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Biodiversity and phylogeny of Marasmius (Agaricales, Basidiomycota) from Madagascar

January 2017
Phytotaxa 292(2):101

DOI:10.11646/phytotaxa.292.2.1

Authors:

Jackie E. Shay

University of California, Merced

Dennis E Desjardin

San Francisco State University

Brian A. Perry

California State University, East Bay

Christopher L. Grace

California Academy of Sciences

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Prior to this monographic treatment, limited research on the genus Marasmius (Basidiomycota, Agaricales) had been conducted in Madagascar. Based on field work in January 2013 and January–February 2014, which generated 45 specimens of Marasmius sensu stricto, supplemented by herbarium exsiccatae and published literature, 35 species of Marasmius are documented from Madagascar. Of these, 5 species are recognized herein as new to science, viz., Marasmius madagascariensis, M. rubrobrunneus, M. dendrosetosus, M. sokola and M. tanaensis; an additional 11 species represent new distribution records for Madagascar. Species delimitations are based on comprehensive morphological descriptions and molecular sequences (ITS) data. Line drawings of salient micromorphological features, color photographs of basidiomes, comparisons with allied taxa, a key to aid in identification, and phylogenetic inferences are provided.

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Phytotaxa 292 (2): 101–149

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Accepted by Genevieve Gates: 23 Dec. 2016; published: 25 Jan. 2017

https://doi.org/10.11646/phytotaxa.292.2.1

101

Biodiversity and phylogeny of Marasmius (Agaricales, Basidiomycota) from Madagascar

JACKIE E. SHAY1*, DENNIS E. DESJARDIN1, BRIAN A. PERRY2, CHRIS L. GRACE1 & DANNY S.

NEWMAN1

1Department of Biology, San Francisco State University, 1600 Holloway Ave., San Francisco, CA 94132, USA

2Department of Biological Sciences, California State University East Bay, 25800 Carlos Bee Blvd., Hayward, CA 94542 USA

*Corresponding Author: J.E. Shay, jeshay@mail.sfsu.edu

Abstract

Prior to this monographic treatment, limited research on the genus Marasmius (Basidiomycota, Agaricales) had been con-

ducted in Madagascar. Based on field work in January 2013 and January–February 2014, which generated 45 specimens

of Marasmius sensu stricto, supplemented by herbarium exsiccatae and published literature, 35 species of Marasmius are

documented from Madagascar. Of these, 5 species are recognized herein as new to science, viz., Marasmius madagascarien-

sis, M. rubrobrunneus, M. dendrosetosus, M. sokola and M. tanaensis; an additional 11 species represent new distribution

records for Madagascar. Species delimitations are based on comprehensive morphological descriptions and molecular se-

quences (ITS) data. Line drawings of salient micromorphological features, color photographs of basidiomes, comparisons

with allied taxa, a key to aid in identification, and phylogenetic inferences are provided.

Keywords: endemism, internal transcribed spacer, marasmioid fungi, saprotrophic, taxonomy

Introduction

Members of the mushroom genus Marasmius (Marasmiaceae) Fries (1836:339) play key roles in the decomposition of

wood and leaf litter, nutrient cycling, and soil genesis. Marasmius diversity has been found to strongly correlate with

plant host diversity, especially in tropical habitats (Lodge et al. 1995). There has been recent interest in understanding

the global biodiversity and evolution of Marasmius (Antonín, 2003, 2004a, b, 2007, 2013; Antonín & Buyck 2006;

Antonín & Noordeloos 2010; Desjardin 1990, Desjardin & Horak 1997, Desjardin & Ovrebo 2006, Desjardin et al.

2000, Tan et al. 2007, 2009; Wannathes et al. 2004, 2007, 2009a, b; Wilson & Desjardin 2005), although only limited

attention has been paid to the Marasmius of Madagascar. The island of Madagascar is a model tropical system that has

evolved in isolation, characterized by high levels of endemism and species diversification (Vences et al, 2009).

The earliest records of Marasmius from Madagascar were reported by Hennings (1893) based on collections

made by J. Braun in 1891 of Marasmius foetidus (Sowerby) Fries (1838:380) [= Gymnopus foetidus (Sowerby) P.M.

Kirk (2014:120)] from Ankoraka, and M. oreades (Bolton) Fries (1836:510). from a meadow at “Champ de Meclas”.

Hennings (1908) reported Marasmius rotula (Scop.) Fries (1838:385) and M. rhodocephalus Fries (1851:31) [= M.

haematocephalus (Mont.) Fries (1838:382)], both from SW Madagascar near Andranohinaly.

Collections of Marasmius and Androsaceus (a synonym of Gymnopus) made by Raymond Decary in 1923–1924

and sent to the Natural History Museum in Paris were studied by Patouillard (1928). These collections were made on the

central plateau of the island towards Antananarivo and Maromandia, and included the European Marasmius candidus

Fries (1838:381) [= Marasmiellus candidus (Fries) Singer (1948:129)] and M. ramealis (Bull.) Fries (1838:381) [=

Marasmiellus ramealis (Bull.) Singer (1948:130)], and tropical M. aureotomentosus (Kalchbr.) Patouillard (1928:23),

M. aculeatus Patouillard (1990:175) and M. sulcatipes Patouillard (1924:526) (nom. illeg. non M. sulcatipes

Murrill 1915). In addition, Patouillard (1928) reported four taxa of Androsaceus, viz., A. rhodocephalus (Fr.) Pat.

[= Marasmius haematocephalus (Montagne) Fries 1838:382)], A. haematocephalus (Montagne) Pat. (= Marasmius

haematocephalus), A. polyadelphus (Lasch) Patuyyar (1887:105) [= Mycena polyadelpha (Lasch) Kühner (1938:262)]

and introduced a new variety “substipitatus” for A. sessilis Patouillard [= Marasmius sessilis Patouillard (1896:132)].

George Métrod, who documented many fungi from Madagascar, studied Decary’s specimens, but his observations

SHAY ET AL.

were never published. The Decary specimens were kept in formol or alcohol and subsequently dried out in later years,

making them unsuitable for any descriptive purposes, particularly for describing new taxa.

In a preliminary investigation of Marasmius from the region, Antonín & Buyck (2006) reported 19 species of

Marasmius from Madagascar, Mauritius and Réunion, and noted that approximately one fourth of the Malagasy fungal

taxa appeared to be endemic. They documented the macro- and micromorphological characteristics for these taxa,

and provided a key to aid in identification. They did not provide molecular sequence data for any specimens. In

2007, Antonín published the first part of a monograph representing marasmioid genera from tropical Africa. This

study documented 110 taxa of Marasmius, however it was estimated that there might be 2–3 times more taxa. The

geographic region encompassed the continent of Africa between the Tropics of Capricorn and Cancer, but excluded the

island of Madagascar. Antonín (2013) followed with a supplement to the African Marasmius monograph in which he

reported an additional 40 taxa.

Fieldwork was conducted in Madagascar January–February 2014 by J.E. Shay and D.S. Newman, and in January

2013 by Taylor Lockwood. These excursions yielded 83 specimens of Marasmius sensu lato, of which 45 specimens

represented Marasmius sensu stricto treated herein. Analyses of the latter material resulted in the recognition of five

new species, 11 new distribution records for Madagascar, and recollection of five of the species reported by Antonín &

Buyck (2006). Species reported herein are delineated using morphological characteristics and DNA sequence data from

the nuclear ribosomal internal transcribed spacer region (ITS), the universal DNA barcode marker for Fungi (Schoch

et al. 2012). Comprehensive descriptions of macro- and micromorphological features, illustrations, photographs, a

dichotomous key to aid in identification, comparisons with similar taxa, and phylogenetic inferences are provided.

Materials and Methods

Sampling protocols

A variety of eastern rainforests in Madagascar were surveyed, chosen for their diversity of plants, habitats types, and

accessibility. Sites sampled include Ranomafana National Park, the forests near Andasibe, the littoral forests of the

east coast near Brickaville, and the Ambohitantily Reserve on the northern central plateau. All specimens matching the

morphotype of Marasmius sensu lato were collected, totaling 83 by the end of the expedition.

Morphological protocols

Data on substrate-type, GPS coordinates of collection site, and photographs of fresh material were documented in the

field at the time of collection. Soon after collection, macromorphological features of the basidiomes were recorded.

Portions of basidiomes were subjected to 3% potassium hydroxide (KOH) and Melzer’s reagent and observed staining

reactions were recorded. Colors terms and notations are from Kornerup & Wanscher (1978). Photographic images

were made with a Canon Rebel T3i EF–S 60MM F/2.8 Macro USM lens. Specimens were dried overnight using a

food dehydrator and packaged promptly in plastic bags to avoid condensation and subsequent mold contamination.

Micromorphological features were analyzed at SFSU by hand-sectioning dried material, rehydrating in 95% ethanol

followed by 3% KOH or Melzer’s reagent, and observed with an Olympus CH-30 or Nikon Optiphot-2 compound

microscope. Line drawings were made with the aid of a drawing tube. The term “inamyloid” refers to a negative reaction

in Melzer’s reagent (neither amyloid nor dextrinoid). Spore statistics include: xm, the arithmetic mean of the spore

length by spore width (± standard deviation) for n spores measured in a single specimen; xmr, the range of xm values and

xmm, the mean of xm values where more than one specimen is available; Q, the quotient of spore length by spore width,

indicated as a range in variation in n spores measured; Qm, the mean of Q values (± standard deviation) in a single

specimen; Qmr, the range of Qm values and Qmm, the mean of Qm values where more than one specimen is available;

n, the number of spores measured per specimen; s, the number of specimens involved. The micromorphological

analyses indicated that only 45 of the 83 specimens of Marasmius sensu lato represented Marasmius sensu stricto

and are included in this monographic treatment. The remaining 38 specimens represent members of the Gymnopus/

Marasmiellus lineage and will be treated in the future.

Herbarium specimens of Madagascar material previously collected by Antonín and Buyck (2006) were borrowed

from the Moravian Museum (BR–Brno, Czech Republic) and the Muséum National D´Histoire Naturelle (PC–Paris,

France), examined, sequenced, and compared with material collected during our expedition.

Molecular protocols

Genomic DNA was extracted from dried herbarium specimens using the E.Z.N.A. Forensic DNA Extraction Kit (Omega

Bio-tek Inc., Norcross, GA) following provided protocol. PCR was performed with 4AccuPower® HotStart PCR

PreMix (Bioneer, Daejeon, Korea) following the methods outlined in Cubero et al. (1999). The internal transcribed

spacer (ITS) regions 1 and 2, as well as 5.8S rDNA, were amplified using primers ITS1-F and ITS4 (Gardes & Bruns

1993; White et al. 1990). DNA fragments were amplified on an MJ Research PTC-200 Peltier Thermal Cycler (GMI,

Ramsey, Minnesota, USA). The thermocycling profile was as follows: an initial denature at 94˚C for five minutes, 39

cycles of denaturing at 94˚C for 30 seconds, annealing at 57˚C for 30 seconds, and extension at 72˚C for 45 seconds.

The final extension was at 72˚C for seven minutes. PCR products were purified using ExoSAPIT (USB Corporation,

Cleveland, OH, USA). Cycle sequencing was performed with ITS1-F and ITS4 primers using BigDye® Terminator

v3.1 (Thermo Fisher Scientific Inc., Life Technologies Corporation, Grand Island, NY, USA) and visualized on an ABI

PRISM® 3100 Genetic Analyzer (PE Biosystems and Hitachi, Ltd., Life Technologies, Carlsbad, CA, USA). Sequence

editing was performed with Geneious v.7 software (Kearse et al. 2012; Biomatters, Auckland 1010, New Zealand).

Maximum likelihood analysis (Felsenstein 1981) was conducted using RAxML 8.1.11 (Stamatakis 2014) under

the GTRGAMMA model using default parameters and run on the CIPRES Science Gateway (Miller et al. 2010), with

node support estimated via 1000 RAxML rapid bootstrap (BS) replicates. Bayesian analyses were performed using

Metropolis Coupled MCMC methods as implemented in MrBayes 3.2.6 (Huelsenbeck & Ronquist 2001; Ronquist

& Huelsenbeck 2003; Ronquist et al. 2012) under a GTR+I+G model of sequence evolution as determined under

the Akaike Information Criterion in PAUP* (Swofford 2002). Analyses consisted of two parallel searches, run for 25

million generations on the CIPRES Science Gateway (Miller et al. 2010), and initiated with random starting trees. Eight

chains (temp = 0.3) were sampled every 2500 generations for a total of 10,001 trees each, sampled from the posterior

distribution. Those topologies sampled prior to the runs reaching a split deviation frequency of 0.1 were discarded,

while the remaining topologies were used to calculate the posterior probabilities (PP) of the individual clades. The

default settings were used in MrBayes to set unconstrained branch lengths and uninformative topology priors. All ITS

sequences generated as part of this study are deposited in GenBank (Table 1; accessions KX148977–KX149019). The

aligned ITS dataset and associated tree files are deposited in TreeBase (submission ID 19993).

TABLE 1 List of Marasmius specimens sequenced (ITS1–5.8S–ITS2) for this study

Species Section/Subsection or Section/Series Collection No. GenBank No.

M. andasibensis var. obscurostipitatus Marasmius/Marasmius Buyck 00.1699b KX149005

M. apatelius Marasmius/Marasmius JES 150 KX148997

M. apatelius Marasmius/Marasmius JES 203 KX148998

M. bambusiniformis Sicci/Haematocephali JES 199 KX148990

M. bekolacongoli Globulares Lockwood 2131638 KX148982

M. brunneoaurantiacus Marasmius/Sicciformes Buyck 99.450 KX148978

M. brunneoaurantiacus Marasmius/Sicciformes JES 113 KX149009

M. brunneoaurantiacus Marasmius/Sicciformes JES 125 KX149010

M. brunneoaurantiacus Marasmius/Sicciformes JES 115 KX149011

M. brunneoaurantiacus Marasmius/Sicciformes JES 137 KX149012

M. brunneoaurantiacus Marasmius/Sicciformes JES 166 KX149013

M. brunneoaurantiacus Marasmius/Sicciformes JES 218 KX149014

M. brunneoaurantiacus Marasmius/Sicciformes JES 133 KX149016

M. corrugatiformis Sicci/Atrorubentes Buyck 97.425 KX148981

M. curreyi Marasmius/Sicciformes Buyck 97.374 KX148980

M. aff. curreyi Marasmius/Sicciformes JES 135 KX149008

M. dendrosetosus Sicci/Spinulosi JES 205 KX148995

M. dendrosetosus Sicci/Spinulosi JES 211 KX148996

...Continued on next page

SHAY ET AL.

TABLE 1. (Continued)

Species Section/Subsection or Section/Series Collection No. GenBank No.

M. ferruginoides Sicci/Haematocephali JES 209 KX148983

M. haematocephalus Sicci/Haematocephali Buyck 00.1820 KX148977

M. haematocephalus Sicci/Haematocephali JES 110 KX148984

M. haematocephalus Sicci/Haematocephali JES 193 KX148985

M. haematocephalus Sicci/Haematocephali JES 202 KX148986

M. haematocephalus Sicci/Haematocephali JES 142 KX148987

M. hinnuleus Sicci/Haematocephali JES 217 KX148988

M. katangensis Sicci/Atrorubentes JES 227 KX148991

M. madagascariensis Marasmius/Sicciformes JES 225 KX149006

M. madagascariensis Marasmius/Sicciformes JES 139 KX149015

M. megistus Sicci/Leonini JES 163 KX148992

M. megistus Sicci/Leonini Lockwood 2132155 KX148993

M. neosessiliformis Neosessiles Buyck 97.615 KX149007

M. nummularius Sicci/Spinulosi JES 121 KX148979

M. rotalis Marasmius/Marasmius JES 141 KX148999

M. rotalis Marasmius/Marasmius JES 145 KX149000

M. rotalis Marasmius/Marasmius JES 150B KX149001

M. rubrobrunneus Marasmius/Sicciformes JES 191 KX148989

M. sokola Sicci/Leonini JES 154 KX148994

M. somalomoensis Marasmius/Marasmius JES 129 KX149002

M. somalomoensis Marasmius/Marasmius JES 165 KX149003

M. somalomoensis Marasmius/Marasmius JES 181 KX149004

M. cf. subruforotula Marasmius/Sicciformes JES 186 KX149017

M. cf. subruforotula Marasmius/Sicciformes JES 192 KX149018

M. cf. subruforotula Marasmius/Sicciformes JES 190 KX149019

Results

Phylogenetic Analysis

Phylogenetic analyses were performed on 135 taxa of Marasmius sensu stricto representing a global sampling of the

genus, and including representatives of all infrageneric groups. Of the 222 ITS sequences included in the dataset, 154

were downloaded from GenBank, and 68 were generated for this study, including 43 sequences of Marasmius from

Madagascar (Table 1). Crinipellis malesiana Kerekes, Desjardin & Vikineswary (2009:125) was chosen as the outgroup

based on the results of previous research (Tan et al. 2009, Wannathes et al. 2009a). A total of 343 ambiguously aligned

nucleotides were removed prior to all analyses, resulting in a final dataset of 687 nucleotide characters.

FIGURE 1a. Maximum likelihood phylogeny based on ITS sequence data. Marasmius from Madagascar are indicated in bold type. Values

separated by / refer to nonparametric ML bootstrap proportions and Bayesian posterior probabilities, respectively. Values greater than

70/0.70 are shown (- designates a value below 70% or below 0.70). Nodes receiving support values greater than 90/0.95 are represented

by bold branches. G—sect. Globulares; N—sect. Neosessiles; SA—sect. Sicci ser. Atrorubentes; SL—sect. Sicci ser. Leonini; SH—sect.

Sicci ser. Haematocephali.

SHAY ET AL.

FIGURE 1b. Maximum likelihood phylogeny based on ITS sequence data. Marasmius from Madagascar are indicated in bold type. Values

separated by / refer to nonparametric ML bootstrap proportions and Bayesian posterior probabilities, respectively. Values greater than

70/0.70 are shown (- designates a value below 70% or below 0.70). Nodes receiving support values greater than 90/0.95 are represented

by bold branches. G—sect. Globulares; SA—sect. Sicci ser. Atrorubentes; SL—sect. Sicci ser. Leonini; SS—sect. Sicci ser. Spinulosi;

SH—sect. Sicci ser. Haematocephali.

