walled. Hymenial elements not observable: no
basidia, basidioles or cystidia seen. Pileipellis
of tangled to subparallel and agglutinated,
thick-walled hairs. Hairs > 300 × 2.2–3.5(–5.5)
μm, cylindrical to sinuous, gradually narrowed
to an acute or subacute tip, hyaline, inamyloid
or dextrinoid; walls smooth, 0.5–1.3 μm thick;
no secondary septa seen near the tips.
Habitat: Fallen leaves.
Known distribution: New Zealand.
Notes: Data on hymenial cells were not
obtainable because of the poor quality of the
limited material in the holotype specimen.
Features of the cheilocystidia are required for
accurate taxonomic placement.
Status: Crinipellis filiformis will remain
an insufficiently known species of Crinipellis
until more material becomes available for
analysis.
29. Crinipellis novae-zelandiae G. Stev., Kew
Bull. 19: 45. 1964.
≡ Collybia novae-zelandiae (G. Stev.) E. Horak,
N. Zealand J. Bot. 9: 440. 1971.
≡ Lentinula novae-zelandiae (G. Stev.) Pegler,
Sydowia 36: 234. 1983.
Type: NEW ZEALAND, Otari, Wilton's
Bush, 18 May 1947, G. Stevenson Cone #150,
on fallen logs (Holotype: K).
The holotype specimen consists of
portions of the pilei of two basidiomes plus one
broken stipe, in good condition. Material not
examined.
Notes: Pegler (1983) provides a good
description of this species and makes the
formal transfer to Lentinula Earle. Hibbett et al.
(1995) included New Zealand Lentinula
isolates in their molecular phylogenetic
analysis of shiitake mushrooms, and found that
the NZ material, referable to L. novae-
zelandiae, represents a monophyletic lineage,
distinct from other Lentinula species.
Status: Currently accepted as Lentinula
novae-zelandiae.
30. Crinipellis patouillardii Singer, Lilloa 8:
483. 1943. (Fig. 27: 1–3)
Type: JAPAN, Bonin Islands, 1.11.4, coll.
by C. Wright, “on leaves in shaded ravine.”

146
Fig. 27. Crinipellis patouillardii (Holotype). – 1. Basidiospores. – 2. Cheilocystidia. – 3. Pileipellis hairs. – Bar: 1-3 =
10 µm.
U.S. North Pacific Exploring Expedition #25 as
Agaricus stipitarius (Holotype: FH!).
Analysis of the holotype specimen
The holotype specimen consists of a
mixed collection. Three basidiomes (one loose
in the packet and two attached to woody sticks)
represent the material described by Singer as C.
patouillardii. Two additional basidiomes
represent an agaric species in the Corti-
nariaceae. As dried, the holotype basidiomes as
designated by Singer: Pileus 5 mm diam.,
convex with two concentric depressions
surrounding a small papilla, radially shaggy-
hairy, brown. Lamellae adnexed, subdistant,
broad, pale brown. Stipe 7–8 × 0.5–0.75 mm,
shaggy-fibrillose, cream-beige, arising from
woody (not leafy) substrate; rhizomorphs
absent.
Basidiospores 7.7–10.2 × 4.5–6 (–6.4)
μm, [xm = 8.9 ± 0.8 × 5.4 ± 0.4 μm, Q = 1.4–
1.9, Qm = 1.7 ± 0.1; n = 20 spores per 1
collection], broadly ellipsoid to subfusoid,
smooth, hyaline, inamyloid, thick-walled.
Basidia, basidioles and pleurocystidia
collapsed; data unobtainable. Cheilocystidia
agglutinated, many collapsed, common, 22–32
× 7–10 μm, clavate to broadly clavate, lacking
diverticula or apical appendages, thick-walled,
hyaline. Pileipellis of tangled to agglutinated
and radially arranged hairs. Hairs > 200 × 2.2–
4 μm (relatively narrow), single or in tufts of
5–10, cylindrical, not appreciably narrowed
near the apex, obtuse, relatively thick-walled
over the upper 30% (< 0.5 μm), thick-walled at
the base (0.5–1.5 μm), hyaline to pale brown,
dextrinoid, non-incrusted, secondary septa rare.
Stipitipellis similar to the pileipellis; hairs
similar but with walls more evenly 1 μm thick.
Habitat: On woody debris and leaves.
Known distribution: Japan.