FIGURE 1c. Maximum likelihood phylogeny based on ITS sequence data. Marasmius from Madagascar are indicated in bold type. Values

separated by / refer to nonparametric ML bootstrap proportions and Bayesian posterior probabilities, respectively. Values greater than

70/0.70 are shown (- designates a value below 70% or below 0.70). Nodes receiving support values greater than 90/0.95 are represented by

bold branches. G—sect. Globulares; N—sect. Neosessiles; L—sect. Leveilleani; MM—sect. Marasmius subsect. Marasmius; MS—sect.

Marasmius subsect. Sicciformes; SA—sect. Sicci ser. Atrorubentes; SL—sect. Sicci ser. Leonini; SS—sect. Sicci ser. Spinulosi; SH—sect.

Sicci ser. Haematocephali.

The RAxML tree with best ML score is presented in Fig. 1 (-lnL = -13820.463). In general, statistical support for

the deep nodes distinguishing major lineages was low, indicating that the ITS gene region is too variable to delimit

infrageneric clades in Marasmius. Only sect. Marasmius subsect. Marasmius, including the type species of the genus

Marasmius (M. rotula) (Fig. 1c), was monophyletic (98% BS, 1.0 PP). All members of this group form lamellae

attached to a collarium, have an insititious stipe and Rotalis-type broom cells in the pileipellis. Members of sect.

Marasmius subsect. Sicciformes, which also form a collarium and an insititious stipe, but have Siccus-type broom

SHAY ET AL.

cells, formed a grade basal to subsect. Marasmius (Fig. 1c), and included a few members of sects. Globulares + Sicci.

There was low support for these relationships. The historically recognized sects. Globulares + Sicci (lamellae lacking

a collarium, with a non-insititious stipe, Globulares-type or Siccus-type boom cells) formed a clade with low support

(Figs. 1a, 1b) and included most members traditionally placed there, except for a few species that appear to be more

closely related to members of sect. Marasmius subsect. Sicciformes (Fig. 1c), as noted above.

Members of sect. Neosessiles occur in two distinct parts of the tree. The recently described M. griseoroseus

(Montagne) Singer (1976:256) and M. conchiformis de Oliveira & Capelari (2014:5), plus M. elaeocephalus Singer

(1964:384) (a member of sect. Sicci ser. Haematocephali) align with members of sect. Globulares + Sicci (Fig. 1a),

whereas M. tenuissimus (Jungh.) Singer (1976:258) and M. neosessiliformis Antonín & Buyck (2006:34) are sister

to sect. Leveilleani and together are sister to one clade of sect. Marasmius subsect. Sicciformes (Fig.1c). Marasmius

nodulocystis Pegler (1977:200) (sect. Sicci ser. Leonini) is morphologically nearly indistinguishable from M.

leveilleanus (Berk.) Saccardo (1925:149) (type species of sect. Leveilleani) and the ITS sequences of three specimens

of M. nodulocystis are sister to a sequence of M. leveilleanus from Thailand.

None of the historically recognized infrageneric groups within sect. Sicci, viz., ser. Atrorubentes (SA),

Haematocephali (SH), Leonini (SL), and Spinulosi (SS) are monophyletic in the ITS phylogeny (Figs. 1a, 1b), although

small groups of species within each of these infrageneric groups do form clades but with limited support. For the most

part, members of these series are scattered throughout a clade containing most of sects. Globulares (G) + Sicci.

Concerning Madagascar taxa in Fig.1a: Marasmius katangensis (SA) Singer (1964:375) is sister to a clade containing

M. occultatiformis (SL) Antonín, Ryoo & H.D. Shin (2012:616), M. cf. cladophyllus (SL) Berkeley (1856:138) and

M. aurantioferrugineus (G) Hongo (1965:74) with low support. Marasmius corrugatiformis (SA) Singer (1964:374)

is sister to M. subarborescens (SA) Singer (1964:364) with low support. Marasmius bambusiniformis (SL) Singer

(1976:1C7) is sister to M. berteroi (SL) Murrill (1915:267) with 100% BS and 1.0 PP support. Marasmius ferruginoides

(SH) Antonín (2004:403) holds an isolated position basal to numerous members of sects. Globulares + Sicci, in a grade

with other members of ser. Haematocephali. Marasmius hinnuleus (SH) Berkeley & Curtis (1868:297) is on a long

branch sister to M. grandisetulosus (SH) Singer (1964:379) with low support. Two sequences of Marasmius megistus

(SL) Singer (1964:356) from Madagascar form a well-supported clade with a sequence from São Tomé (100% BS, 1.0

PP) and are on a long branch sister to members of ser. Haematocephali.

Concerning Madagascar taxa in Fig. 1b: The new species M. dendrosetosus (SS) is sister to M. longisetosus (SS)

de Oliveira & Capelari (2014:19) with 99% BS and 1.0 PP support, in a clade with other members of ser. Spinulosi. A

sequence of Marasmius nummularius (SS) Berkeley & Broome (1873:33) from Madagascar forms a well-supported

clade with two sequences from Thailand (90% BS, 1.0 PP), in a clade with other members of ser. Spinulosi. Five

sequences of M. haematocephalus (SH) align with several sequences from Thailand with 99% BS support, and are

sister to M. pulcherripes (SH) Peck (1872:77) but with low support. Marasmius bekolacongoli (G) Beeli (1928:157)

forms an unresolved trichotomy with other members of sect. Globulares plus M. coarctatus (SS) Wannathes, Desjardin

& Lumyong (2009:251). The new species M. sokola (SL) is on a long branch sister to M. imitarius (SL) Wannathes,

Desjardin & Lumyong (2009:279) with low support.

Concerning Madagascar taxa in Fig. 1c: Three sequences of M. somalomoensis (MM) Antonín (2003:66) are

unresolved in a clade containing other members of sect. Marasmius subsect. Marasmius with low support. Three

sequences of Madagascar M. rotalis (MM) Berkeley and Broome (1873:40) form a well-supported clade (98% BS, 1.0

PP) sister to a GenBank sequence of M. rotalis plus other members of subsect. Marasmius. Two sequences of Madagascar

M. apatelius (MM) Singer (1964:332) form a grade with other M. apatelius sequences from Thailand and Príncipe and

M. andasibensis var. obscurostipitatus (MM) Antonín & Buyck (2006:23) from Madagascar, but with low support. Three

sequences of M. cf. subruforotula (MS) Singer (1964:339) form a well-supported clade (99% BS, 1.0 PP) and are sister

to several sequences of M. subruforotula (MS) from Príncipe with 100% BS and 1.0 PP support. The new species M.

rubrobrunneus (MS) is on a long branch sister to M. purpureobrunneolus (MS) Hennings (1899:151) with 81% BS and

1.0 PP support. A sequence of Marasmius curreyi (MS) Berkeley & Broome (1879:209) from Madagascar (KX148980)

forms an unresolved clade with a sequence of M. curreyi from North Carolina (FJ431237) plus M. graminum (MS)

Berkeley (1860:22) with low support. This clade is sister to a well-supported clade (98% BS, 1.0 PP) clade containing

a Madagascar sequence of M. aff. curreyi (MS) plus a Korean sequence of M. curreyi (FJ936152). Eight sequences of

M. brunneoaurantiacus (MS) Antonín & Buyck (2006:24) form a well-supported clade (100% BS, 1.0 PP) sister to

other members of subsect. Sicciformes. Marasmius neosessiliformis (N) forms an unresolved clade with Malaysian and

Thailand sequences of M. tenuissimus (N) plus a Malaysian sequence of M. leveillianus (L) with low support. The new

species M. madagascariensis (MS) is on a long branch sister to a clade containing Malaysian sequences of M. guyanensis

(MS) Montagne (1854:114) and M. crinisequi (MS) with 0.97 PP support.

Tests of Sectional Monophyly

Two maximum likelihood analyses were conducted in RAxML: 1) an unconstrained analysis; and 2) an analysis that

constrained the tree into two monophyletic groups representing the major division of the genus Marasmius into section

Globulares and section Marasmius. The resulting constrained and unconstrained topologies were compared using the

Shimodaira-Hasegawa (SH) test (Simodaira & Hasegawa 1999) as implemented in PAUP* (Swofford 2002) using

1000 RELL bootstrap replicates. Results of the SH test indicate no statistically significant difference between these

topologies, and we are therefore unable to reject the monophyly of these sections given the data presented.

Artificial Key to Marasmius of Madagascar

1. Lamellae attached to a distinct collarium; stipe insititious (sect. Marasmius) ...................................................................................2

- Lamellae attached directly to the stipe or free and collarium absent; stipe not insititious ...............................................................17

2. Pileipellis composed of Rotalis-type broom cells (subsect. Marasmius) ...........................................................................................3

- Pileipellis composed of Siccus-type broom cells (subsect. Sicciformes) ...........................................................................................7

3. Pileus white, pale grey, or buff ........................................................................................................................................1. M. rotalis

- Pileus greyish orange, tan, or pale brown ..........................................................................................................................................4

4. Pileus tan to pale brown; lamellae subdistant (11–14) .....................................................................................2. M. somalomoensis

- Pileus greyish orange to buff; lamellae distant (9–12) ......................................................................................................................5

5. Pileus with a dark brown papilla ................................................................................................................................ 3. M. apatelius

- Papilla absent or concolorous with pileus surface ..............................................................................................................................6

6. Papilla absent; stipe pale brown ............................................................................................... 4. M. andasibensis var. andasibensis

- Papilla present, concolorous with pileus surface; stipe brown-black ................................5. M. andasibensis var. obscurostipitatus

7. Pileus brownish black or black ...........................................................................................................................................................8

- Pileus differently colored (reddish brown, brown, greyish orange, tan or cream) .............................................................................9

8. Basidiospores 8.5–10.5 × 4–5 µm; cheilocystidia 10–25 × 7–12 µm; pileus bicolored black and grey ............... 6. M. nigrogriseus

- Basidiospores 9–12 × 4.5–6.5 µm; cheilocystidia 10–14 × 5–8 µm; pileus brownish black ............................ 7. M. nigrobrunneus

9. Pileus reddish brown, or white, cream, tan to light brown ...............................................................................................................10

- Pileus greyish orange to brownish orange ........................................................................................................................................15

10. Pileus white to cream with prominent, dark brown conical papilla ...............................................................8. M. conicopapillatus

- Pileus more deeply pigmented .........................................................................................................................................................11

11. Pileus reddish brown with pinkish tinge at center and paler yellow-brown towards margin, or tan to light brown .......................12

- Pileus dark reddish brown ................................................................................................................................................................14

12. Pileus tan to light brown .........................................................................................................................................10. M. aff. curreyi

- Pileus reddish brown with or without pink tones .............................................................................................................................13

13. Basidiospores 8.5–10.5 × 4.5–5.5 µm; cheilocystidia of one type ................................................................................. 9. M. curreyi

- Basidiospores 10–12.5 × 4.5–6 µm; cheilocystidia of two types ..................................................... 11. M. curreyi var. bicystidiatus

14. Basidiospores mean range 14.9–16.5 × 3.2 µm; lamellae distant (11–12) ...................................................... 12. M. rubrobrunneus

- Basidiospores mean range 8.8–9.5 × 4.1–4.6 µm; lamellae subdistant (12–16) .................................... 13. M. brunneoaurantiacus

15. Basidiomes arising directly from black rhizomorphs; pileus 1–2 mm diam; lamellae distant (6); stipe 2–4 mm long .......................

..................................................................................................................................................................................14. M. crinisequi

- Basidiomes not arising directly from black rhizomorphs; pileus 1–7 mm diam; lamellae subdistant (9–12); stipe 6–40 mm long ...

..........................................................................................................................................................................................................16

16. Basidiospores mean range 8–8.4 × 3.8–3.9 µm; lamellae white to buff ...................................................... 15. M. cf. subruforotula

- Basidiospores mean range 11.3–11.6 × 4.8–5 µm; lamellae light orange to cream ....................................16. M. madagascariensis

17. Stipe absent or very short and lateral to strongly eccentric (sect. Neosessiles) ...............................................................................18

- Stipe well-developed and central (sect. Globulares + Sicci) ...........................................................................................................19

18. Pileus 1–4 mm diam, reddish brown; basidiospores 10–11 × 5–6 µm, ellipsoid; pleurocystidia absent .......17. M. neosessiliformis

SHAY ET AL.

- Pileus 5–8 mm diam, orangish brown to yellowish brown or pink; basidiospores 14.5–17 × 6.5–8 µm, lacrimoid or clavate-fusoid;

pleurocystidia present ........................................................................................................................................ 18. M. cecropiformis

19. Pileipellis composed of smooth, broadly clavate, non-setulose cells (sect. Globulares) .................................................................20

- Pileipellis composed of Siccus-type broom cells (sect. Sicci) ..........................................................................................................21

20. Pileus 100–120 mm diam, pale mouse grey overall; pleurocystidia present ...........................................................19. M. sulcatipes

- Pileus up to 85 mm diam, striped, brown to violaceous brown with pinkish-violaceous sulcae and cream ridges; pleurocystidia

absent .................................................................................................................................................................20. M. bekolacongoli

21. Stipe pruinose to hispid, ornamented with simple smooth cystidia or setae ....................................................................................22

- Stipe glabrous, simple cystidia and setae absent ..............................................................................................................................25

22. Setae present on pileus, lamellae and/or stipe (ser. Spinulosi) .........................................................................................................23

- Setae absent, stipe with simple smooth cystidia (ser. Atrorubentes) ................................................................................................24

23. Pileosetae branched, common, up to 300 µm long; pleurocystidia absent; basidiospores mean range 8.2–9.1 × 3.9 µm; caulosetae

absent, caulocystidia of Siccus-type broom cells ............................................................................................. 21. M. dendrosetosus

- Pileosetae unbranched, rare; pleurocystidia present; basidiospores mean range 12.4–12.7 × 4.3–4.8 µm; caulosetae abundant .......

............................................................................................................................................................................22. M. nummularius

24. Basidiospore mean 9.0 × 4.0 µm; caulocystidia dextrinoid, of one type (non-setulose cells) ...................... 23. M. corrugatiformis

- Basidiospores mean 7.8 × 4.0 µm; caulocystidia inamyloid, of two types (non-setulose cells and Siccus-type broom cells) ............

...............................................................................................................................................................................24. M. katangensis

25. Pleurocystidia absent (ser. Leonini) ..................................................................................................................................................26

- Pleurocystidia present (ser. Haematocephali) ..................................................................................................................................29

26. Pileus 20 mm diam, dark brown ................................................................................................................................... 25. M. sokola

- Pileus 4–9 mm diam, orangish ochraceous, reddish orange to orange or pale violet brown with reddish grey sulcae ...................27

27. Stipe 3–4 mm long; pileus pale orangish ochraceous ............................................................................................ 26. M. rammelooi

- Stipe 25–115 mm long; pileus reddish orange to orange, or pale violet brown with reddish grey sulcae ......................................28

28. Pileus 6.5–9 mm diam, violet brown with reddish grey sulcae; stipe 104–115 mm long; basidiospore mean 32 × 2.5 µm ...............

....................................................................................................................................................................................27. M. megistus

- Pileus 4–5 mm diam, reddish orange to orange; stipe 25–30 mm long; basidiospore mean 16.4 × 3.5 µm ........................................

...................................................................................................................................................................... 28. M. bambusiniformis

29. Basidiospores 12–22 µm long, with mean length 13.4–21 µm ........................................................................................................30

- Basidiospores 8–12.8 µm long, with mean length 10–12 µm ..........................................................................................................33

30. Pileus pale orangish pink, pinkish purple, dull reddish pink or red; basidiospores 16–22 µm long ........... 29. M. haematocephalus

- Pileus orangish pink, dark brown, light brown or reddish brown; basidiospore 11.2–18 µm long..................................................31

31. Pileus orangish pink, 1–3 mm diam; stipe up to 22 mm long; pleurocystidia 6–9.5 µm diam .................................30. M. tanaensis

- Pileus brown to reddish brown, 3–15 mm diam; stipe up to 40 mm long; pleurocystidia 8–15 µm diam ......................................32

32. Pileus 5–15 mm diam, dark brown at center, light brown towards the margin; basidiospores 15–18 × 4–5 µm; pleurocystidia 38–65

× 9–15 µm, subfusoid, often rostrate; pileipellis cells with 8–20 projections up to 6 µm long ................31. M. cf. grandisetulosus

- Pileus 3–5 mm diam, brown to reddish brown; basidiospores 13.5–16 × 3.5–5 µm; pleurocystidia 30–50 × 8–14 µm, clavate to

subfusoid, sometimes rostrate; pileipellis cells with up to 40 projections up 10 µm long .....................................32. M. eyssartieri

33. Pileus up to 3 mm diam; Siccus-type broom cells present on stipe apex ................................. 33. M. cf. confertus var. parvisporus

- Pileus 6–9 mm diam; Siccus-type broom cells absent on stipe apex ...............................................................................................34

34. Pileus orange to reddish orange; lamellae close (17–20); pleurocystidia 5–7.2 µm diam ................................34. M. ferruginoides

- Pileus brownish orange to reddish brown; lamellae distant (15–17); pleurocystidia 7–10 µm diam ..................... 35. M. hinnuleus

Taxonomy

I. Sect. Marasmius

Marasmius, II. Mycena, 2. Rotulae Fr., Epicr.: 384. 1838.

= Marasmius B. Rotulae Quél., Enchir.: 145. 1886.

= Marasmius sect. Rotulae Kühner, Botaniste 25: 98. 1933.

= Marasmius, I. Rotularia J. Schröt. in Cohn, Kryptog.-Fl. Schles. 3(1): 556. 1889.

= Marasmius, “sect.” Setipedes, α Collariati Bataille, Marasmes Eur.: 26. 1919.

= Marasmius sect. Pararotulae Singer, Sydowia 18: 140. 1965.

– Type species: Marasmius rotula (Scop.) Fr.

Ia. Subsect. Marasmius

= Marasmius sect. Pararotulae Singer, Sydowia 18: 336. 1965.

= Marasmius sect. Marasmius, subsect. Pararotulae (Singer) Singer, Fl. Neotrop. Monogr. 17: 92. 1976.

– Type species: Marasmius rotula (Scop.) Fr.

1. Marasmius rotalis Berk. & Broome, J. Linn. Soc., Bot. 14: 40. 1873 (1875). (Fig. 2, Plate 1)

Type:—SRI LANKA. Peradeniya, Thwaites 810 (K!)

Description:—Pileus 1–5 mm diam, campanulate to umbilicate, shallowly depressed; margin plicate to sulcate,

crenate; surface dry, glabrous; white to buff or pale gray (5B2–3). Context thin, concolorous. Lamellae adnate to a

collarium, distant (8–11), no lamellulae, non-intervenose, broad (0.5–0.8 mm), white, non-marginate. Stipe 8–54 × 0.2

mm, central, cylindrical, hollow, wiry, insititious; surface glabrous; dark brown to black overall. Odor and taste not

distinctive.