Notes: This represents a good species of
Crinipellis, distinctive because of the simple,
broadly clavate cheilocystidia, relatively broad
basidiospores, cylindrical and relatively narrow
and thin-walled pileipellis hairs. Although
Wright’s notes state “on leaves” the material in
the holotype packet is on sticks. In addition,
two non-Crinipellis cortinariaceous basidiomes
are in the holotype packet.
Status: Crinipellis patouillardii is known
currently only from the Bonin Islands, Japan.
31. Crinipellis procera G. Stev., Kew Bull. 19:
43. 1964. (Fig. 28: 1–3)
Type: NEW ZEALAND, Levin, Ohau
River, 22 July 1949, coll. by G. Stevenson
Cone #743, inserted on dead twigs of forest
litter (Holotype: K!).
Analysis of the holotype specimen:
The holotype specimen consists of 6
intact basidiomes in fair condition. As dried:
Pileus 4–8 mm diam., conical with a conical
papilla surrounded by a zone of long tufted
hairs, margin with shorter and smaller tufts,
yellowish brown to orangish brown. Lamellae
free, close with 1 series of lamellulae, narrow,
cream-colored. Stipe 60–100 × 1 mm,
pubescent, greyish brown; rhizomorphs absent.

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147
Fig. 28. Crinipellis procera (Holotype). – 1. Basidiospores. – 2. Cheilocystidia. – 3. Pileipellis hair. – Bar: 1-3 = 10 µm.
Basidiospores (7.7–) 8–9.6 × 4.2–5.5 μm
[xm = 8.8 ± 0.5 × 4.7 ± 0.4 μm, Q = 1.7–2.2,
Qm = 1.9 ± 0.1, n = 20 spores per 1 collection],
ellipsoid, smooth, hyaline, inamyloid, thin-
walled. Basidia 2- and 4-sterigmate, clavate,
clamped. Basidioles subfusoid. Pleurocystidia
absent. Cheilocystidia common, lamellar edge
sterile; main body 11–20 × 4.5–8 μm,
irregularly cylindrical to clavate with numerous
apical appendages; apical appendages 1.5–6.5
(–10) × 1–3 μm, finger-like or irregular in
outline, obtuse, sometimes forked, hyaline,
inamyloid, thin-walled. Pileipellis of tangled to
parallel and agglutinated, thick-walled hairs.
Hairs up to 700 μm long × 2.5–4.5 (–5) μm
diam., cylindrical, gradually narrowed to a
subacute tip; walls smooth, 0.5–2 μm thick,
hyaline to yellow, inamyloid to dextrinoid.
Stipitipellis with hairs like the pileipellis.
Habitat: Inserted on dead twigs of forest
litter. Known distribution: New Zealand.
Notes: This represents a good species of
Crinipellis distinctive because of the conical-
papillate, ochraceous pileus, long greyish
brown stipe (up to 100 mm), setulose
cheilocystidia, and the absence of rhizomorphs.
It is allied with C. setipes (Peck) Singer from
North America.
Status: Crinipellis procera is known
currently only from New Zealand.
32. Crinipellis readiae G. Stev., Kew Bull. 19:
43. 1964 (as Crinipellis readii). (Fig. 29: 1)
≡ Collybia stevensoniae E. Horak, N. Zealand J.
Bot. 9: 450. 1971 (as C. stevensonii comb. et nom. nov.),
non Collybia readiae G. Stev., Kew Bull. 19: 9. 1964.
Nom. illegit., a later homonym of Collybia stevensonii
(Berk. & Broome) Sacc., Syllog. Fung. 5: 226. 1887.
≡ Gymnopus readiae (G. Stev.) J.L. Mata,
Sydowia 58: 289. 2007.
Type: NEW ZEALAND, Puramahoi, 20
April 1955, coll. by D. Reid, G. Stevenson
Cone #985, in litter under Leptospermum
ericoides (Holotype: K!).
Analysis of the holotype specimen
The holotype specimen consists of about
2 very fragmented basidiomes in fair condition.
As dried: Pileus striate, pruinose overall,
reddish brown. Lamellae adnexed, subdistant
with 2 series of lamellulae, broad, cream-
coloured. Stipe 1 mm diam, pubescent,
yellowish grey-brown.
Basidiospores 8–9 × 3.8–4.5 µm [xm =
8.5 ± 0.3 × 4.1 ± 0.2 μm, Q = 1.8–2.3, Qm = 2.1
± 0.2, n = 10 spores per 1 collection], ellipsoid,
smooth, hyaline, inamyloid, thin-walled.
Basidia clavate, 4-spored, clamped. Basidioles
clavate. Pleurocystidia absent. Cheilocystidia
scattered, lamellar edge fertile; 22–40 × 6.5–10
µm, irregularly clavate to lageniform or
ventricose, mucronate or forked, projections
obtuse, hyaline, inamyloid, thin-walled.
Pileipellis a cutis of repent, subparallel,
cylindrical hyphae 3–5 µm diam, non-

148
Fig. 29. 1. Crinipellis readiae – a. Basidiospores b. Cheilocystidia – 2. Crinipellis roseola – a. Basidiospores b.
Cheilocystidia. – 3. Crinipellis velutipes – a. Basidiospores. – 4. Crinipellis vinacea – a. Basidiospores b. Cheilocystidia.
– Bar: 1-4 = 10 µm.
diverticulate, often with brown pigment
incrustations.
Notes: Horak (1971: 450) transferred the
species into Collybia as C. stevensonii because
the name Collybia readiae was already taken.
However, Horak was unaware that there was
already a Collybia stevensonii, so his nom. nov.
created an illegitimate homonym. Correcting
the gender ending to the feminine
“stevensoniae” for Stevenson’s species versus
“stevensonii” for the Berkeley & Broome taxon
does not legitimize the illegitimate homonym.
Status: This New Zealand species was
transferred recently into Gymnopus. It does not
represent a species of Crinipellis, but rather
belongs in Gymnopus sect. Vestipedes.
33. Crinipellis roseola G. Stev., Kew Bull. 19:
42. 1964. (Fig. 29: 2)
Type: NEW ZEALAND, Arthur’s Pass, 7
June 1954, coll. by R. Mason, det. by G.
Stevenson Cole #961, inserted on fallen dead
leaves of Nothofagus clifortioides (Holotype:
K!).
Analysis of the holotype specimen
The holotype specimen consists of about
12 intact to fragmented basidiomes in fair
condition. As dried: Pileus 1.5–2 mm diam.,
hemispherical, glabrous, pinkish orange.
Lamellae adnate with a short tooth to arcuate or
subdecurrent, subdistant, broad, pinkish orange,
edges concolorous. Stipe 10–14 × 0.5 mm,
glabrous, orange, non-insititious, on leaves and
small twigs.
Basidiospores 4.5–6.4 × 2.8–3.8 μm [xm
= 5.7 ± 0.7 × 3.4 ± 0.3 μm, Q = 1.4–1.8, Qm =
1.7 ± 0.1, n = 10 spores per 1 collection],
ellipsoid to pip-shaped, smooth, hyaline,
distinctly amyloid, thin-walled, mostly
collapsed. Basidia clavate, 4-spored.
Basidioles clavate. Pleurocystidia absent.
Cheilocystidia scattered, edge heteromorphous;
main body 22–25 × 5.4–6.4 μm, fusoid to
irregular in outline, simple, hyaline, inamyloid,
thin-walled. Pileipellis a cutis of repent hyphae
with a few erect terminal cells; hyphae 1.5–2.5
μm diam., cylindrical, smooth, hyaline,

Fungal Diversity
149
inamyloid, mostly repent and agglutinated;
terminal cells not differentiated, cylindrical,
mostly repent. Pileus trama little differentiated,
no distinct hypodermium; hyphae gelatinous,
dextrinoid. Stipe cortical hyphae 2.2–4 μm
diam., parallel, cylindrical, smooth, non-
gelatinous, hyaline, dextrinoid, clamped;
medullary hyphae similar but up to 10 μm
diam.
Habitat: On fallen dead leaves.
Known distribution: New Zealand.
Notes and Status: The holotype specimen
represents a small, pink species of Mycena.
Until further material is collected to better
diagnose the pileipellis anatomy, and until an
accounting of previously described Austra-
lasian Mycena species is evaluated, a formal
transfer will not be made.
34. Crinipellis substipitaria G. Stev., Kew
Bull. 19: 43. 1964. (Fig. 30: 1–4)
Type: NEW ZEALAND, Levin, Lake
Papaitonga, Buller Bush, 26 Oct. 1947, coll. by
G. Stevenson Cone #211, on fallen dead leaves
under tawa (Holotype: K!).