FIGURE 2. Marasmius rotalis (JES 141, JES 145, JES 150B) a) basidiospores; b) basidioles; c) cheilocystidia; d) pileipellis cells. Scale

bar = 10 µm. Illustrated by J.E. Shay.

Basidiospores (6.4–) 7.2–10.4 (–11.2) × 3.2–4.8 (–5) µm [xmr = 8.4–8.9 × 3.7–4.2 µm; xmm = 8.7 ± 0.3 × 4.0 ± 0.3

µm; Q = 1.5–3.3; Qmr = 2.09–2.28; Qmm = 2.19 ± 0.1, n = 24–25, s = 3], ellipsoid, smooth, hyaline, inamyloid, thin-

walled. Basidia not observed. Basidioles 13.6–23.2 × 4–8 µm, clavate to fusoid, some utriform, hyaline, inamyloid,

thin-walled. Cheilocystidia numerous, of Rotalis-type broom cells; main body 6.8–20 × 8–22.4 µm, clavate to broadly

clavate, globose, subglobose or obpyriform, hyaline, inamyloid, thin-walled; divergent setulae 0.5–2.4 × 0.5–2.4

µm, cylindrical to conical, obtuse, hyaline, inamyloid, thin-walled. Pleurocystidia absent. Pileipellis not mottled, a

hymeniform layer of Rotalis-type broom cells; main body 8–34 × 8–28 µm, clavate to broadly clavate, subglobose or

globose, pale yellowish brown to hyaline, inamyloid, thin-walled; divergent setulae 0.5–3 × 0.2–1.6 µm, numerous,

cylindrical to conical, pale yellowish brown to hyaline, inamyloid, thin-walled. Pileus trama interwoven; hyphae 1.6–

4.8 µm diam, cylindrical, smooth, hyaline, inamyloid, thin-walled. Lamellar trama regular; hyphae 1.6–9.6 µm diam,

cylindrical to slightly inflated, smooth, hyaline, inamyloid, thin-walled. Stipe tissue monomitic; cortical hyphae 3–4

SHAY ET AL.

µm diam, parallel, cylindrical, smooth, yellowish brown to brown, dextrinoid, thick-walled; medullary hyphae 1.6–6

µm diam, parallel, cylindrical, smooth, hyaline, inamyloid, thin-walled. Caulocystidia absent. Clamp connections

present.

Habit, habitat, and known distribution:—Solitary or gregarious on leaves of Cephalostachium vigueri (bamboo),

Cynodon dactylon (Poaceae), and various unknown dicotyledonous leaves and stems. Africa (Benin, Cameroon, DR

Congo, Kenya, Malawi, Nigeria, Tanzania, Uganda), Indonesia (Java), Madagascar, Papua New Guinea, South America

(Colombia), Sri Lanka.

Material examined:—MADAGASCAR. Commune Ranomafana, District Ifanadiana, Region Vatovavy-Fitovinany,

Ranomafana National Park, Circuit Vohiparara, elev.1062 m, GPS: 21˚ 14.255’ S, 47˚ 23.409’ E, 21 January 2014, J.E.

Shay 141 (SFSU) & J.E. Shay 145 (SFSU); Piste B, elev.1004 m, GPS: 21˚ 15.413’ S, 47˚ 25.253’ E, 22 January 2014,

J.E. Shay 150B (SFSU).

Notes:—Marasmius rotalis forms small (1–5 mm diam), white to pale grayish pilei, distant (8–11), collariate

lamellae, dark brown, wiry insititious stipe, basidiospores in the range 7.2–10.4 × 3.2–5 µm, Rotalis-type broom cells,

and growth on dicotyledonous leaves, bamboo or various grasses. A quick comparison with M. apatelius indicates that

they differ primarily in pileus color, paler and whitish in M. rotalis and more brownish in M. apatelius. ITS sequences

of the Madagascar specimens of M. rotalis (KX148999, KX149000, KX149001) align with GenBank sequences of M.

rotalis and M. rotula, in a clade with other members of sect. Marasmius subsect. Marasmius. (Fig. 1c).

2. Marasmius somalomoensis Antonín, Mycotaxon 88: 66. 2003. (Fig. 3, Plate 1)

Type:—CAMEROON. Sud Province, Somalomo, Dja Biosphere Reserve, 8 April 2001, V. Antonín Cm 01.42 (BRNM 666108).

Description:—Pileus 2–9 mm diam, plano-convex to campanulate, umbilicate, with a brown papilla; margin plicate to

sulcate; surface dry, glabrous; tan to pale brown (4–5A3). Context thin (<1 mm), white. Lamellae adnate to a collarium,

distant to subdistant (11–14), no lamellulae, broad, buff (4A2), non-marginate. Stipe 13–32 × <0.5 mm, central, narrow,

wiry, twisted, insititious; surface glabrous; dark brown (6F8). Odor and taste not distinctive.

FIGURE 3. Marasmius somalomoensis (JES 129, JES 165, JES 181) a) basidiospores; b) basidioles; c) basidia; d) cheilocystidia;

e) pileipellis cells. Scale bar = 10 µm. Illustrated by J.E. Shay.

Basidiospores (6.4–) 7.2–10.4 × 3.2–4.8 µm [xmr = 7.9–9.3 × 3.6–3.9 µm; xmm = 8.53 ± 0.70 × 3.79 ± 0.10; Q =

1.5–2.8; Qmr = 2.18–2.44; Qmm = 2.27 ± 0.10, n = 25, s = 3], ellipsoid to narrowly ellipsoid, smooth, hyaline, inamyloid,

thin-walled. Basidia 16.8–22.4 × 6.4–7.2 μm, cylindrical to subclavate, 4-spored, hyaline, inamyloid, thin-walled;

sterigmata 3.2–4.8 × 0.8 μm. Basidioles 18.4–24 × 5.6–8 μm, clavate to fusoid, hyaline, inamyloid, thin-walled.

Cheilocystidia few, of Rotalis-type broom cells; main body 7.2–28.8 × 6.4–21.6 μm, cylindrical to clavate, subglobose

or irregular, hyaline, inamyloid, thin-walled; divergent setulae 0.3–1.6 × 0.8 μm, dense, cylindrical, hyaline, inamyloid,

thin-walled. Pleurocystidia absent. Pileipellis mottled, a hymeniform layer of Rotalis-type broom cells; main body

12–31.2 × 8.8–24 μm, globose to subglobose, broadly clavate or irregular, hyaline, inamyloid, thin-walled; divergent

setulae 0.5–1.6 × 0.5–1.6 μm, dense, cylindrical, hyaline to brown, inamyloid, thin-walled. Pileus trama interwoven;

hyphae 1.6–12 μm diam, cylindrical to inflated, smooth, hyaline, inamyloid, thin-walled. Lamellar trama regular;

hyphae 1.6–14.4 μm diam, cylindrical to inflated, smooth, inamyloid, thin-walled. Stipe tissue monomitic; cortical

hyphae 2–8.8 μm diam, parallel, cylindrical, dark brown, dextrinoid, thick-walled; medullary hyphae 1.6–8 μm diam,

parallel, cylindrical to inflated, hyaline, inamyloid, thin-walled. Caulocystidia absent. Clamp connections present.

PLATE 1. Basidiocarps representing sect. Marasmius subsect. Marasmius a) Marasmius rotalis (JES 154) b) Marasmius apatelius (JES

203) c) Marasmius somalomoensis (JES 129). Scale bar = 5 mm (a); = 10 mm (b, c). Photography by D.S. Newman.

SHAY ET AL.

Habit, habitat and known distribution:—Solitary or in small gregarious clusters on Uapaca densifolia (dicot,

Phyllanthaceae). Africa (Cameroon, DR Congo), Madagascar.

Material examined:—MADAGASCAR. Commune Ranomafana, District Ifanadiana, Region Vatovavy-

Fitovinany, Ranomafana National Park, Talatakely Trail, elev.973 m, GPS: 21˚ 15.44’ S, 47˚ 25.116’ E, 20 January

2014, J.E. Shay 129 (SFSU); Ranomafana National Park, Piste B, elev.1004 m, GPS 21˚ 15.413’ S, 47˚ 25.253’ E, 22

January 2014, J.E. Shay 165 (SFSU); District Moramanga, Region Alaotra-Mangoro, Commune Andasibe, Vohimana

forest, Piste 5, elev.844 m, GPS 18˚ 55.422’ S, 48˚ 30.201’ E, 27 January 2014, J.E. Shay 181 (SFSU).

Notes:—Described originally from Cameroon, Marasmius somalomoensis is characterized by a tan to pale brown

pileus, subdistant (11–14), collariate, non-marginate lamellae, a brown stipe with cortical hyphae that do not turn olive

in KOH, basidiospores in the range 7.2–10.4 × 3.2–4.8 µm with mean 8.5 × 3.8 µm, Rotalis-type broom cells, and

growth on dicotyledonous leaves. It is nearly indistinguishable from M. colorimarginatus, which differs in forming a

darker brown pileus, greyish brown lamellae with brown margins, and stipe cortical hyphae that are olive in KOH.

ITS sequences of the Madagascar material of M. somalomoensis (KX149002, KX149003, KX149004) are more

than 3% different from the single available sequence of M. somalomoensis (EU935559) derived from a specimen from

Thailand (NW 232), but it should be noted that the Thai material is reported as forming pilei with more reddish brown

tones and more lamellae (12–18) with brown edges. It is possible that the Thai material represents a species different

from the African M. somalomoensis. (Fig. 1c).

3. Marasmius apatelius Singer, Bull. Jard. Bot. État Brux. 34: 332. 1964. (Fig. 4, Plate 1)

Type:—DR CONGO, Kisantu, 20 March 1907, H. Vanderyst s.n. (BR 11377–28, as M. friesianus).

Description:—Pileus 2–5 mm diam, campanulate to umbilicate, with a dark brown (5F4) papilla; margin sulcate to

plicate; surface dry, glabrous; greyish orange (5B4–5, 6A–C2) to buff, dries dark brown. Context thin, buff. Lamellae

adnate to a collarium, distant (9–12), broad (2–3 mm), white to orange-white (5A2), non-marginate. Stipe 18–54 ×

<0.5 mm, central, hollow, thin, wiry, insititious; surface glabrous, dark brown (6F8). Odor and taste not distinctive.

FIGURE 4. Marasmius apatelius (JES 150, JES 203) a) basidiospores; b) basidia; c) basidioles; d) cheilocystidia; e) pileipellis cells.

Scale bar = 10 µm. Illustrated by J.E. Shay.

Basidiospores (7.2–) 8–10.4 (–11.2) × 3.2–4.8 (–5) μm [xmr = 8.9–9.0 × 4.0–4.1 µm; xmm = 8.94 ± 0.02 × 4.02

± 0.03; Q = 1.8–2.8; Qmr = 2.23–2.25; Qmm = 2.24 ± 0.02, n = 25, s =2], ellipsoid, smooth, hyaline, inamyloid, thin-

walled. Basidia 23.2–24.8 × 6.4–8 μm, clavate, 4-spored, hyaline, inamyloid, thin-walled. Basidioles 12–22.4 × 4–8

μm, clavate to fusoid or cylindrical, hyaline, inamyloid, thin-walled. Cheilocystidia abundant, of Rotalis-type broom

cells; main body 7.6–23.2 × 8–14.4 μm, globose to subglobose or broadly clavate, hyaline, inamyloid, thin-walled;

divergent setulae 0.5–3.2 × 0.5–2 μm, cylindrical to conical, hyaline, inamyloid, thin-walled. Pleurocystidia absent.

Pileipellis not mottled, a hymeniform layer of Rotalis-type broom cells; main body 12–27.2 × 8–27.2 μm, globose to

broadly clavate, hyaline, inamyloid, thin-walled; divergent setulae 0.5–2 × 0.5–2 μm, numerous, cylindrical, hyaline

to brown, dextrinoid, thin-walled. Pileus trama interwoven; hyphae 1.6–12.8 μm diam, cylindrical to inflated, smooth,

hyaline, inamyloid, thin-walled. Lamellar trama regular; hyphae 1.6–8 µm diam, cylindrical to inflated, smooth,

hyaline, inamyloid, thin-walled. Stipe tissue monomitic; cortical hyphae 2.4–4 μm diam, parallel, cylindrical, yellow,

dextrinoid, thick-walled; medullary hyphae 3.2–6 μm diam, parallel, cylindrical, hyaline, inamyloid, thin-walled.

Caulocystidia absent. Clamp connections present.

Habit, habitat and known distribution:—In gregarious clusters on Uapaca littoralis (dicot, Phyllanthaceae). Africa

(DR Congo, Tanzania, Uganda), Madagascar, Thailand.

Material examined:—MADAGASCAR. Region Atsinanana, District Brickaville, Commune Andevoranto,

Andavakimena Forest, elev.-1 m, GPS: 18˚ 53.231’ S, 49˚ 07.490’ E, 28 January 2014, J.E. Shay 203 (SFSU); Region

Vatovavy-Fitovinany, District Ifanadiana, Commune Randomafana, Ranomafana National Park, Piste B, elev. 1004 m,

GPS: 21˚ 15.413’ S, 47˚ 25.253’ E, 22 January 2014, J.E. Shay 150 (SFSU).

Notes:—Marasmius apatelius is characterized by small (2–5 mm diam), greyish orange to buff pileus, distant

(9–12) collariate lamellae, a dark brown, wiry insititious stipe, basidiospores in the range 8–10.4 × 3.2–5 µm with

mean 8.9 × 4.0 µm, Rotalis-type broom cells, and growth on dicotyledonous leaves. Described originally from the DR

Congo, the Madagascar material matches nicely the African specimens reported by Antonín (2007) and Thai material

reported by Wannathes et al. (2009a). An ITS sequence of the holotype of M. andasibensis var. obscurostipitatus

Antonín & Buyck (KX149005) places the taxon in a clade with M. apatelius (Fig. 1c), and differs primarily in several

21–25 base pair insertions. For a comparison with numerous other members of Marasmius sect. Marasmius subsect.

Marasmius, refer to Antonín (2007).

4. Marasmius andasibensis var. andasibensis Antonín & Buyck, Fungal Diversity 23: 21. 2006

Type:—MADAGASCAR. Andasibe, 23 February 2000, B. Buyck 00.1704 (PC!).

For a description and illustrations of Madagascar material, refer to Antonín and Buyck (2006). Repeated attempts to

sequence material from collection Buyck 00.1704 (PC) were unsuccessful.

5. Marasmius andasibensis var. obscurostipitatus Antonín & Buyck, Fungal Diversity 23: 23. 2006

Type:—MADAGASCAR. Andasibe, 21 February 1997, B. Buyck 00.1699b (PC!).

For a description and illustrations of Madagascar material, refer to Antonín & Buyck (2006). The holotype collection

Buyck 00.1699b (PC) was sequenced (KX149005) and falls in a clade with M. apatelius but with low support.

Ib. Subsect. Sicciformes Antonín

Marasmius sect. Marasmius, subsect. Sicciformes Antonín, Acta Mus. Moraviae, Sci. Nat., 76: 145. 1991.

= subsect. Penicillati Singer sensu Singer, Fl. Neotrop. Monogr. 17: 121. 1976.

– Type species: Marasmius curreyi Berk. & Broome.

6. Marasmius nigrogriseus Antonín & Buyck, Fungal Diversity 23: 29. 2006

Type:—MADAGASCAR. Andasibe, 22 February 1997, B. Buyck 97.011 (PC!).

For a description and illustrations of Madagascar material, refer to Antonín & Buyck (2006). Repeated attempts to

sequence material from collection Buyck 97.011 (PC) were unsuccessful.

7. Marasmius nigrobrunneus (Pat.) Sacc., Syll. Fung. (Abellini) 11: 37. 1895

Type:—VIETNAM. Hanoi, Keso, 31 May 1890, Bon 4397 (FH).

For a description and illustrations of Madagascar material, refer to Antonín & Buyck (2006). Repeated attempts to

sequence material from collection Buyck 97.156 (PC) were unsuccessful.

SHAY ET AL.

8. Marasmius conicopapillatus Henn., Bot. Jb. 22: 100. 1895. (Fig. 5, Plate 2)

Type:—CAMEROON. Ekundu–Liongo, 20 May 1892, P. Dusén 41 (UPS).

Description:—Pileus 1–4 mm diam, convex to plano-convex, umbilicate, with prominent dark brown conical papilla;

margin sulcate; surface dull, dry, glabrous; white at margins becoming tan to cream towards disk. Context thin (<1 mm),

buff. Lamellae adnate to a collarium, distant (11–12), no lamellulae, ventricose, buff to cream (5A2), non-marginate.

Stipe 2–9 × <0.5 mm, central, wiry, pliant, hollow; surface glabrous; initially white darkening to light brown or brown

(6E8) at the base. Odor and taste not distinctive.

FIGURE 5. Marasmius conicopapillatus (JES 180) a) basidiospores; b) basidia; c) basidioles; d) cheilocystidia; e) pileipellis cells. Scale

bar = 10 µm. Illustrated by J.E. Shay.

Basidiospores (7.2–) 8–9.6 × (3.8–) 4–4.4 µm [xm = 8.50 ± 8.4 × 4.02 ± 0.12 µm; Q = 1.36–2.40; Qm = 2.11

± 0.18, n = 25, s = 1], ellipsoid, smooth, hyaline, inamyloid, thin-walled. Basidia 19.2–28 × 4.8–6.4 μm, clavate,

4-spored, hyaline, inamyloid, thin-walled; sterigmata 3.2–4.8 × 1.6 μm. Basidioles 12–29.6 × 3.3–5.6 μm, clavate,

hyaline, inamyloid, thin-walled. Cheilocystidia of Siccus-type broom cells; main body 9.6–16 × 7.2–8 μm, clavate to

subglobose or irregular, seldom lobed, hyaline, inamyloid, thin-walled; apical setulae 0.8–2.4 × 0.8 μm, cylindrical,

hyaline, inamyloid, thin-walled. Pleurocystidia absent. Pileipellis not mottled, a hymeniform layer of Siccus-type

broom cells; main body 7.2–20 × 4.8–10.4 μm, clavate to subglobose, seldom lobed, hyaline, inamyloid, thick-walled;

apical setulae 0.8–3.2 × 0.8 μm, cylindrical, hyaline, inamyloid, thick-walled. Pileus trama interwoven; hyphae

3.2–14.4 μm diam, cylindrical to inflated, smooth, hyaline, inamyloid, thin-walled. Lamellar trama regular; hyphae

2.4–6.4 μm diam, cylindrical, hyaline, inamyloid, thin-walled. Stipe tissue monomitic; cortical hyphae 3.2–8.8 μm

diam, cylindrical to slightly inflated, smooth, hyaline, dextrinoid, thick-walled; medullary hyphae 0.8–22.4 μm diam,

cylindrical to inflated, smooth, hyaline, inamyloid, thin-walled. Caulocystidia absent. Clamp connections present.

Habit, habitat and known distribution:—Gregarious, in clusters on leaves of Eugenia (dicot, Myrtaceae).

Africa (Burundi, DR Congo, Cameroon, Ghana, Ivory Coast, Nigeria, Uganda, Sierra Leone), Indonesia (Java),

Madagascar.

Material examined:—MADAGASCAR. Region Alaotra-Mangora, District Moramanga, Commune Andasibe,

Vohimana Forest, Piste 5, elev. 820–860 m, GPS: 18˚ 55.422’ S, 48˚ 30.201’ E, 26 January. 2014, J.E. Shay 180 (SFSU).

Notes:—Marasmius conicopapillatus is distinguished by small (1–4 mm diam), sulcate, umbilicate pileus with a

prominent dark brown conical papilla and initially white margin that develops tan to cream tones in age, distant (11–12)

non-marginate collariate lamellae, a short stipe initially white to cream and becoming brown in age, basidiospores with

mean 8.5 × 4 µm, Siccus-type cheilocystidia and pileipellis broom cells, and growth on dicotyledonous leaves. The

Madagascar specimen (JES 180) matches nicely the description of African material by Antonín (2007). Unfortunately,

repeated attempts to obtain an ITS sequence were unsuccessful.

PLATE 2. Basidiocarps representing sect. Marasmius subsect. Sicciformes a) Marasmius conicopapillatus (JES 180) b) Marasmius aff.

curreyi (JES 135) c) Marasmius rubrobrunneus (JES 183). Scale bar = 5 mm (a); = 10 mm (b, c). Photography by D.S. Newman.

9. Marasmius curreyi Berk. & Broome, Ann. Mag. Nat. Hist., Ser. 5, 3: 209. 1879

Type:—UNITED KINGDOM. Fineshade, 1859, M.J. Berkeley (K).

For a description and illustrations of Madagascar material, refer to Antonín & Buyck (2006). An ITS sequence from

the Madagascar collection Buyck 97.374 (PC) (KX148980) formed a weakly supported trichotomy with a Malaysian

sequence of M. curreyi (FJ431237) and M. graminum (JN943595) from Denmark.

SHAY ET AL.

10. Marasmius aff. curreyi (Fig. 6, Plate 2)

Description:—Pileus 2–7 mm diam, campanulate, umbilicate, with a dark brown papilla; margin plicate to sulcate;

surface dry, glabrous; tan to light brown with greyish tones (6D6) or cream (5A2) becoming paler at the margin.

Context thin, white. Lamellae adnate to a collarium, distant (10–11), no lamellulae, narrow (0.3–0.5), white to cream,

non-marginate. Stipe 5–30 × <0.5 mm, central, cylindrical, hollow, wiry, insititious; surface glabrous; dark brown.

Odor and taste not distinctive.

FIGURE 6. Marasmius aff. curreyi (JES 135) a) basidiospores; b) basidioles; c) pileipellis cells. Scale bar = 10 µm. Illustrated by J.E.

Shay.

Basidiospores (7.2–) 8–9.6 × 4–4.8 µm [xm = 9.02 ± 0.71 × 4.29 ± 0.37 µm; Q = 1.80–2.40; Qm = 2.11 ± 0.20, n

= 25, s = 1], ellipsoid, smooth, hyaline, inamyloid, thin-walled. Basidia 4–14.4 μm, clavate to cylindrical, 4-spored,

hyaline, inamyloid, thin-walled; sterigmata 0.8 × 0.2 µm. Basidioles 12–20 × 4.8–8 μm, clavate to cylindrical, hyaline,

inamyloid, thin-walled. Cheilocystidia not observed. Pleurocystidia absent. Pileipellis mottled, a hymeniform layer of

Siccus-type broom cells; main body 8–16 × 6.4–8.8 μm, clavate, hyaline, inamyloid, thin-walled; apical setulae 0.8–

2.4 × 0.8–1.6 μm, cylindrical to conical, brown to hyaline, inamyloid, thin-walled. Pileus trama interwoven; hyphae

2.4–8.8 μm diam, cylindrical to inflated, smooth, hyaline, inamyloid, thin-walled. Lamellar trama regular; hyphae

2.4–12 μm diam, cylindrical, smooth, hyaline, dextrinoid, thin-walled. Stipe tissue monomitic; cortical hyphae 2–8

μm diam, parallel, cylindrical, smooth, dark brown to brownish yellow, sometimes dextrinoid, thin-walled; medullary

hyphae 2–6 μm diam, parallel, cylindrical, smooth, hyaline, weakly dextrinoid, thin-walled. Caulocystidia absent.

Clamp connections present.

Habit, habitat, and known distribution:—Solitary to gregarious on stems of Justicia (Acanthaceae). Madagascar.

Material examined:—MADAGASCAR. Region Vatovavy-Fitovinany, District Ifanadiana, Commune Ranomafana,

Ranomafana National Park, Circuit Vohiparara, elev. 1062 m, GPS: 21˚ 14.255’ S, 47˚ 23.409’ E, 21 January 2014, J.E.

Shay 135 (SFSU).

Notes:—Distinctive features of Madagascar populations of Marasmius aff. curreyi are a rather small (2–7 mm

diam) tan to light brown pileus with yellowish brown tones that soon fades to tan, distant (10–11) collariate non-

marginate lamellae, a relatively short (<30 mm) stipe, basidiospores in the range 8–9.6 × 4–4.8 µm, and growth

on dead stems. An ITS sequence of material from Madagascar (KX148980) identified as M. curreyi by Antonín &

Buyck (2006) was only 96.3% similar to JES 135 (KX149008). The latter Madagascar sequence, however, was nearly

indistinguishable from material determined as M. curreyi from Korea (FJ936152) (Fig. 1c). Although the morphology

of the Madagascar material is similar to that of M. curreyi, this species was described originally from England and

no sequences of U.K. or European material determined as M. curreyi are available for comparison. Accordingly, we

recognize our material as M. aff. curreyi.

11. Marasmius curreyi var. bicystidiatus Antonín & Hauskn., Fungal Diversity 23: 26. 2006

Type:—MAURITIUS. Rivière du Rempart, Grand Baie, Bougain Villas, 4 March 1993, leg. Hausknecht MA04a (WU 14896).

For a description and illustrations of Madagascar material, refer to Antonín & Buyck (2006). Material from collection

A. Hausknecht MA04a (WU 14896) was unavailable for sequencing.

12. Marasmius rubrobrunneus J.E. Shay & Desjardin, sp. nov. (Fig. 7, Plate 2)

MycoBank no.: MB818617

Holotype:—MADAGASCAR. District Moramanga, Region Alaotra-Mangoro, Commune Andasibe, Vohimana Forest, Piste 5, elev. 844

m, GPS: 18˚ 55.422’ S, 48˚ 30.201’ E, 27 January 2014, J.E. Shay 191 (SFSU).

Etymology:—rubro–reddish, -brunneus–brown, referring to the dark reddish brown pileus.

Description:—Pileus 4–9 mm diam, campanulate, umbilicate, with a dark brown papilla; margin sulcate; surface

glabrous; dark reddish brown (7–8E–F5–8). Context thin, dark reddish brown. Lamellae adnate to a collarium, distant

(11–12), broad, no lamellulae; white to buff (4A2) with reddish brown edges. Stipe 16–26 × <0.5 mm, central, hollow,

twisted, wiry, insititious; surface glabrous; dark brown. Odor and taste not distinctive.

FIGURE 7. Marasmius rubrobrunneus (JES 183, JES 191) a) basidiospores; b) basidioles; c) cheilocystidia; d) pileipellis cells. Scale bar

= 10 µm. Illustrated by J.E. Shay.

Basidiospores (12–) 13.6–19.2 × 2.4–4 µm [xmr = 14.9–16.5 × 3.2 µm; xmm = 15.71 ± 1.13 × 3.2 ± 0.06; Q = 2.8–6.7;

Qmr = 4.6–5.18; Qmm = 4.89 ± 0.41, n = 25, s = 2], narrowly fusoid, smooth, hyaline, inamyloid, thin-walled. Basidia

not observed. Basidioles 17.6–26.4 × 4.8–7.2 μm, clavate to fusoid, hyaline, inamyloid, thin-walled. Cheilocystidia

abundant, of Siccus-type broom cells; main body 12–20 × 6.4–8 μm, clavate to cylindrical or subglobose, seldom

bilobed, light brown, inamyloid, thick-walled; apical setulae 0.8–4.8 × 0.8–2.4 µm, cylindrical to conical, sometimes

branched, light brown, inamyloid, thick-walled. Pleurocystidia absent. Pileipellis mottled, a hymeniform layer of

Siccus-type broom cells; main body 9.6–19.2 × 6.4–12 μm, clavate to cylindrical or subglobose, brown to hyaline,

inamyloid, thick-walled; apical setulae 0.8–6.4 × 0.8–2.4 µm, cylindrical to conical, obtuse, seldom branched, brown

to hyaline, inamyloid, thick-walled. Pileus trama interwoven; hyphae 3.2–8.8 μm diam, smooth, hyaline, inamyloid,

thin-walled. Lamellar trama regular; hyphae 2.4–16 μm diam, cylindrical, smooth, hyaline, inamyloid, thin-walled.

Stipe tissue monomitic; cortical hyphae 2.4–8 μm diam, parallel, cylindrical, smooth, light brown to brown, dextrinoid,

thick-walled; medullary hyphae 4–10.4 μm diam, parallel, cylindrical, smooth, light yellow to hyaline, inamyloid,

thin-walled. Caulocystidia absent. Clamp connections present.

Habit, habitat and known distribution:—Solitary or in small gregarious clusters on a variety of leaf litter and stems

of Uapaca densifolia (Phyllanthaceae), Canarium boivinii (Burseraceae), Pandanus, Contium and other unknown

dicots. Madagascar.

Material examined:—MADAGASCAR. District Moramanga, Region Alaotra-Mangoro, Commune Andasibe,

Vohimana Forest, Piste 5, elev. 844 m, GPS: 18˚ 55.422’ S, 48˚ 30.201’ E, 27 January 2014, J.E. Shay 191 (SFSU) &

J.E. Shay 183 (SFSU).

Notes:—Features of Marasmius rubrobrunneus include a small (4–9 mm diam), sulcate, dark reddish brown

pileus, distant collariate lamellae with reddish brown edges, a black wiry insititious stipe, basidiospores with mean

15.7 × 3.2 µm (Qm = 4.9), Siccus-type broom cells, and growth on dicotyledonous leaves and twigs. Morphologically,

SHAY ET AL.

the new species is nearly indistinguishable from M. purpureobrunneolus Henn., described originally from Java. The

latter species has been redescribed several times (Desjardin et al. 2000, Tan et al. 2009, Wannathes et al. 2009a)

from material collected in southeast Asia, and is distinguished by a dark purplish brown pileus and basidiospores in

the range 12–17 (–19) × 2.5–5 µm, with means in the range 14.0–14.8 × 3.2–4.1µm (Qmr = 3.6–4.5). In comparison,

Marasmius rubrobrunneus has a dark reddish brown pileus lacking purple tones, and basidiospores with means in the

range 14.9–16.5 × 3.2 µm, i.e., slightly longer and narrower than in M. purpureobrunneolus. Although this variability

may seem trivial, a comparison of the ITS sequence of the Madagascar holotype specimen (KX148989) with two

specimens of M. purpureobrunneolus from Thailand (EU935556, EU935557) show only 85% similarity. In the ITS

phylogenetic analysis (Fig. 1c), M. rubrobrunneus is sister to M. purpureobrunneolus with good support (81% BS; 1.0

PP).

13. Marasmius brunneoaurantiacus Antonín & Buyck, Fungal Diversity 23: 24. 2006. (Fig. 8, Plate 3)

Type:—MADAGASCAR. Ranomafana National Park, 4 February 1999, leg. B. Buyck & G. Eyssartier, Buyck 99.450 (PC!).

Description:—Pileus 5–20 mm diam, campanulate to hemispherical, umbilicate, with a dark brown papilla; margin

sulcate to plicate; surface dry, glabrous; reddish brown (7C–E7–8, 8D8, 6D6). Context thin (<1 mm), white to cream

(4A3). Lamellae adnate to a collarium, subdistant (12–16), broad (0.5–1mm), cream, non-marginate or with brown

edges. Stipe 16–67 × 0.2–0.4 mm, central, cylindrical, wiry, pliant, insititious; surface glabrous, light brown at apex to

dark brown (7E6) towards the base. Odor and taste not distinctive.

FIGURE 8. Marasmius brunneoaurantiacus (JES 113, JES 137, JES 166, JES 218) a) basidiospores; b) basidia; c) basidioles; d)

cheilocystidia; e) pileipellis cells. Scale bar = 10 µm. Illustrated by J.E. Shay.

Basidiospores (6.2–) 8–10.4 (–11.2) × (3.2–) 4–4.8 (–5.6) μm [xmr = 8.8–9.5 × 4.1–4.6 µm; xmm = 9.03 ± 0.3

× 4.28 ± 0.2; Q = 1.3–3.0; Qmr = 2.0–2.3; Qmm = 2.14 ± 0.1, n = 25, s = 6], ellipsoid, smooth, hyaline, inamyloid,

thin-walled. Basidia 16.8–27.2 × 5.6–6.4 μm, clavate, 4-spored, hyaline, inamyloid, thin-walled; sterigmata 3.2–4

× 1.6 µm. Basidioles 14.4–30.4 × 3.2–8 μm, clavate to fusoid, hyaline, inamyloid, thin-walled. Cheilocystidia few

to abundant, of Siccus-type broom cells; main body 12–29.6 × 4.8–14.4 μm, cylindrical to clavate, subglobose or

irregular, often 2–3-lobed, hyaline, inamyloid, thin-walled; apical setulae dense, 0.8–5.6 × 0.8–2.4 μm, cylindrical to

conical, hyaline, inamyloid, thick-walled. Pleurocystidia absent. Pileipellis mottled, a hymeniform layer of Siccus-

type broom cells; main body 8.8–20 × 6.4–8.8 μm, clavate to subglobose or irregular, seldom 2–4-lobed, hyaline,

inamyloid, thin-walled; apical setulae 0.8–7.2 × 0.8–2.4 μm, cylindrical to conical, seldom branched, hyaline to dark

brown, inamyloid, thick-walled. Pileus trama interwoven; hyphae 1.6–16 μm diam, cylindrical to inflated, smooth,

hyaline, inamyloid, thin-walled. Lamellar trama regular; hyphae 2.4–14.4 μm diam, cylindrical to inflated, smooth,

hyaline, inamyloid, thin-walled. Stipe tissue monomitic; cortical hyphae 3.2–10.4 μm diam, parallel, cylindrical to

inflated, smooth, brown, dextrinoid, thick-walled; medullary hyphae 3.2–27 μm diam, parallel, cylindrical to inflated,

smooth, hyaline, dextrinoid, thick-walled. Caulocystidia absent. Clamp connections present.

Habit, habitat and known distribution:—Solitary or gregarious on leaves and stems of Uapaca (Phyllanthaceae),

Cryptocarya (Lauraceae), Maesa (Primulaceae), Volina madagascariensis (bamboo) and Cephalostachium vigueri

(bamboo). Madagascar.

Material examined:—MADAGASCAR. Commune Ranomafana, District Ifanadiana, Region Vatovavy-Fitovinany,

Ranomafana National Park, Talatakely Trail, elev. 937–973 m, GPS: 21˚ 15.237’ S, 47˚ 25.183’ E, 20 January 2014,

J.E. Shay 113 (SFSU), J.E. Shay 115 (SFSU), J.E. Shay 125 (SFSU) & J.E. Shay 131 (SFSU); Ranomafana National

Park, Circuit Vohiparara, elev. 1062 m, GPS: 21˚ 14.255’ S, 47˚ 23.409’ E, 21 January 2014, J.E. Shay 133 (SFSU) & J.

E. Shay 137 (SFSU); Ranomafana National Park, Piste B, elev. 1004 m, GPS: 21˚ 15.413’ S, 47˚ 25.253’ E, 22 January

2014, J.E. Shay 166 (TAN); Region Analamanga, City of Antananarivo, Parc Botanique de Zoologique (P.B.Z.T.), near

the garden of Crops Wild and Relatives (CWR), elev. 1270 m, GPS: 18˚ 55.53’ S, 47˚ 31.35’ E, J.E. Shay 218 (SFSU);

Ranomafana National Park, along crest near roadside just outside Park area, 4 February 1999, leg. B. Buyck & G.

Eyssartier, Buyck 99.450 (PC, holotype).

Notes:—Marasmius brunneoaurantiacus is characterized by a rather large (5–20 mm diam), light brown to brown

or reddish brown pileus, subdistant (12–16), collariate, non-marginate or brown-marginate lamellae, a relatively long

stipe (up to 67 mm), basidiospores in the range 8–10.4 (–11) × 4–4.8 µm, Siccus-type broom cells, and growth mainly

on bamboo leaves, occasionally on dicot leaves.

Antonín & Buyck (2006) described the species as having brown-marginate lamellae, basidiospores in the range

9.5–11 × 4.5–6 µm, and growth on dead leaves of Uapaca ferruginea (Buyck 99.439). The holotype specimen (Buyck

99.450), however, is undoubtedly growing on bamboo leaves. Our material of M. brunneoaurantiacus occurs mainly on

bamboo leaves, (although several specimens are on both grasses and dicot leaves (JES 115, JES 133)), has basidiomes

with brown-marginate or non-marginate lamellae, and the basidiospores are in the shorter end of the range. In other

features, our material is indistinguishable from the holotype. An ITS sequence of the holotype specimen (KX148978)

shows 99.1–99.6% similarity to seven additional specimens from Madagascar (KX149009–KX149014, KX149016),

forming a clade with 100% BS and 1.0 PP support (Fig. 1c).

14. Marasmius crinisequi F. Muell. ex Kalchbr., in Kalchbrenner, Grevillea 8(48): 153. 1880. (Fig. 9, Plate 3)

= Marasmius equicrinis F. Muell. ex Berk., J. Linn. Soc. Bot. 18: 383. 1881.