Analysis of the holotype specimen:
The holotype specimen consists of 5
intact or fragmented basidiomes in fair
condition, one attached to a leaf. As dried:
Pileus 3–4 mm diam., conical, radially
fibrillose, pale brown. Lamellae subdistant to
close, broad, cream-colored. Stipe 10–25 × 0.5
mm, shaggy, ochraceous; rhizomorphs absent.
One Mycena basidiome is also in the collection.
Basidiospores 8.6–10.5 × 6.4–7.4 (–7.7)
μm, [xm = 9.5 ± 0.7 × 7.0 ± 0.4 μm, Q = 1.2–
1.5, Qm = 1.4 ± 0.1; n = 10 spores per 1
collection], broadly ovoid, smooth, hyaline,
inamyloid, thin- to firm-walled. Basidia
clavate, clamped. Basidioles broadly fusoid.
Pleurocystidia absent. Cheilocystidia common,
lamellar edge heteromorphous; main body 14–
23 × 4.5–6.5 μm, subcylindrical to clavate with
numerous apical appendages; apical
appendages 1.5–4 × 1.2–1.5 μm, rod-like,
obtuse, seldom forked, hyaline, inamyloid,
thin-walled. Pileipellis of tangled to subparallel,
thick-walled hairs. Hairs > 300 × 3.8–5 μm,
cylindrical to sinuous, narrowed near the apex
to an obtuse tip, hyaline to yellowish brown,
inamyloid to strongly dextrinoid; walls smooth,
0.5–2.5 μm thick. Stipitipellis with hairs like
the pileipellis.
Habitat: On fallen dead leaves.
Known distribution: New Zealand.
Notes: This represents a good species of
Crinipellis, distinctive because of the very
broad ovoid spores and the clavate-setulose
cheilocystidia.
Status: Crinipellis substipitaria is known
currently only from New Zealand.
Fig. 30. Crinipellis substipitaria (Holotype). – 1. Basidiospores. – 2. Cheilocystidia. – 3. Basidioles. – 4. Pileipellis
hairs. – Bar: 1-4 = 10 µm.

150
35. Crinipellis velutipes G. Stev., Kew Bull.
19: 45. 1964. (Fig. 29: 3)
Type: NEW ZEALAND, Waikanae, 1
Jan. 1951, G. Stevenson Cone #792, on fallen
rotten log under Dysoxylon spectabile
(Holotype: K!).
Analysis of the holotype specimen
The holotype specimen consists of about
5, large, fragmented basidiomes in fair
condition, loose in the packet. As dried:
Pileus up to 30 mm diam., plano-convex,
striate, pellucid, pruinose, dark ferruginous.
Lamellae adnexed, distant with 2–3 series of
lamellulae, broad, concolorous with pileus.
Stipe 22–40 × 2–3 mm, cleft, pubescent,
mustard yellow-brown.
Basidiospores (6.5–) 7–9 × 4–5 μm [xm
= 7.9 ± 0.6 × 4.5 ± 0.3 μm, Q = 1.5–1.9, Qm =
1.7 ± 0.1, n = 20 spores per 1 collection],
subcylindrical to ellipsoid, smooth, hyaline,
strongly amyloid, thin-walled. Other
microdata not obtained.
Habitat: On fallen rotting log.
Known distribution: New Zealand.
Notes: Horak (1971: 459) indicated that
the holotype specimen was sterile. This is not
true. Basidiospores are very common and
strongly amyloid. Subsequently, Horak (1979:
150) considered the species as a synonym of
Heimiomyces neovelutipes (Hongo) E. Horak, a
species based on Collybia neovelutipes Hongo,
dating from 1974. Pegler (1977: 244)
considered C. velutipes as a synonym of
Heimiomyces tenuipes (Schwein.) Singer (as
Xeromphalina tenuipes), but apparently did not
examine the type collection of C. velutipes, and
after Horak published his report, Pegler (1983:
248) removed C. velutipes from his list of
synonyms. We agree that the old world,
Australasian and Southeast Asian taxon is
different from the New World H. tenuipes (as
suggested by Horak 1979), and that C.
velutipes (dating from 1964) may be
conspecific with H. neovelutipes.