= Androsaceus crinisequi (F. Muell. ex Kalchbr.) Overeem, De nuttige planten van Nederlandsch Indië 1: 69. 1927.

= Marasmius graminum var. equicrinis (F. Muell. ex Berk.) Dennis, Trans. Brit. Mycol. Soc. 34: 416. 1951.

= Marasmius repens Henn., Bot. Jb. 23: 548. 1897 (nom. illeg., non Marasmius repens (Bull.) Quél. 1886).

= Marasmius ramentaceus (Pat.) Sacc. & Traverso, Syll. Fung. (Abellini) 20: 21. 1911.

= Androsaceus ramentaceus Pat., Ann. Jard. Bot. Buitenzorg, Suppl. 1: 107. 1897.

Type:—AUSTRALIA. North Queensland, Rockingham Bay, F. von Mueller s.n. (K(M) 99658, lectotype).

Description:—Pileus 1–2 mm diam, convex to campanulate, umbilicate, with a dark brown papilla; margin plicate to

sulcate; surface dull, dry, glabrous; light brownish orange (5B5–6). Context thin, buff. Lamellae adnate to a collarium,

distant (6), no lamellulae, broad, buff (5A3), non-marginate. Stipe 2–4 × <0.5 mm, central, wiry, pliant, arising directly

from coarse black rhizomorphs; surface glabrous; dark brown. Odor and taste not distinctive.

Basidiospores (8.8–) 9.6–13.6 × 4–5.6 μm [xm = 10.14 ± 1.24 × 4.54 ± 0.50 µm; Q = 1.57–3.40; Qm = 2.27 ±

0.18, n = 25, s = 1], ellipsoid, smooth, hyaline, inamyloid, thin-walled. Basidia 18.4–28 × 8.8–9.6 μm, clavate to

broadly clavate, 4-spored, hyaline, inamyloid, thin-walled; sterigmata 4–4.8 × 1.6 μm. Basidioles 16.8–24 × 5.6–7.2

μm, clavate to fusoid, hyaline, inamyloid, thin-walled. Cheilocystidia of Siccus-type broom cells; main body 8.8–15.2

× 6.4–10.4 μm, clavate to broadly clavate, seldom 2–3-lobed, hyaline, inamyloid, thin-walled; apical setulae 0.8–4.8

× 0.8–1.6 µm, cylindrical, obtuse, sometimes branched, hyaline, inamyloid. Pleurocystidia absent. Pileipellis not

mottled, a hymeniform layer of Siccus-type broom cells; main body 10.4–16 × 6.4–10.4 μm, cylindrical to clavate or

broadly clavate, seldom 2–3-lobed, hyaline, inamyloid, thin-walled; apical setulae 0.8–5.6 × 0.8–1.6 μm, cylindrical,

seldom branched, pale brown, inamyloid, thin-walled. Pileus trama interwoven; hyphae 2.4–6.4 μm diam, cylindrical,

smooth, hyaline, inamyloid, thin-walled. Lamellar trama regular; hyphae 1.6–7.2 μm diam, cylindrical, smooth, hyaline,

SHAY ET AL.

inamyloid, thin-walled. Stipe tissue monomitic; cortical hyphae 3.2–4.8 μm diam, cylindrical, smooth, dark brown,

inamyloid, thick-walled; medullary hyphae 2.4–9.6 μm diam, cylindrical to inflated, smooth, hyaline, dextrinoid, thin-

walled. Caulocystidia absent. Clamp connections present.

FIGURE 9. Marasmius crinisequi (JES 176) a) basidiospores; b) basidia; c) basidioles; d) cheilocystidia; e) pileipellis cells. Scale bar =

10 µm. Illustrated by J.E. Shay.

Habit, habitat and known distribution:—Solitary arising directly from rhizomorphs, attached to debris of Uapaca

densifolia, Anthocleista madagascarensis, Omphalea oppositefolia, and Noronhia. Pantropical, common in Africa

(Burundi, Cameroon, DR Congo, Ghana, Ivory Coast, Kenya, Nigeria, Sierra Leone), Asia, Australia, Caribbean

region, Madagascar.

Material examined:—MADAGASCAR. Region Alaotra-Mangora, District Moramanga, Commune Andasibe,

Vohimana Forest, Piste 2, elev. 820–860 m, GPS: 18˚ 55.422’ S, 48˚ 30.201’ E, 26 January 2014, J.E. Shay 176

(SFSU).

Notes:—Marasmius crinisequi, commonly called the horsehair fungus, forms basidiomes that arise directly

from coarse black rhizomorphs. The species is often arboreal, with the rhizomorphs forming a net-like structure that

captures falling leaves. The orange to light brown pileus is less than 2 mm diam, sulcate, with a small dark papilla in

the umbilicus, distant (6) lamellae, a short (2–4 mm), dark brown stipe, basidiospores with mean 10.1 × 4.5 µm, and

Siccus-type cheilocystidia and pileipellis broom cells. It represents a pantropical species or complex of species in need

of more phylogenetic analyses with multiple genes. Repeated attempts to obtain ITS sequences from the Madagascar

material were unsuccessful.

15. Marasmius cf. subruforotula Singer, Bull. Jard. Bot. État Brux. 34: 339. 1964. (Fig. 10, Plate 3)

Type:—DR CONGO. Equateur Province, Eala, July 1907, L. Pynaert 1608 (BR 11515–69).

Description:—Pileus 2–7 mm diam, campanulate to convex, umbilicate, with a dark brown (6F8) conical papilla;

margin plicate to sulcate; surface dry, glabrous; greyish orange to brownish orange or light brown (5–6B–D4–8).

Context thin, white. Lamellae adnate to a collarium, distant (9–12), no lamellulae, broad, white to buff (4A2), non-

marginate. Stipe 6–40 × <0.5 mm, central, cylindrical, wiry, hollow, insititious; surface glabrous; dark brown to black.

Odor and taste not distinctive.

Basidiospores 7.2–9.6 (–10.4) × 3.2–4 µm [xmr = 8–8.4 × 3.8–3.9 µm; xmm = 8.23 ± 0.2 × 3.87 ± 0.1 µm; Q = 1.8–

2.6; Qmr = 2.06–2.19; Qmm = 2.14 ± 0.1, n = 24–25, s = 3], ellipsoid, smooth, hyaline, inamyloid, thin-walled. Basidia

19.2–32 × 4–8 μm, clavate 4-spored, hyaline, inamyloid, thin-walled; sterigmata 1.6–4.8 × 0.8–1.6 μm. Basidioles

16.8–26.4 × 4–7.2 μm, clavate, hyaline, inamyloid, thin-walled. Cheilocystidia numerous, of Siccus-type broom cells;

main body 8.8–18.4 × 5.6–10.4 μm, clavate to cylindrical or subglobose, hyaline, inamyloid, thin-walled; apical

setulae 0.5–5.6 × 0.8–1.6 μm, conical to cylindrical, hyaline, inamyloid, thin-walled. Pleurocystidia absent. Pileipellis

mottled, a hymeniform layer of Siccus-type broom cells; main body 11.2–20 × 5.6–12.8 μm, clavate to cylindrical,

subglobose or irregular, hyaline, inamyloid, thick-walled; apical setulae 0.8–5.6 × 0.8–1.6 μm, cylindrical to conical,

sometimes branching, hyaline to yellow or brown, inamyloid, thin-walled. Pileus trama interwoven; hyphae 1.6–11.2

µm diam, cylindrical to inflated, smooth, hyaline, inamyloid, thin-walled. Lamellar trama regular; hyphae 1.6–9.6

µm diam, cylindrical to inflated, smooth, hyaline, inamyloid, thin-walled. Stipe tissue monomitic; cortical hyphae

3–5.6 µm diam, parallel, cylindrical, smooth, light brown to brown, dextrinoid, thick-walled; medullary hyphae 1.6–

9.6 µm diam, parallel, cylindrical to inflated, smooth, hyaline, inamyloid, thin-walled. Caulocystidia absent. Clamp

connections present.

PLATE 3. Basidiocarps representing sect. Marasmius subsect. Sicciformes a) Marasmius brunneoaurantiacus (JES 113) b) Marasmius

crinisequi (JES 176) c) Marasmius cf. subruforotula (JES 186). Scale bar = 10 mm. Photography by D.S. Newman.

Habit, habitat and known distribution:—Solitary or gregarious on a variety of monocotyledonous and

dicotyledonous leaves including, but not limited to Mallotus (Euphorbiaceae), Pandanus (Pandanaceae), Vernonia

(Asteraceae), Noronhia (Oleaceae), Blotia (Euphorbiaceae), Coffea mangoroensis (Rubiaceae), Alafia (Apocynaceae),

Uapaca thouarai, Uapaca densifolia (Phyllanthaceae), Ambavia (Annonaceae), Psychotria (Rubiaceae) and Carallia

brachiata (Rhizophoraceae). Africa (Cameroon, DR Congo, Nigeria, Tanzania, Uganda), Madagascar, Thailand.

SHAY ET AL.

FIGURE 10. Marasmius cf. subruforotula (JES 186, JES 190, JES 192) a) basidiospores; b) basidia; c) basidioles; d) cheilocystidia;

d) pileipellis cells. Scale bar = 10 µm. Illustrated by J.E. Shay.

Material examined:—MADAGASCAR. District Moramanga, Region Alaotra-Mangoro, Commune Andasibe,

Vohimana forest, Piste 5, elev. 844 m, GPS: 18˚ 55.422’ S, 48˚ 30.201’ E, 27 January 2014, J.E. Shay 186 (SFSU), J.E.

Shay 190 (SFSU) & J.E. Shay 192 (SFSU).

Notes:—The material from Madagascar shows closest phenetic similarity to Marasmius subruforotula. The

Madagascar taxon is distinguished by a small (2–7 mm diam), brownish orange pileus with a distinct dark brown

papilla, distant (9–12), collariate, non-marginate lamellae, dark brown insititious stipe, basidiospores in the range 7.2–

10.4 × 3.2–4 µm, Siccus-type broom cells, and growth on dicotyledonous leaves and twigs. Marasmius subruforotula,

described from the DR Congo, has reddish brown pilei, reddish brown-marginate lamellae and broader basidiospores

(4–5 µm wide). Our material matches that reported from Madagascar by Antonín & Buyck (2006) as M. cf. subruforotula.

Pegler (1977) and Antonin (2007) report M. subruforotula from throughout Africa with morphology that overlaps that

reported here; however, ITS sequences of material from Príncipe (Grace et al. unpubl.) are quite different from those

reported here from Madagascar specimens, although they are sister to each other (100% BS, 1.0 PP support; Fig.

1c), suggesting that they represent different species. Until additional materials representing a wide distribution of

specimens from the African continent are compared, we tentatively recognize the Madagascar taxon as Marasmius cf.

subruforotula.

16. Marasmius madagascariensis J.E. Shay & Desjardin, sp. nov. (Fig. 11, Plate 4)

MycoBank no.: MB 818620

Holotype:—MADAGASCAR. Region Vatovavy-Fitovinany, District Ifanadiana, Commune Ranomafana, Ranomafana National Park,

Circuit Vohiparara, elev. 1062 m, GPS: 21˚ 14.255’ S, 47˚ 23.409’ E, 21 January 2014, J.E. Shay 139 (SFSU).

Etymology:—madagascar-iensis–occurring in Madagascar.

Description:—Pileus 2–6 mm diam, convex to campanulate, umbilicate, with a reddish brown papilla; margin sulcate;

surface dry, glabrous; orangish brown (6C–D7, 5B6–7). Context thin (<1 mm), concolorous with pileus. Lamellae

adnate to a collarium, subdistant (9–11), broad (0.5–1 mm), light orange (5A4) to cream, non-marginate. Stipe 10–

23 × 0.1–1 mm, central, hollow, wiry, insititious; surface glabrous; black; rhizomorphs present. Odor and taste not

distinctive.

Basidiospores 8.8–12.8 × 4–5.6 (–7.2) µm [xmr = 11.3–11.6 × 4.8–5.0 µm; xmm = 11.46 ± 0.17 × 4.90 ± 0.09

µm; Q = 1.8–3.2; Qmr = 2.36–2.38; Qmm = 2.37 ± 0.02, n = 21–22, s = 2], ellipsoid, smooth, hyaline, inamyloid, thin-

walled. Basidia 18.4–26.4 × 9.6–11.2 µm, clavate, 4-spored, hyaline, inamyloid. Basidioles 13.6–24.8 × 4.8–8.5 µm,

clavate to cylindrical, hyaline, inamyloid, thin-walled. Cheilocystidia of Siccus-type broom cells; main body 8–19.2

× 6.4–11.2 µm, clavate to cylindrical, seldom lobed, hyaline, inamyloid, thin-walled; apical setulae 0.8–6.4 × 0.5–2

µm, cylindrical to conical, often branched, hyaline, inamyloid, thin-walled. Pleurocystidia absent. Pileipellis mottled,

a hymeniform layer of Siccus-type broom cells; main body 8–16 × 6.4–16 µm, clavate or irregular, 2–3-lobed, hyaline,

inamyloid, thin-walled; apical setulae 1.6–5.6 × 0.5–2 µm, cylindrical to conical, sometimes branched, yellowish

brown to hyaline, inamyloid, thin-walled. Pileus trama interwoven; hyphae 2.4–4.8 µm diam, cylindrical, smooth,

hyaline, inamyloid, thin-walled. Lamellar trama regular; hyphae 1.6–12 µm diam, cylindrical to inflated, smooth,

hyaline, inamyloid, thin-walled. Stipe tissue monomitic; cortical hyphae 2–4 µm diam, parallel, cylindrical, smooth,

brown, dextrinoid, thick-walled; medullary hyphae 2.4–8 µm diam, parallel, cylindrical, smooth, hyaline, weakly

dextrinoid, thin-walled. Caulocystidia absent. Clamp connections present.

FIGURE 11. Marasmius madagascariensis (JES 139, JES 225) a) basidiospores; b) basidia; c) basidioles; d) cheilocystidia; e) pileipellis

cells. Scale bar = 10 µm. Illustrated by J.E. Shay.

Habit, habitat and known distribution:—Solitary or gregarious on stems of Cyathea (tree fern, Cyatheaceae) and

on debris of an unknown grass (Poaceae). Madagascar.

Material examined:—MADAGASCAR. Region Vatovavy-Fitovinany, District Ifanadiana, Commune Ranomafana,

Ranomafana National Park, Circuit Vohiparara, elev. 1062 m, GPS: 21˚ 14.255’ S, 47˚ 23.409’ E, 21 January 2014,

J.E. Shay 139 (SFSU); Region Analamanga, District Ankazobe, Commune Ambatoharanama, Ambohitantely Forest

Reserve Sentier Botanique, 1574 m, GPS: 18˚ 11.504’ S, 47˚ 17.074’ E, 8 Feb. 2014, J.E. Shay 225 (SFSU).

Notes:—Marasmius madagascariensis is characterized by small (2–6 mm diam), orangish brown pileus with a

reddish brown central papilla, distant (9–11), collariate, non-marginate lamellae, a wiry stipe <23 mm long, abundant

rhizomorphs, basidiospores with mean 11.5 × 4.9 µm, Siccus-type broom cells, and growth on grass leaves and tree

fern stems. Morphologically it is similar to M. guyanensis Mont., a species described originally from French Guyana

(South America), and subsequently reported from the Caribbean region, Indonesia, Singapore, Malaysia, Thailand and

Africa. Morphologically, the Madagascar specimens are most similar to Thailand M. guyanensis where basidiomes are

formed on dicot leaves and have basidiospores with mean width 3.8 µm. In comparison, the Madagascar specimens

differ in growing on grass leaves and tree fern stems and have basidiospores with mean width 4.9 µm. ITS sequences

of Madagascar material of M. madagascariensis (KX149015, KX149006) are on a long branch embedded in a clade

with two Malaysian specimens determined as M. guyanensis (FJ431246, FJ431247; Tan et al. 2009), two specimens of

M. crinisequi, and two Thai specimens of M. guyanensis (EU935552, EU935553; Wannathes et al. 2009a). In addition,

the Madagascar specimens are morphologically similar to M. aff. guyanensis reported from the island of Princípe

(DED 8285, Grace et al., unpubl.), but the latter has longer and narrower basidiospores (12.5–15 × 3.5–4.5 µm), and

SHAY ET AL.

an insertion of 38 base pairs (between 593–630) in the ITS region. Because of its distribution in Madagascar, subtle

differences in basidiospore size and substrate, and ITS sequence differences, we recognize the Madagascar material as

a distinct species.

II. Sect. Neosessiles Singer

Marasmius sect. Neosessiles Singer, Mycologia 50: 104 (1958).

– Type species: Marasmius neosessilis Singer

17. Marasmius neosessiliformis Antonín & Buyck nom. prov., Fungal Diversity 23: 34. 2006

For a description and illustrations of Madagascar material of this provisionally described species, refer to Antonín &

Buyck (2006). An ITS sequence of collection Buyck 97.615 (KX149007) formed an unresolved clade with sequences

of M. tenuissimus and M. leveilleanus.

18. Marasmius cecropiformis Antonín & Hauskn., Fungal Diversity 23: 33. 2006

Type:—FRANCE. La Réunion, Forêt de Belouve, ca. 140 m elev., 13 March 1996, leg. A. Hausknecht RE 59/96 and G. Wölfel (WU

25700).

For a description and illustrations of Madagascar material, refer to Antonín & Buyck (2006). Material from collection

A. Hausknecht RE 59/96 and G. Wölfel (WU 25700) was unavailable for sequencing.

III. Sect. Globulares Kühner

Marasmius sect. Globulares Kühner, Botaniste 25: 100. 1933 (ut Globularinae).

– Type species: Marasmius globularis (Weinm.) Fr. (= M. wynneae Berk. & Broome)

IIIa. ser. Globulares

19. Marasmius sulcatipes Pat., Bull. Mus. Nat. Hist. Natur. 13: 526. 1924

Type:—MADAGASCAR. Massif de l’Ankaizniana, on old stump of tree in a humid mountain forest at 1500 m alt., leg. M. Decary

(PC).

For a descriptions and illustrations of Madagascar material, refer to Antonín & Buyck (2006). Material from collection

M. Decary (PC) was unavailable for sequencing. It should be noted that M. sulcatipes Pat. (1924) is an illegitimate

name, a later homonym of Marasmius sulcatipes Murrill [N.Amer. Fl. (New York) 9(4): 259. 1915], a species described

from Cuba now recognized as belonging to the genus Gymnopus.

20. Marasmius bekolacongoli Beeli, Bull. Soc. R. Bot. Belg. 60(2): 157. 1928. (Fig. 12, Plate 4)

Type:—DR CONGO. Equateur Province, Eala, October 1923, M. Goossens–Fontana 204 (BR 11406–57).

Description:—Macromorphological features derived from a photograph and dried material. Pileus 85 mm diam,

obtusely conical; disc rugulose; margin sulcate; surface dry, glabrous; striped, disc brown to violaceous brown,

sulcae pinkish-violaceous brown, ridges pale cream to buff. Context thin. Lamellae adnexed, distant (15), cream, non-

marginate. Stipe about 120 × 10 mm, central, cylindrical; surface longitudinally ridged; cream to tan or pale brown.

Odor and taste not distinctive.

Basidiospores (21.6–) 24–29.4 × (3.2–) 4–6.4 μm [x m = 26.05 ± 2.19 × 5.17 ± 0.98 µm; Q = 3.5–7.5; Qm = 5.24 ±

1.41, n = 25, s = 1], clavate, smooth, hyaline, inamyloid, thin-walled. Basidia not observed. Basidioles 35–42.4 × 8–11.2

μm, clavate, hyaline, inamyloid, thin-walled. Cheilocystidia few, 18.4–26.4 × 8–12 μm, clavate to broadly clavate or

cylindrical, smooth, hyaline, inamyloid, thin-walled. Pleurocystidia absent. Pileipellis not mottled, a hymeniform layer

of Globulares-type cells; main body 14.4–28 × 9.6–16 μm, clavate to broadly clavate, pyriform or subglobose, smooth,

hyaline, inamyloid, thin-walled. Pileus trama interwoven; hyphae 3.2–16 μm diam, cylindrical to inflated, smooth,

hyaline to pale light brown, dextrinoid, thin-walled. Lamellar trama regular; hyphae 3.2–16 μm diam, cylindrical to

inflated, smooth, hyaline, dextrinoid, thin-walled. Stipe tissue monomitic; cortical hyphae 3.2–8.8 μm diam, parallel,

cylindrical, yellowish brown, dextrinoid, thick-walled; medullary hyphae 2.4–10.4 μm diam, parallel, cylindrical, pale

yellowish, inamyloid, thick-walled. Caulocystidia absent. Clamp connections present.

FIGURE 12. Marasmius bekolacongoli (Lockwood 2131638) a) basidiospores; b) basidioles; c) cheilocystidia; d) pileipellis cells. Scale

bar = 10 µm. Illustrated by J.E. Shay.

PLATE 4. Basidiocarps representing sect. Marasmius subsect. Sicciformes. a) Marasmius madagascariensis (JES 225); sect. Globulares

b) Marasmius bekolacongoli (Lockwood 2131638), photo generously donated by Taylor Lockwood. Scale bar = 10 mm (a); 20 mm (b).

Photography by D.S. Newman.

SHAY ET AL.

Habit, habitat and known distribution:—Solitary on undetermined dicotyledonous debris. Africa (Burundi,

Cameroon, DR Congo, Kenya, Malawi, Nigeria, Tanzania, Uganda, Zimbabwe), Madagascar.

Material examined:—MADAGASCAR. Region Vatovavy-Fitovinany, District Ifanadiana, Commune Ranomafana,

February 2013, T. Lockwood 2131638 (SFSU).

Notes:—The single specimen from Madagascar (Lockwood 2131638) matches nicely the descriptions of African

material (Singer 1965, Pegler 1977, Antonín 2007), although the basidiospores are slightly longer. Our specimen has

a violaceous brown and cream striped pileus 85 mm diam, a large (120 × 10 mm) cream longitudinally ridged stipe,

basidiospores in the range 24–29.6 × 4–6.4 µm, no pleurocystidia, clavate cheilocystidia, Globulares-type pileipellis

cells, no caulocystidia, and growth in leafy debris. African M. bekolacongoli are reported to have basidiospores 17.5–

26 × 3.8–5.4 µm (Antonín 2007), otherwise the morphology is indistinguishable from Lockwood 2131638. Because

of the basidiospore size, the Madagascar specimen would key to M. camerunensis Antonín & Mossebo, but the latter

has a smaller, non-striped pileus lacking violaceous tones, and a smaller stipe (40–70 × 4–6 mm) and grows on

woody debris (Antonín 2007). Only a single basidiome of M. bekolacongoli was collected and photographed by Taylor

Lockwood. An ITS sequence of this specimen (KX148982) formed a weakly supported clade with other members of

sect. Globulares plus M. coarctatus (sect. Sicci, ser. Spinulosi) (Fig. 1b).

The following four series were formally described within Sect. Sicci Singer, but because they represent non-

monophyletic lineages (see Results above) they are herein informally placed in Sect. Globulares for pragmatic reasons;

no formal transfers are implied.

IIIb. ser. Spinulosi

Sect. Sicci Singer, Subsect. Siccini Singer, ser. Spinulosi (Clémençon) Desjardin in Antonín & Noordeloos, Liberi. Bot.

8: 179. 1993.

= Subsect. Spinulosi Clémençon, Z. Mykol. 48: 15. 1982.

= Ser. Actinopodes Singer pro parte, Fl. Neotrop. Monogr. 17: 236. 1976.

– Type species: Marasmius cohaerens (Pers.) Cooke & Quél.

21. Marasmius dendrosetosus J.E. Shay & Desjardin, sp. nov. (Fig. 13, Plate 5)

MycoBank no.: MB 818619

Holotype:—MADAGASCAR. Region Atsinanana, District Brickaville, Commune Andevoranto, Andavakimena Forest, elev. 0–8 m,

GPS: 18˚ 53.231’ S, 49˚ 07.490’ E, 28 January 2014, J.E. Shay 205 (SFSU).

Etymology:—dendro-setosus–referring to the branched, tree-shaped pileosetae.

Description:—Pileus 2–9 mm diam, convex to plano-convex; disc rugulose; margin smooth; surface dry, glabrous;

cream to orangish white (4A2–3, 5A2–3) or greyish orange (5B3), lighter towards the margin. Context thin (<1 mm),

concolorous. Lamellae adnate, distant (8–12) with 3–6 series of lamellulae, narrow, buff to cream (4A2–3, 5A2–3),

non-marginate. Stipe 3–11 × 0.5–0.8 mm, central, cylindrical, hollow, non-insititious; surface pruinose; apex buff

(4A3, 5A3), centrally light brown (6D5), base dark brown (6F5–7). Odor and taste not distinctive.

Basidiospores 7.2–9.6 (–10.4) × 3.2–4 (–4.8) µm [xmr = 8.2–9.1 × 3.9 µm; xmm = 8.66 ± 0.68 × 3.88 ± 0.02 µm;

Q = 1.6–2.8; Qmr = 2.12–2.36; Qmm = 2.24 ± 0.17, n = 7–25; s = 2], ellipsoid, smooth, hyaline, inamyloid, thin-walled.

Basidia 20–24 × 6.4–8 μm, clavate, 4-spored, hyaline, inamyloid, thin-walled. Basidioles 16–21.6 × 5.6–7.2 μm,

clavate to fusoid, hyaline, inamyloid, thin-walled. Cheilocystidia abundant, of Siccus-type broom cells; main body

14.4–18.4 × 5.6–7.2 μm, clavate to cylindrical or irregular, seldom 2–3-lobed, hyaline, inamyloid, thin-walled; apical

setulae dense, 0.8–1.6 × 0.8–1.6 μm, cylindrical or irregular, sometimes branched, hyaline, inamyloid, thin-walled.

Pleurocystidia absent. Pileipellis mottled, a hymeniform layer of three types of cells: 1) thin-walled Siccus-type broom

cells with main body 6–20 × 5.6–8.8 μm, clavate or irregular, seldom 2–3-lobed, hyaline, inamyloid; apical setulae

0.8–8 × 0.8–1.6 μm, cylindrical to conical or irregular, hyaline, inamyloid, thin-walled, branched; 2) thick-walled

Siccus-type broom cells with main body 16.8–21.6 × 5.6–8 µm, clavate, lobed, hyaline, inamyloid; apical setulae 0.8–8

× 0.8–3.2 μm, cylindrical to conical, hyaline, inamyloid, thick-walled; 3) pileosetae 40–300 × 1.6–8 μm, cylindrical to

antler-like, often highly branched, clustered, hyaline, inamyloid, thick-walled. Pileus trama interwoven; hyphae 2.4–

7.2 μm diam, smooth, hyaline, dextrinoid, thin-walled. Lamellar trama regular; hyphae 2.4–7.2 μm diam, cylindrical,

smooth, hyaline, inamyloid, thin-walled. Stipe tissue monomitic; cortical hyphae 2.4–7.2 μm diam, parallel, cylindrical,

smooth, light brown to pale yellow, dextrinoid, thick-walled; medullary hyphae 2.4–8 μm diam, parallel, cylindrical,

hyaline, dextrinoid, thin-walled. Caulocystidia of Siccus-type broom cells emerging directly from hyphae; main body

3.2–4 × 1.6–5.6 μm, clavate or irregular, frequently lobed, hyaline, inamyloid, thin-walled; apical setulae cylindrical

or irregular, branched, hyaline, inamyloid, thin-walled. Clamp connections present.

FIGURE 13. Marasmius dendrosetosus (JES 205, JES 211) a) basidiospores; b) basidia; c) basidioles; d) cheilocystidia; e) thick-walled

pileipellis cells; f) thin-walled pileipellis cells; g) pileosetae and stipe surface broom cells (bottom center). Scale bar = 10 µm. Illustrated

by J.E. Shay.

SHAY ET AL.

Habit, habitat and known distribution:—Solitary or in gregarious clusters on stems and leaves of Uapaca littoralis

and Pandanus. Madagascar.

Material examined:—MADAGASCAR. Region Atsinanana, District Brickaville, Commune Andevoranto,

Andavakimena Forest, elev. 0–8 m, GPS: 18˚ 53.231’ S, 49˚ 07.490’ E, 28 January 2014, J.E. Shay 205 (SFSU); same

location, GPS: 18˚ 53.082’ S, 49˚ 07.559’ E, 30 January 2014, J.E. Shay 211 (SFSU).

Notes:—Marasmius dendrosetosus is characterized by a small (<10 mm) smooth cream to orangish white pileus,

subdistant non-marginate lamellae, a minutely pruinose stipe, basidiospores in the range 7.2–10.4 × 3.2–4.8 µm,

Siccus-type cheilocystidia and caulocystidia, a pileipellis composed of Siccus-type broom cells and scattered hyaline

branched pileosetae up to 300 µm long, and growth on dicotyledonous leaves and sticks. Morphology and molecular

data indicate that this new species is allied with M. longisetosus J.S. Oliveira & Capelari, describe recently from Brazil

(Oliveira et al. 2014). Marasmius longisetosus differs in forming a more deeply pigmented pileus (yellowish orange

to pure orange), slightly longer basidiospores (mean 10.5 µm), and shorter, unbranched pileosetae. ITS sequences

of M. dendrosetosus (KX148995, KX148996) are only 95% similar to the holotype specimen of M. longisetosus

(JX424040), and align sister to the latter in the ITS phylogenetic analysis (99% BS, 1.0 PP; Fig. 1b). Marasmius

jalapensis Murrill, reported from tropical Africa, is also similar, but forms more crowded lamellae, a longer stipe

(40–60 mm), has hymenial setae, shorter and broader pileosetae, and numerous caulosetae (Antonín 2007).

PLATE 5. Basidiocarps representing sect. Sicci ser. Spinulosi a) Marasmius dendrosetosus (JES 205) b) Marasmius nummularius (JES

124). Scale bar = 10 mm. Photography by D.S. Newman.

22. Marasmius nummularius Berk. & Broome, J. Linn. Soc., Bot. 14 (73): 33. 1873 (1875). (Fig. 14, Plate 5)

Type:—SRI LANKA. Kandy District, Peradeniya, Thwaites 102 cum icone (K!).

Description:—Pileus 2–7 mm diam, convex to plano-convex; margin striate; surface glabrous; orangish brown to

ochraceous (5B8). Context thin. Lamellae adnexed, subdistant (14–16) with 3–4 series of lamellulae, white with brown

to orangish brown edges. Stipe 20–48 × 1 mm, central, tough, non-insititious; surface dull, hispid; apex white, base

light brown to orange-brown (6D6). Odor and taste not distinctive.

FIGURE 14. Marasmius nummularius (JES 121, JES 124) a) basidiospores; b) basidia; c) basidioles; d) cheilocystidia; e) pleurocystidia;

f) pileipellis cells; g) caulosetae. Scale bar = 10 µm. Illustrated by J.E. Shay.

Basidiospores (10.4–) 11.2–14.4 × 3.2–5.6 μm [xmr = 12.4–12.7 × 4.3–4.8 µm; xmm = 12.56 ± 0.21 × 4.56 ± 0.39

µm; Q = 1.9–3.6; Qmr = 2.69–2.92; Qmm = 2.80 ± 0.16, n = 25, s = 2], ellipsoid, smooth, hyaline, inamyloid, thin-walled.

Basidia 17.6–25.6 × 6.4–8 μm, clavate to broadly clavate or cylindrical, 4-spored, hyaline, inamyloid, thin-walled.

Basidioles 16.8–25.6 × 5.6–8.8 μm, clavate to broadly clavate, cylindrical or fusoid, hyaline, inamyloid, thin-walled.

Cheilocystidia of Siccus-type broom cells; main body 6–20 × 6–9.6 μm, broadly clavate, hyaline to pale yellow brown,

dextrinoid, thick-walled; apical setulae 0.8–10.4 × 0.8–2.4 μm, numerous, cylindrical to conical, subacute, hyaline,

inamyloid, thick-walled. Pleurocystidia scattered, not conspicuous, 22–28 × 6.5–7.5 µm, fusoid, hyaline, refractive,

inamyloid, thin-walled. Pileipellis mottled, a hymeniform layer of Siccus-type broom cells; main body 9.6–20 × 6.4–

8.8 μm, cylindrical to clavate or subglobose, hyaline to light brown, dextrinoid, thick-walled; apical setulae 1.6–12.8

× 0.5–2.4 μm, cylindrical to conical, hyaline, inamyloid, thick-walled. Pileus trama interwoven; hyphae 3.2–7.2

μm diam, cylindrical, smooth, hyaline, dextrinoid, thin-walled. Lamellar trama regular; hyphae 3.2–5.6 μm diam,

cylindrical, smooth, hyaline, weakly dextrinoid, thin-walled. Stipe tissue monomitic; cortical hyphae 2.4–4 μm diam,

SHAY ET AL.

parallel, cylindrical, hyaline to pale yellow, dextrinoid, thick-walled; medullary hyphae 3.2–9.6 μm diam, parallel,

cylindrical to inflated, hyaline, inamyloid, thick-walled. Caulosetae 28–109 × 7.2–36 μm, conical to cylindrical, fusoid

or lageniform, hyaline, inamyloid, thick-walled. Clamp connections present.

Habit, habitat and known distribution:—Solitary to gregarious on leaf litter of Dalberjia (Fabaceae) and bark of

unknown trees. Indonesia (Java), Madagascar, Sri Lanka, Thailand.

Material examined:—MADAGASCAR. Commune Ranomafana, District Ifanadiana, Region Vatovavy-

Fitovinany, Ranomafana National Park, Talatakely Trail, elev. 937–973 m, GPS: 21˚ 15.237’ S, 47˚ 25.183’ E, 20

January 2014, J.E. Shay 121 (SFSU); same location and date, J.E. Shay 124 (SFSU).

Notes:—The Madagascar material is characterized by small (2–7 mm diam) orangish brown to ochraceous pileus,

subdistant orangish brown-marginate lamellae, a hispid orangish brown to brown non-insititious stipe, basidiospores

with mean 12.4–12.7 × 4.3–4.8 µm, Siccus-type cheilocystidia and pileipellis cells with setulae up to 12 mm long,

small fusoid pleurocystidia, rare pileosetae, conspicuous caulosetae, and growth on woody debris. Populations of M.

nummularius in Thailand (Wannathes et al. 2009a) and Indonesia (Desjardin et al. 2000) differ in lacking the small

pleurocystidia, and often have more reddish brown tones to the pileus. An ITS sequence of Madagascar material

(KX148979) shows 98% similarity to two Thai sequences (EU935492, EU935493) forming a well-supported clade

(90% BS, 1.0 PP) (Fig. 1b). The African species M. castaneovelutinus Henn. and M. fulvovelutinus Beeli differ in

forming larger (4–35 mm diam), chestnut brown to reddish brown pilei, non-marginate lamellae, larger basidiospores

(14–18 × 4–6 µm), and more conspicuous pleurocystidia (Antonín 2007).

IIIc. ser. Atrorubentes

Sect. Sicci Singer, Ser. Atrorubentes Desjardin & E. Horak, Bibl. Mycol. 168: 27. 1997.

= Ser. Actinopodes Singer pro parte, Fl. Neotrop. Monogr. 17: 236. 1976.

– Type species: Marasmius atrorubens (Berk.) Mont.

23. Marasmius corrugatiformis Singer, Bull. Jard. Bot. État Brux. 34: 374. 1964. (Fig. 15, Plate 6)

Type:—DR CONGO. Near Yambao, 21 June 1939, J. Louis 15275 (BR 11426–77).

Description:—Pileus 12–17 mm diam, convex to plano-convex; disc and margin smooth to rugulose; surface, dry,

glabrous; orangish red or orange. Context thin, buff. Lamellae subfree, close, narrow, white to buff (5A4–5), non-

marginate. Stipe 30–49 × 1 mm, central, cylindrical, hollow; surface pruinose; apex cream to yellow, becoming orange

(7C–E7–8) to brown (6D7) towards the base. Odor and taste not distinctive.