Status: If the above-stated premise is true,
then the oldest epithet at species rank for this
taxon is velutipes, and the correct name should
be Heimiomyces velutipes (G. Stev.) comb.
prov. (with H. neovelutipes as a synonym).
Until further material of this species is
collected from New Zealand and compared
with Papua New Guinea material (type locality
of H. neovelutipes), a formal combination will
not be made.
36. Crinipellis vinacea G. Stev., Kew Bull. 19:
44. 1964. (Fig. 29: 4)
≡ Collybia vinacea (G. Stev.) E. Horak, N.
Zealand J. Bot. 9: 459. 1971.
Type: NEW ZEALAND, Nelson,
Wakapuaka, Palmer’s Bush, 14 April 1956,
coll. by G. Stevenson Cone #1029, in litter on
fallen rotting log (Holotype: K!).
Analysis of the holotype specimen:
The holotype specimen consists of two
different species: About 10 basidiomes in total
in fair condition, loose in the packet.
Taxon 1: As dried: Pileus 7–15 mm
diam., plano-convex, striate, appressed-
fibrillose, dark brown. Lamellae narrowly
adnexed, subdistant with 2 series of lamellulae,
narrow, greyish brown. Stipe up to 25 × 1 mm,
pruinose, dark brown. Cheilocystidia simple,
fusoid to clavate. Pileipellis a cutis of
cylindrical hyphae with dark brown
incrustations.
Notes and Status: This dark brown,
possibly vinaceous, species is the one reported
by Horak (1971: 459, Fig. 27:342) and may
represent what Stevenson had in mind for
Crinipellis vinacea. If the appropriate
basidiomes could be separated, this taxon
should be transferred as Gymnopus vinacea (G.
Stevenson) comb. prov. and belongs in Sect.
Vestipedes. A formal transfer will not be made
here. Taxon 2: As dried: Pileus 10–12 mm
diam., plano-convex, non-striate or striatulate,
glabrous, ochraceous. Lamellae adnexed,
subdistant with 2 series of lamellulae, broad,
cream- to clay-colored. Stipe 12–15 × 1 mm,
glabrous, reddish brown. Basidiospores 9.3–
11.2 × 4.8–5.7 μm [xm = 10.3 ± 0.7 × 5.3 ± 0.3
μm, Q = 1.8–2.3, Qm = 2.0 ± 0.1, n = 10 spores
per 1 collection], elongate-ellipsoid, smooth,
hyaline, inamyloid, thin-walled. Cheilocystidia
with main body 11–16 × 4–6.5 μm,
subcylindrical to clavate with numerous apical
diverticula; diverticula 1.5–5 × 1–2 μm, knob–
like, hyaline, thin-walled. Pileipellis a
Dryophila-structure of broad, short, branched
cells with clavate terminal cells, hyaline or
with brown plasmatic pigments.

Fungal Diversity
151
Notes and Status: This second entity
represents a species of Gymnopus (Pers.)
Roussel in Sect. Levipedes. A formal transfer
will not be made here.
Acknowledgments:
This research was funded in part by a National
Science Foundation PEET grant (DEB-0118776) to
Desjardin and a NSF GK12 grant (DGE-0337949 and
DGE-0086467) to Kerekes. We are most grateful to the
following institutions and colleagues for providing
essential Material Transfer Agreements (MTA) that
allowed us the ability to transfer specimens from their
countries to the US for study and deposition: Dr.
Saisamorn Lumyong, Chiang Mai University, Thailand;
Dr. S. Vikineswary, University of Malaya, Malaysia;
Atik Retnowati, Bogor Herbarium and Indonesian
Institute of Sciences, Java, Indonesia. We thank Dr.
Kevin D. Hyde, Mushroom Research Foundation,
Thailand, for providing reseach facilities at the
Mushroom Research Centre. We are thankful to Dr.
Brian Perry (SFSU) for helping with the molecular
analysis and for critical comments on the manuscript.
We thank two anonymous reviewers for providing
critical comments that helped improve the manuscript.
We would also like to thank all of our in-country hosts
and companions in the field.
References
Aime, M.C. and Phillips-Mora, W. (2005). The causal
agents of witches’ broom and frosty pod rot of
cacao (chocolate, Theobroma cacao) from a new
lineage of Marasmiaceae. Mycologia 97: 1012-
1022.