Basidiospores (6.4–) 8–11.2 × 3.2–4.8 µm [xm = 9.08 ± 1.08 × 4.04 ± 0.33 µm; Q = 1.67–2.80; Qm = 2.26 ±

0.14, n = 25, s = 1], ellipsoid, smooth, hyaline, inamyloid, thin-walled. Basidia 16.8–17.6 × 7.2 μm, clavate, 4-

spored, hyaline, inamyloid, thin-walled. Basidioles 8–15.2 × 3.2–6.4 μm, clavate to fusoid, hyaline, inamyloid, thin-

walled. Cheilocystidia of two types of cells: 1) Siccus-type broom cells with main body 8–17.6 × 6–12 μm, clavate to

cylindrical or irregular, hyaline, inamyloid, apically thick-walled; apical setulae 0.8–10.4 × 0.8–1.6 μm, cylindrical to

conical, hyaline to golden brown, inamyloid, thick-walled; 2) interspersed non-setulose cells, clavate, smooth, hyaline,

inamyloid, thin-walled. Pleurocystidia absent. Pileipellis mottled, a hymeniform layer of Siccus-type broom cells; main

body 8–20.8 × 3.2–9.6 μm, clavate to broadly clavate, cylindrical or irregular, seldom 2–3-lobed, smooth, hyaline to

brown, inamyloid, apically thick-walled; apical setulae 0.8–12.8 × 0.8–1.6 μm, cylindrical to conical, seldom branched,

erect, obtuse, pale brown to hyaline, inamyloid, thick-walled. Pileus trama interwoven; hyphae 1.6–12 μm diam,

cylindrical, smooth, hyaline, dextrinoid, thin-walled. Lamellar trama regular; hyphae 3.2–19.2 μm diam, cylindrical

to inflated, smooth, pale yellowish brown to hyaline, weakly dextrinoid, thin-walled. Stipe tissue monomitic; cortical

hyphae 2.4–12 μm diam, parallel, cylindrical to inflated, smooth, pale yellowish brown to hyaline, dextrinoid, thick-

walled; medullary hyphae 2.4–9.6 μm diam, parallel, cylindrical to inflated, smooth, hyaline, dextrinoid, thin-walled.

Caulocystidia 11.2–24 × 5.6–7.2 μm, versiform, clavate to lageniform or irregular, seldom lobed, smooth, hyaline,

dextrinoid, thin-walled. Clamp connections present.

Habit, habitat and known distribution:—Solitary on leaves of Cryptocarya (Lauraceae). Africa (Cameroon, DR

Congo, Ghana, Ivory coast, Uganda), Madagascar.

Material examined:—MADAGASCAR. Region Vatovavy-Fitovinany, District Ifanadiana, Commune Ranomafana,

Ranomafana National Park, Piste B, elev. 1004 m, GPS: 21˚ 15.413’ S, 47˚ 25.253’ E, 22 January 2014, J.E. Shay 164

(SFSU); Ranomafana National Park, by the river, January 2013, Lockwood 2132268W250 (SFSU); Andasibe, moist

mountain forest, 12 February 1997, leg. B. Buyck, G. Eyssartier and P.-A. Moreau, Buyck 97.425 (PC).

FIGURE 15. Marasmius corrugatiformis (JES 164, Lockwood 2132268W250) a) basidiospores; b) basidia; c) basidioles; d)

cheilocystidia; e) pileipellis cells; f) caulocystidia. Scale bar = 10 µm. Illustrated by J.E. Shay.

Notes:—Marasmius corrugatiformis is characterized by a relatively small (12–17 mm), rugulose, reddish orange

to orange pileus, close, non-marginate lamellae, a pruinose stipe with cream-yellow apex and brownish orange base,

basidiospores with mean 9 × 4 µm, Siccus-type cheilocystidia plus a few smooth, clavate cells interspersed, simple,

broadly rounded, cylindrical to clavate caulocystidia, and a lack of Siccus-type broom cells on the stipe. It is similar to

M. katangensis Singer, but the latter has only one type of cheilocystidia (Siccus-type), and two types of caulocystidia

(broom cells and simple cylindrical cells). Repeated attempts to generate ITS sequences from JES 164 were unsuccessful;

however, an ITS sequence of Madagascar material determined as M. corrugatiformis (Buyck 97.425, KX148981)

formed a clade with several sequences of M. corrugatiformis from São Tomé (KX953756, KX953757) but with low

support (Fig. 1b).

24. Marasmius katangensis Singer, Bull. Jard. Bot. État Brux. 34: 375. 1964. (Fig. 16, Plate 6)

Type:—DR CONGO. Shaba Province, Kipopo, 10 January 1961, M.C. Schmitz–Levecq 315 (BR 11476–30).

Description:—Pileus 6–16 mm diam, plano-convex to umbonate, with a dark brown papilla; margin striate; surface

dry, glabrous; dark orangish brown to orange (6B–D6–8). Context thin, light brown (6B3). Lamellae adnate, close

with 6 series of lamellulae, narrow (0.7–1.5 mm), cream (5A2), non-marginate. Stipe 16–50 × 1–2 mm diam, central,

cylindrical, hollow; surface pruinose; apex orangish white (5A3–4), centrally yellow (5B6), base orangish brown to

brown (6D6–8). Odor and taste not distinctive.

Basidiospores 7.2–8.8 × 3.2–4.8 µm [xm = 7.79 ± 0.48 × 3.99 ± 0.38 µm; Q = 1.67–2.50; Qm = 1.97 ± 0.21,

n = 25, s = 1], ellipsoid, smooth, hyaline, inamyloid, thin-walled. Basidia 16–23.2 × 7.2 µm, clavate, 4-spored,

hyaline, inamyloid, thin-walled; sterigmata 2.4–4.8 × 0.8–1.6 µm. Basidioles 13.6–24.8 × 4–5.6 µm, clavate, hyaline,

inamyloid, thin-walled. Cheilocystidia abundant, of Siccus-type broom cells; main body 9.6–23.2 × 4.8–7.2 µm,

clavate or irregular, seldom 2-lobed, hyaline, inamyloid, apically thick-walled; apical setulae 3.2–8.8 × 0.8–1.6

µm, cylindrical to conical, seldom branched, light yellowish brown, inamyloid, thick-walled. Pleurocystidia absent.

Pileipellis mottled, a hymeniform layer of Siccus-type broom cells; main body 8–15.2 × 7.2–8 µm, clavate or irregular,

SHAY ET AL.

seldom 2-lobed, hyaline, inamyloid, thin-walled; apical setulae 2.4–20 × 0.8–1.6 µm, 7–9 setulae per cell, cylindrical

to conical, light yellowish brown, inamyloid, thick-walled. Pileus trama interwoven; hyphae 1.6–12.8 µm diam,

cylindrical to inflated, smooth, hyaline, strongly dextrinoid, thin-walled. Lamellar trama regular; hyphae 3.2–6.4 µm

diam, cylindrical to inflated, smooth, hyaline, strongly dextrinoid, thin-walled. Stipe tissue monomitic; cortical hyphae

4–9.6 µm diam, parallel, cylindrical, smooth, hyaline, dextrinoid, thick-walled; medullary hyphae 3.2–14.4 µm diam,

parallel, cylindrical to inflated, smooth, hyaline, dextrinoid, thin-walled. Caulocystidia of two types; 1) scattered

Siccus-type broom cells; main body 10.4–20 × 5.6–7.2 µm, clavate or irregular, seldom lobed, hyaline, inamyloid,

thin-walled; apical setulae 0.8–14.4 × 0.8–2.4 µm, cylindrical to conical, seldom branched, hyaline, inamyloid, thick-

walled; 2) smooth non-setulose cystidia with main body 20.8–48 × 7.2–11.2 µm, clavate to cylindrical or irregular,

smooth, hyaline, inamyloid, thick-walled. Clamp connections present.

FIGURE 16. Marasmius katangensis (JES 227) a) basidiospores; b) basidia; c) basidioles; d) cheilocystidia; e) pileipellis cells; f) Siccus-

type caulocystidia; g) non-setulose caulocystidia. Scale bar = 10 µm. Illustrated by J.E. Shay.

Habit, habitat and known distribution:—Solitary or in small clusters on wood and leaf litter of undetermined trees.

Africa (Benin, DR Congo, Kenya, Malawi, Nigeria, Tanzania, Uganda), Madagascar.

Material examined:—MADAGASCAR. Region Analamanga, District Ankazobe, Commune Ambatoharanama,

Ambohitantely Forest Reserve, Sentier Botanique, elev. 1574 m, GPS: 18˚ 11.504’ S, 47˚ 17.074’ E, 6 February 2014,

J.E. Shay 227 (SFSU).

Notes:—Marasmius katangensis has a centrally rugulose, dark orangish brown pileus with orange margin,

close lamellae, a pruinose stipe with two types of caulocystidia, relatively small basidiospores with mean 7.8 × 4.0

µm, no pleurocystidia, and no setae. Morphologically, the species is similar to M. corrugatiformis, but the latter

has cheilocystidia of two types, and caulocystidia of only one type (non-setulose, cylindrical to clavate). JES 227,

determined here as M. katangensis, is easily confused with Madagascar material determined by Antonín & Buyck

(2006) as M. corrugatiformis (KX148981), but ITS sequences clearly distinguish the two (Fig. 1a).

Based solely on ITS data, JES 227 (KX148991) is basal to a clade containing M. occultatiformis Antonín, Ryoo

& H.D. Shin, described from Korea on detritus of Acer and Juglans. Morphologically, JES 227 is very similar to M.

occultatiformis, differing primarily in the latter species having a glabrous stipe lacking caulocystidia; other features are

indistinguishable.

PLATE 6. Basidiocarps representing sect. Sicci ser. Atrorubentes a) Marasmius corrugatiformis (JES 164) b) Marasmius katangensis

(JES 227). Scale bar = 10 mm. Photography by D.S. Newman.

IIId. ser. Leonini

Sect. Sicci Singer, Subsect. Siccini Singer, ser. Leonini Singer, Fl. Neotrop. Monogr. 17: 160. 1976.

–Type species: Marasmius leoninus Berk.

25. Marasmius sokola J.E. Shay & Desjardin, sp. nov. (Fig. 17)

MycoBank no.: MB 818621

Holotype:—MADAGASCAR. Commune Ranomafana, District Ifanadiana, Region Vatovavy-Fitovinany, Ranomafana National Park, 22

Jan. 2014, J.E. Shay 154 (SFSU).

Etymology:—sokola–Malagasy for chocolate, referring the dark chocolate brown colored basidiomes.

Description:—Pileus 20 mm diam, campanulate; disc rugulose; margin sulcate; surface dull, glabrous; dark brown (5E7).

Context thin. Lamellae subfree, distant (11), no lamellulae, broad (4 mm), light grey (5C3), with brown edges. Stipe 45 ×

1 mm, central, cylindrical, hollow, pliant; surface glabrous; dark greyish brown (6F4). Odor and taste not distinctive.

Basidiospores (16.8–) 18.4–23.2 (–26.4) × 4–4.8 μm [xm = 21.93 ± 2.36 × 4.40 ± 0.40 µm; Q = 3.50–6.40; Qm

= 5.03 ± 1.20, n = 25, s =1], subcylindrical to subfusoid, smooth, hyaline, inamyloid, thin-walled. Basidia 32.8–

46.4 × 6.4–8 μm, clavate, 4-spored, hyaline, inamyloid, thin-walled. Basidioles 29.6–42.4 × 7.2–8.8 μm, clavate,

hyaline, inamyloid, thin-walled. Cheilocystidia abundant, of Siccus-type broom cells; main body 14.4–22.4 × 6.4–8.8

μm, clavate to subglobose or irregular, seldom 2–3-lobed, hyaline, inamyloid, thin-walled; apical setulae 0.8–13.6 ×

0.8–1.6 μm, cylindrical to conical, often branched, brown, inamyloid, thick-walled. Pleurocystidia absent. Pileipellis

mottled, a hymeniform layer of Siccus-type broom cells; main body 8.8–22.4 × 5.6–10.4 μm, cylindrical to clavate

or irregular, seldom 2–3-lobed, hyaline, inamyloid, thick-walled; apical setulae 0.8–9.6 × 0.8–1.6 μm, cylindrical to

conical, seldom branched, hyaline to brown, inamyloid, thick-walled. Pileus trama interwoven; hyphae 2.4–17.6 μm

diam, cylindrical to inflated, rough, hyaline, dextrinoid, thick-walled. Lamellar trama regular; hyphae 3.2–8.8 μm

diam, cylindrical, smooth, hyaline, dextrinoid, thin-walled. Stipe tissue monomitic; cortical hyphae 3.2–11.2 μm diam,

cylindrical, smooth, pale greenish brown, dextrinoid, thick-walled; medullary hyphae 2.4–12.8 μm diam, cylindrical

to inflated, smooth, hyaline to pale brown, inamyloid, thin-walled. Caulocystidia absent. Clamp connections present.

Habit, habitat and known distribution:—Solitary on woody sticks of Weinmannia (Cunoniaceae). Madagascar.

Material examined:—MADAGASCAR. Region Vatovavy-Fitovinany, District Ifanadiana, Commune Ranomafana,

Ranomafana National Park, Piste B, elev. 1004 m, GPS: 21˚ 15.413’ S, 47˚ 25.253’ E, 22 January 2014, J.E. Shay 154

(SFSU).

SHAY ET AL.

FIGURE 17. Marasmius sokola (JES 154) a) basidiospores; b) basidia; c) basidioles; d) cheilocystidia; e) pileipellis cells. Scale bar = 10

µm. Illustrated by J.E. Shay.

Notes:—Marasmius sokola is characterized by a dark brown, smooth to rugulose pileus 20 mm diam, distant

(11), broad (4 mm), brown-marginate lamellae, a glabrous, dark brown stipe 45 mm long, basidiospores with mean

21.9 × 4.4 µm (Q = 5.0), basidia 32–46 µm long, Siccus-type cheilocystidia and pileipellis cells with brown setulae

up to 12 µm long, no pleurocystidia or caulocystidia, and growth on woody dicotyledonous debris. The new species

shows closest phenetic similarity to M. carcharus Singer, M. selangorensis Y.S. Tan & Desjardin, and M. mazatecus

Singer. Marasmius carcharus, described from the DR Congo, differs in a pinkish brown pileus, broader (6–7 mm),

non-marginate lamellae, cheilocystidia with shorter apical setulae (up to 8 µm), and growth on dicotyledonous leaves

(Antonín 2007). Marasmius selangorensis, described from Malaysia, differs in a paler brown pileus with pink tones,

narrower and more numerous (12–18) lamellae, and growth on dead dicotyledonous leaves (Tan et al. 2009); its

micromorphology is indistinguishable from M. sokola. ITS sequences of two specimens of M. selangorensis from

Malaysia (Fig. 1a), however, show only 70% similarity to JES 154 (KX148994) (Fig. 1b) and are distant in the ITS

phylogeny. Marasmius mazatecus, described from Mexico, differs in an orange-ferruginous pileus, fewer (9) and

narrower (2 mm) lamellae with orange-ferruginous edges, a shorter (20 mm) stipe, and slightly shorter basidiospores

(17–21 µm) (Singer 1976). Marasmius sokola is on a long branch in the ITS phylogeny, sister to M. imitarius Wannathes,

Desjardin & Lumyong, a species described from Thailand.

26. Marasmius rammelooi Antonín, Mycotaxon 89(2): 410. 2004

Type:—MAURITIUS. Trou d’ eau douce, 10 June 1990, leg. J. Rammeloo 9251 (BR 6902-15).

For descriptions and illustrations of Madagascar material, refer to Antonín (2004a) and Antonín & Buyck (2006).

Material was not available for sequencing.

27. Marasmius megistus Singer, Bull. Jard. Bot. État Brux. 34: 356. 1964. (Fig. 18, Plate 7)

Type:—DR CONGO. Binga, 7 May 1928, M. Goossens–Fontana 733 (BR 11492–46).

Description:—Pileus 6.5–15 mm diam, campanulate; margin deeply sulcate; surface dry, glabrous; disc yellowish

grey (4B2), sulcae pale violet brown (10E4) to reddish grey (10B2), ridges and margin buff (4A2) to cream (4A3).

Context thin. Lamellae subfree, distant (14) broad (1–2 mm wide), buff (4A2), non-marginate. Stipe 104–115 × 1–2

mm, central, cylindrical, hollow; surface glabrous; brownish grey (10D2) to brownish red (10E6). Odor and taste not

distinctive.

FIGURE 18. Marasmius megistus (JES 163, Lockwood 2132155) a) basidiospores; b) basidioles; c) cheilocystidia; d) pileipellis cells.

Scale bar = 10 µm. Illustrated by J.E. Shay.

Basidiospores (26.4–) 29.6–32.8 (–40) × 4.8–7.2 µm [xmr = 30.7–34.2 × 5.8–6.7 µm; xmm = 32.44 ± 2.49 ×

6.23 ± 0.66 µm; Q = 2.8–6.8; Qmr = 5.09–5.33; Qmm = 5.21 ± 0.17, n = 2–27, s =2], narrowly ellipsoid to clavate,

smooth, hyaline, inamyloid, thin-walled. Basidia not observed. Basidioles 18.4–57 × 5.6–10.4 μm, clavate to fusoid,

hyaline, inamyloid, thin-walled. Cheilocystidia evenly distributed, of Siccus-type broom cells; main body 16–25.6

× 4.8–9.6 μm, clavate or irregular, 2–3-lobed, hyaline, inamyloid, thin-walled; apical setulae 0.8–8 × 0.8–1.6 μm,

cylindrical to conical, sometimes branched, hyaline, inamyloid, thin-walled. Pleurocystidia absent. Pileipellis not

mottled, a hymeniform layer of Siccus-type broom cells; main body 16–32 × 6.4–10.4 μm, clavate or irregular, hyaline,

inamyloid, thin-walled; apical setulae 2.4–8 × 0.8–3.2 μm, few per cell, broadly conical to cylindrical or utriform,

seldom branched, hyaline, inamyloid, thin-walled. Pileus trama interwoven; hyphae 1.6–8 μm diam, cylindrical

smooth, hyaline, dextrinoid, thick-walled. Lamellar trama regular; hyphae 2–7.2 μm diam, cylindrical to inflated,

smooth, hyaline to pale yellow, dextrinoid, thin-walled. Stipe tissue monomitic; cortical hyphae 1.6–8 μm diam,

parallel, cylindrical, smooth, pale yellow, green brown, dextrinoid, thick-walled; medullary hyphae 1.6–9.6 μm diam,

parallel, cylindrical to inflated, smooth, hyaline, dextrinoid, thin-walled. Caulocystidia absent. Clamp connections

present.

Habit, habitat and known distribution:—Solitary on dicotyledonous leaves. Africa (Burundi, Cameroon, DR

Congo, Tanzania, Uganda), Madagascar.

Material examined:—MADAGASCAR. Region Vatovavy-Fitovinany, District Ifanadiana, Commune Ranomafana,

Ranomafana National Park, Piste B, elev. 1004 m, GPS: 21˚ 15.413’ S, 47˚ 25.253’ E, 22 January 2014, J.E. Shay

SHAY ET AL.

163 (TAN); Region Vatovavy-Fitovinany, District Ifanadiana, Commune Ranomafana, February 2013, T. Lockwood

2132155 (SFSU).