Bodensteiner, P., Binder, M., Moncalvo, J.M., Agerer,
R.S., and Hibbett, D. (2004). Phylogenetic
relationships of cyphelloid homobasidiomycetes.
Molecular Phylogenetics and Evolution 33: 501-
515.
Corner, E.J.H. (1996). The agaric genera Marasmius,
Chaetocalathus, Crinipellis, Heimiomyces,
Resupinatus, Xerula and Xerulina in Malesia,
Beiheft Nova Hedwigia 111: 1-175.
Desjardin, D.E. and Horak, E. (1999). Agaricales of
Indonesia. 1: A new cyphelloid genus
(Lecanocybe lateralis gen. et sp. nov.) from Java
and the Hawaiian Islands. Sydowia 51: 20-26.
Desjardin, D.E., Retnowait, A. and Horak, E. (2000).
Agaricales of Indonesia. 2. A preliminary
monograph of Marasmius from Java and Bali.
Sydowia 54: 92-194.
Gardes, M. and Bruns, T.B. (1993). ITS primers with
enhanced specificity for basidiomycetes -
application to the identification of mycorrhizae
and rusts. Molecular Ecology 2: 113-118.
Hibbett, D.S., Fukumasa-Nakai, Y., Tsuneda, A. and
Donoghue, M.J. (1995). Phylogenetic diversity in
shiitake inferred from nuclear ribosomal DNA
sequences. Mycologia 87: 618-638.
Holmgren, P.K., Keuken, W. and Schofield, E.K.
(1981).Index Herbariorum. Part I. The Herbaria
of the world. 7th edn. Regnum Vegetabile, Utrecht,
Netherlands.
Horak, E. (1971). A contribution towards the revision of
the Agaricales (Fungi) from New Zealand. New
Zealand Journal of Botany 93: 403-462.
Huelsenbeck, J. P. and F. Ronquist. (2001). MRBAYES:
Bayesian inference of phylogeny. Bioinformatics
17: 754-755.
Kirk P.M., Cannon, P.F., David, J.C., Stalpers, J.A.
(2001). Dictionary of the fungi. 9th Ed. United
Kingdom: CAB International: 655
Kornerup, A. and Wanscher, J.H. (1978). Methuen hand-
book of colour. 3rd Ed. London: Eyre Methuen:
252.
Largent, D.L. (1986). How to Identify Mushrooms to
Genus 1: Macroscopic Features. Mad River Press:
166 .
Largent, D.L., Johnson, D. and Watling, R. (1977). How
to identify Mushrooms to Genus. 3. Microscopic
Features. Mad River Press: 148.
Maddison, D.R. and Maddison, W.P. (2001). MacClade
4, version 4.03. Sinauer, Sunderland,
Massachusetts, USA.
Matheny, P.B., Curtis, J.M., Hofstetter, V., Aime, M.C.,
Moncalvo, J.-M., Ge, Z.-W., Yang, Z.-L., Slot,
J.C., Ammirati, J.F., Baroni, T.J., Bougher, N.L.,
Hughes, K.W., Lodge, D.J., Kerrigan, R.W., Seidl,
M.T., Aanen, D.K., DeNitis, M., Daniele, G.M.,
Desjardin, D.E., Kropp, B.R., Norvell, L.L.,
Parker, A., Vellinga, E.C., Vilgalys, R. and
Hibbett, D.S. (2006). Major clades of Agaricales:
a multilocus phylogenetic overview. Mycologia
98: 982-995.
Meyers, N., Mittermeir, R., Mittermeier, C.G., da
Fonseca, G.A.B. and Kent, J. (2000). Biodiversity
hotspots for conservation priorities. Nature 403:
853-858.
Moncalvo, J.M., Vilgalys, R., Redhead, S.A., Johnson,
J.E., James, T.Y., Aime, M.C., Hofstetter, V.,
Verduin, S.J.W., Larsson, E., Baroni, T.J., Thorn,
R.G., Jacobsson, S., Clemencon, H. and Miller,
O.K.Jr. (2002). One hundred and seventeen
clades of euagarics. Molecular Phylogenetics and
Evolution 23: 357-400.
Nylander, J.A.A. (2004). MrModeltest v2. Program
distributed by the author. Evolutionary Biology
Centre, Uppsala University.
Pegler, D.N. (1968). Studies on African Agaricales: I.
Kew Bulletin 21: 499-533.
Pegler, D.N. (1977). A preliminary agaric flora of East
Africa. Kew Bull. Add. Ser. 6: 1-615.
Pegler, D.N. (1983). Agaric flora of the Lesser Antilles.
Kew Bull. Add. Ser. 9: 1-668.
Pegler, D.N. (1986). Agaric flora of Sri Lanka. Kew
Bulletin Additional Series 12: 15-19.
Posada, D. and Crandall, K.A. (1998). Modeltest: testing
the model of DNA substituion. Systematic
Biology 50: 817-818.
Redhead, S.A. (1986). The genus Crinipellis in Canada.
Atti del Centro Studi per la flora Mediterranea 6:
175-199.

152
Redhead, S.A. (1989). A biogeographical overview of
the Canadian mushroom flora. Canadian Journal
of Botany 67: 3003–3062.
Ronquist, F. and Huelsenbeck, J. P. (2003). MRBAYES
3: Bayesian phylogenetic inference under mixed
models. Bioinformatics 19: 1572-1574.
Singer, R. (1943, “1942”). A Monographic Study of the
Genera Crinipellis and Chaetocalathus. Lilloa 8:
441-534.
Singer, R. (1976). Marasmieae (Basidiomycetes-
Tricholomataceae). Flora Neotropica Monograph
17: 1-347.
Singer, R. (1986). The Agaricales in modern taxonomy.
4th Ed. KoeltzScientific Books, Koenigstein,
Germany: 981.
Stevenson, G. (1964). The Agaricales of New Zealand:
V. Tricholomataceae. Kew Bulletin 19: 1-59.
Swofford, D.L. (2002). PAUP*. Phylogenetic analysis
using parsimony (*and other methods), version 4.
Sinauer, Sunderland, Massachusettes, USA.
Takahashi, H. (2000). Three new species of Crinipellis
found in Iriomote Island, southwestern Japan, and
central Honshu, Japan. Mycoscience 41: 171-182.
Takhtajan, A., Crovello, T.J. and Cronquist, A. (1986).
Floristic regions of the world. Berkeley:
University of California Press.
Tan Y.S., Desjardin, D.E., Vikineswary, S. and
Noorlidah, A. (2007). New species and mating
studies of Marasmius from Malaysia. Fungal
Diversity 25: 187-217.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin,
F. and Higgins, D.G. (1997). The ClustalX
windows interface: flexible strategies for multiple
sequence alignment aided by quality analysis
tools. Nucleic Acids Research 24: 4876-4882.
Vellinga, E. (1988). Glossary - Flora Agaricina
Neerlandica 1: 54-64.
Vizzini A., Antonin V., and Noordeloos M.E. (2007).
Crinipellis pedemontana sp. nov. (Agaricomy-
cetes), a new basidiomycete from Italy.
Mycologia 99(5): 786-791.
Wannathes, N., Desjardin, D.E. and Lumyong, S. (2007).
Mating studies, new species, and new reports of
Marasmius from northern Thailand. Mycological
Research 111: 985-996.
Wannathes, N., Desjardin, D.E., Retnowati A., Tan Y.S.,
and Lumyong, S. (2004). A redescription of
Marasmius pellucidus, a species widespread in
South Asia. Fungal Diversity 17: 203-218.
White, T.J., Bruns, T.D., Lee, S.B. and Taylor, J.W.
(1990). Amplification and direct sequencing of
fungal ribosomal genes for phylogenetics. In:
PCR protocols (eds. M.A. Innis, D.H. Gelfand,
J.J. Sninsky and T.J. White). San Diego,
California: Academic Press: 315-322.
Wilson, A.W. and Desjardin, D.E. (2005). Phylogenetic
relationships in the gymnopoid and marasmioid
fungi (Basidiomycetes, euagarics clade).
Mycologia 97: 667-679.
Wilson, A.W., Desjardin, D.E., and Horak, E. (2004).
Agaricales of Indonesia. 5. The genus Gymnopus
from Java and Bali. Sydowia 56: 137-210.
Yang, Z.-L. (2000). Type studies on agarics described by
N. Patouillard (and his co-authors) from Vietnam.
Mycotaxon 75: 431-476.
Zwickl, D.J. (2006). Genetic algorithm approaches for
the phylogenetic analysis of large biological
sequence datasets under the maximum likelihood
criterion. Ph.D. dissertation, The University of
Texas at Austin.