Notes:—Marasmius megistus forms some of the largest basidiospores in the genus. The Madagascan material

is distinguished by a relatively small (up to 15 mm diam) sulcate striped pileus with violet-brown sulcae and cream

ridges, distant non-marginate lamellae, a very long (up to 115 mm) glabrous stipe, basidiospores in the range 29.6–40

× 4.8–7.2 µm, no pleurocystidia, Siccus-type broom cells with few setulae, and growth singly on dicotyledonous

leaves. Antonín (2007) reports the species as forming a larger pileus (26–50 mm diam) but in all other respects the

specimens from Madagascar match those reported from tropical Africa. ITS sequences of two Madagascan specimens

(KX148992, KX148993) are sister to a specimen from São Tomé (KX953750) with strong support (100% BS, 1.0 PP;

Fig. 1a).

28. Marasmius bambusiniformis Singer, Fl. Neotrop. Monogr. 17: 167. 1976. (Fig. 19, Plate 7)

Type:—ECUADOR. Napo, Lago Agrio, 16 May 1973, Singer B7480 (F!)

Description:—Pileus 4–5 mm diam, convex to campanulate; margin smooth to sulcate; surface dull, dry, glabrous;

reddish orange becoming more orange towards the margin (6C–E8). Context thin, white. Lamellae adnate, distant

(12–17), no lamellulae, not intervenose, narrow; white with reddish brown edges. Stipe 25–30 × 0.5–1 mm, central,

cylindrical, hollow, wiry; surface glabrous, apex white (3A3), grading to brownish orange (6C–E5–8) at the base. Odor

and taste not distinctive.

FIGURE 19. Marasmius bambusiniformis (JES 199) a) basidiospores; b) basidioles; c) cheilocystidia; d) pileipellis cells. Scale bar = 10

µm. Illustrated by J.E. Shay.

Basidiospores (14.4–) 16–18.4 × 3.2–4 µm [xm = 16.35 ± 1.44 × 3.46 ± 0.36 µm; Q = 4–5.75; Qm = 4.76 ± 0.57, n =

25, s = 1], narrowly ellipsoid, smooth, hyaline, inamyloid, thin-walled. Basidia not observed. Basidioles 19.2–24 × 5.6–

7.2 μm, clavate to fusoid, hyaline, inamyloid, thin-walled. Cheilocystidia numerous, of Siccus-type broom cells; main

body 9.6–14.4 × 5.6–10.4 μm, clavate to broadly clavate, seldom 2–3-lobed, hyaline, inamyloid, apically thick-walled;

apical setulae 1.6–7.2 × 0.8 μm, dense, cylindrical to conical, seldom branched, hyaline, inamyloid, thick-walled.

Pleurocystidia absent. Pileipellis mottled, a hymeniform layer of Siccus-type broom cells; main body 10.4–17.6 ×

6.4–8 µm, clavate to broadly clavate, seldom 2–3-lobed, hyaline, inamyloid, apically thick-walled; apical setulae 1.6–

4.8 × 0.8 μm, dense, cylindrical to conical, pale yellowish brown, inamyloid, thick-walled. Pileus trama interwoven;

hyphae 1.6–16.8 μm diam, cylindrical, smooth, hyaline, inamyloid, thin-walled. Lamellar trama regular; hyphae 1.6–8

μm diam, cylindrical to inflated, smooth, hyaline, dextrinoid, thick-walled. Stipe tissue monomitic; cortical hyphae

1.6–4 μm diam, parallel, cylindrical, smooth, dark brown, dextrinoid, thick-walled; medullary hyphae 2.4–8 μm diam,

parallel, cylindrical to inflated, hyaline, dextrinoid, thin-walled. Caulocystidia absent. Clamp connections present.

PLATE 7. Basidiocarps representing sect. Sicci ser. Leonini a) Marasmius megistus (JES 163) b) Marasmius megistus (Lockwood

2132155), photo generously donated by Taylor Lockwood, c) Marasmius bambusiniformis (JES 199). Scale bar = 10 mm (a, c); 5 mm (b).

Photography by D.S. Newman.

Habit, habitat, and known distribution:—Solitary on stems of Uapaca (Phyllanthaceae). Malaysia, Madagascar,

Papua New Guinea, South America (Brazil. Ecuador), Thailand, United States (Florida).

Material examined:—MADAGASCAR. Region Alaotra-Mangoro, District Moramanga, Commune Andasibe,

Vohimana Forest, Piste 5, elev. 844 m, GPS: 18˚ 55.422’ S, 48˚ 30.201’ E, 27 January 2014, J.E. Shay 199 (TAN).

Notes:—Marasmius bambusiniformis was described originally from Ecuador (Singer 1976), and subsequently

reported from Papua New Guinea (Desjardin & Horak 1997), Malaysia (Tan et al. 2009) and Thailand (Wannathes et

al. 2009a). Distinctive features include a small (3–10 mm diam), obtusely conical, striate, reddish orange pileus, distant

(12–17) lamellae with reddish orange edges, a glabrous, non-insititious stipe lacking caulocystidia, no pleurocystidia,

Siccus-type broom cells, and growth on dicotyledonous leaves and twigs. The material from Madagascar (JES

199) matches nicely that reported from Southeast Asia and Papua New Guinea, and ITS sequences support this

determination.

Marasmius conicoparvus Antonín, C. Sharp & Stubbe is quite similar, differing primarily in forming non-marginate

lamellae and slightly shorter basidiospores (13–16 µm); this may represent the same taxon as what we report from

Madagascar, but until more material becomes available for comparison and sequencing, we prefer to recognize the

Madagascan taxon as M. bambusiniformis. Marasmius berteroi (Lév.) Murr. described from Puerto Rico, and reported

from Indonesia (Desjardin et al. 2000), is similar but has non-marginate lamellae and shorter basidiospores (12–16

SHAY ET AL.

µm). An ITS sequence of JES199 (KX148990) is sister to a Thai specimen of M. bambusiniformis (EU935521) and

together are sister to M. berteroi (FJ917632) (Fig. 1a)

IIIe. ser. Haematocephali

Sect. Sicci Singer, Subsect. Siccini Singer, ser. Haematocephali Singer, Fl. Neotrop. Monogr. 17: 201. 1976.

–Type species: Marasmius haematocephalus (Mont.) Fr.

29. Marasmius haematocephalus (Mont.) Fr., Epicr. Syst. Mycol. (Upsaliae): 382. 1838 (1836–1838). (Fig. 20, Plate

Type:—BRAZIL. Not preserved. Neotype. Guanabara, Jardin Botånico, 28 January 1961, R. Singer C 3172 (BAFC).

Description:—Pileus 2–12 mm diam, convex to campanulate, umbilicate with age, with or without a papilla; margin

sulcate to plicate; surface dry, glabrous; buff (5A2) with pale orangish pink tones (6–7A3–4), or pinkish purple (11C–

D5–6) to dull reddish purple (9D–E5–8). Context thin, cream to buff. Lamellae subfree to adnexed, distant (10–13),

narrow; buff to pale beige with pink tones, non-marginate. Stipe 12–25 × 0.1–0.2 mm, central, cylindrical or wiry,

hollow; surface glabrous; apex white to light orange (5B5), base brownish orange (6C6) to light brown or dark brown

(9F8). Odor and taste not distinctive.

FIGURE 20. Marasmius haematocephalus (JES 147, JES 202, JES 193) a) basidiospores; b) basidioles; c) cheilocystidia; d)

pleurocystidia; e) pileipellis cells. Scale bar = 10 µm. Illustrated by J.E. Shay.

Basidiospores (13.6–) 16–22 × 3.2–4.8 μm [xmr = 17.1–20.9 × 3.6–4.1 µm; xmm = 18.71 ± 1.7 × 3.84 ± 0.3

µm; Q = 3.7–7.0; Qmr = 4.80–5.08; Qmm = 4.93 ± 0.1, n = 9–25, s = 4], narrowly fusiform to elongate-ellipsoid,

smooth, hyaline, inamyloid, thin-walled. Basidia not observed. Basidioles 16–30.4 × 5.6–8.8 μm, clavate, hyaline,

inamyloid, thin-walled. Cheilocystidia abundant, of Siccus-type broom cells; main body 6.4–24 × 4.8–8.8 μm, clavate

to subglobose, cylindrical or irregular, seldom lobed, hyaline, inamyloid, thin-walled; apical setulae 0.8–7.2 × 0.8–1.6

μm, conical to cylindrical, obtuse, hyaline, inamyloid, thin-walled. Pleurocystidia 20.8–68 × 7.2–15.2 μm, clavate to

fusoid, lecythiform or lageniform, some strangulate, rarely apically bilobed, hyaline, inamyloid, thin-walled. Pileipellis

mottled, a hymeniform layer of Siccus-type broom cells; main body 6.4–24 × 4.8–9.6 μm, clavate to subglobose,

cylindrical or irregular, seldom lobed, hyaline to light brown, inamyloid, thin-walled or apically thick-walled; apical

setulae 0.5–8 × 0.5–2.4 μm, conical to cylindrical, obtuse, hyaline to light brown, inamyloid, thick-walled. Pileus

trama interwoven; hyphae 2.4–9.6 μm diam, cylindrical, smooth, hyaline, dextrinoid, thin-walled. Lamellar trama

regular; hyphae 2.4–8.8 μm diam, cylindrical to inflated, smooth, hyaline, weakly dextrinoid, thin-walled. Stipe

tissue monomitic; cortical hyphae 2.4–9.6 μm diam, parallel, cylindrical to inflated, smooth, hyaline to light yellow

brown, dextrinoid, thin-walled; medullary hyphae 2.4–15.2 μm diam, parallel, cylindrical to inflated, smooth, hyaline,

dextrinoid, thin-walled. Caulocystidia absent. Clamp connections present.

Habit, habitat, and known distribution:—Solitary, scattered on leaves of Aframomium angustifolium

(Zingiberaceae), Psorospermum (Clusiaceae), Uapacca densifolia (Phyllanthaceae), Mammea (Calophyllaceae) and

Garcinia (Clusiaceae). Pantropical: Africa (Cameroon, DR Congo, Gabon, Ghana, Ivory Coast, Kenya, Nigeria, Sierra

Leone, Tanzania, Uganda, Zimbabwe), Caribbean region, Indonesia (Java), Madagascar, Malaysia, Papua New Guinea,

South America (Brazil, Argentina), Sri Lanka, Thailand.

Material examined:—MADAGASCAR. Region Alaotra-Mangoro, District Moramanga, Commune Andasibe,

Vohimana Forest, Piste 5, elev. 844 m, GPS: 18˚ 55.422’ S, 48˚ 30.201’ E, 27 January 2014, J.E. Shay 193 (SFSU);

Region Vatovavy-Fitovinany, District Ifanadiana, Commune Ranomafana, City of Ranomafana near riverbed next to

Forest Service Station, elev. ~900 m, 20 January 2014, J.E. Shay 110 (SFSU), same region Ranomafana National Park,

Circuit Vohipara, elev. 1062 m, GPS: 21˚ 14.255’ S, 47˚ 23.409’ E, 21 January 2014, J.E. Shay 142 (SFSU); Region

Atsinanana, District Brickaville, Commune Andevoranto, Andavakimena Forest, elev. 1 m, GPS 18˚ 53.231’ S, 49˚

7.490’ E, 28 January 2014, J.E. Shay 202 (SFSU); Melville oil palm plantation, near Tamatave, 24 February 2000, B.

Buyck 00.1820 (PC).

Notes:—Marasmius haematocephalus has been reported from tropical habitats around the world and probably

represents a complex of species. Wannathes et al. (2009a) reported multicolored forms from Thailand, with pilei

ranging from yellowish white to olive, red, violet, greyish blue and brown, or a combination of many of these pigments,

all of which grouped in a well-supported clade with 1.0 PP and 99% BS support. The typical form of the species,

described originally from southern Brazil, has a reddish purple to blood red pileus, clavate basidiospores in the range

14–20 × 3.5–5 µm, conspicuous pleurocystidia, and grows on leaves and twigs (Singer 1976). Our material from

Madagascar forms sulcate, pink to pinkish purple pilei, distant (10–12), non-marginate lamellae, a glabrous, non-

insititious stipe, basidiospores with mean range 17.1–20.9 × 3.6–4.1 µm, strangulate pleurocystidia, and growth on

dicotyledonous leaves. It matches well with the material reported from Madagascar by Antonín & Buyck (2006), and

their ITS sequences (KX148977, KX148984, KX148985, KX148986, KX148987) form a well-supported clade (99%

BS, 1.0 PP; Fig. 1b) with several sequences of M. haematocephalus from Thailand (EU935527, EU935532).

Specimen JES 142 (KX148987) shows some differences with the other Madagascar specimens in forming slightly

longer basidiospores with mean 20.8 × 4.1 µm, faintly reddish purple lamellar edges near the pileus margin, and an

ITS sequence that is only 94% similar to other specimens in the well-supported M. haematocephalus clade. JES 142

consists of only a single basidiome with a reddish purple pileus, and until additional material becomes available, it is

accepted as belonging to the M. haematocephalus complex.

30. Marasmius tanaensis J.E. Shay & Desjardin sp. nov. (Fig. 21, Plate 8)

MycoBank no.: MB 818618

Holoype:—MADAGASCAR. Region Analamanga, City of Antananarivo, Parc Botanique de Zoologique (P.B.Z.T.), near the garden of

Crops Wild and Relatives (CWR), elev. 1270 m, GPS: 18˚ 55.530’ S, 47˚ 31.350’ E, 8 Feb. 2014, J.E. Shay 220 (SFSU).

Etymology:—tana-ensis–occurring in “tana”, the local name for Antananarivo where the holotype specimen was collected.

Description:—Pileus 1–3 mm diam, convex to hemispherical; margin smooth to striate; surface dull, dry, glabrous; light

orange (6A4) to orange (6B6). Context thin, pale orangish pink to light orange (6A3–4). Lamellae adnexed, subdistant

(14–15), no lamellulae, narrow (<0.3 mm); white, non-marginate. Stipe 1–22 × 0.1–0.5 mm, central, cylindrical, wiry,

hollow; surface glabrous; white at apex, becoming brownish orange (6B–C4-6) to brown (6E7) at the base. Odor and

taste not distinct.

SHAY ET AL.

FIGURE 21. Marasmius tanaensis (JES 220) a) basidiospores; b) basidia; c) basidioles; d) cheilocystidia; e) pleurocystidia; f) pileipellis

cells. Scale bar = 10 µm. Illustrated by J.E. Shay.

Basidiospores 11.2–16 × (3.2–) 4–4.8 (–5.6) μm [xm = 13.37 ± 1.41 × 4.26 ± 0.49 µm; Q = 2.43–5.0; Q m = 3.19

± 0.26, n = 31, s =1], broadly ellipsoid, smooth, hyaline, inamyloid, thin-walled. Basidia 22.4 × 5.6–6.4 μm clavate,

4-spored, hyaline, inamyloid, thin-walled; sterigmata 2.4–3.2 × 0.8 μm. Basidioles 8.8–28.8 × 4.8–7.2 μm, clavate

to fusoid, hyaline, inamyloid, thin-walled. Cheilocystidia abundant, of Siccus-type broom cells; main body 16–29.6

× 4–8 μm, clavate to broadly clavate, seldom 2–3 lobed, hyaline, inamyloid, thin-walled; apical setulae 0.8–4 × 0.8,

cylindrical to conical, seldom branched, hyaline to light brown, inamyloid, thick-walled. Pleurocystidia 22–48 × 6–9.5

µm, utriform to fusiform, mucronate, hyaline, inamyloid, thin-walled. Pileipellis mottled, a hymeniform layer of Siccus-

type broom cells; main body 12–16.8 × 4.8–9.6 μm, clavate to broadly clavate, seldom 2–3 lobed, hyaline, inamyloid,

thin-walled; apical setulae 0.8–5.6 × 0.8–1.6 μm, clustered, cylindrical to conical, seldom branching, light brown,

inamyloid, thick-walled. Pileus trama interwoven; hyphae 2.4–12 μm diam, cylindrical to inflated, smooth, hyaline,

dextrinoid, thin-walled. Lamellar trama regular; hyphae 3.2–8.8 μm diam, cylindrical to inflated, smooth, hyaline,

dextrinoid, thick-walled. Stipe tissue monomitic; cortical hyphae 2.4–6.4 μm diam, parallel, cylindrical, smooth, pale

light brown, dextrinoid, thick-walled; medullary hyphae 3.2–8.8 μm diam, parallel, cylindrical to inflated, smooth,

hyaline, dextrinoid, thin-walled. Caulocystidia absent. Clamp connections present.

Habit, habitat and known distribution:—Clustered on bamboo debris. Madagascar.

Material examined:—MADAGASCAR. Region Analamanga, City of Antananarivo, Parc Botanique de Zoologique

(P.B.Z.T.), near the garden of Crops Wild and Relatives (CWR), elev. 1270 m, GPS: 18˚ 55.530’ S, 47˚ 31.350’ E, 8

February 2014, J.E. Shay 220 (SFSU).

Notes:—Marasmius tanaensis forms tiny (1–3 mm diam), smooth to striate, orange pilei, subdistant (14–15),

non-marginate lamellae, a short (up to 22 mm), glabrous, white (upper half) to brown (base) stipe that grows on

bamboo debris, basidiospores with mean 13.4 × 4.3 µm, narrow (6–9.5 µm), mucronate pleurocystidia, Siccus-type

cheilocystidia and pileipellis cells, and no caulocystidia. The small orange pilei and growth on bamboo debris is similar

to M. bambusinus (Fr.) Fr., described from Brazil, but the latter forms fewer lamellae (6–13), longer basidiospores

(13.8–22 µm), and broader pleurocystidia (8–12.5 µm diam) (Singer 1976). We were unable to obtain a quality ITS

sequence from the holotype specimen.

31. Marasmius (cf.) grandisetulosus Singer, Bull. Jard. Bot. État Brux. 34: 379. 1964

Type:—DR CONGO. Kivu, Panzi, November 1948, M. Goossens-Fontana 5076 (BR 11460–14).

For a description and illustrations of Madagascan material, refer to Antonín & Buyck (2006). Repeated attempts to

sequence material from collection Buyck 97.004 (PC) were unsuccessful.

32. Marasmius eyssartieri Antonín & Buyck, Fungal Diversity 23: 42. 2006

Type:—MADAGASCAR. Sainte Lucie, near Fort Dauphin, 27 Jan. 1999, leg. B. Buyck & G. Eyssartier, Buyck 99.375 (PC!).

For a description and illustrations of Madagascan material, refer to Antonín & Buyck (2006). Repeated attempts to

sequence material from collection Buyck 99.375 (PC) were unsuccessful.

33. Marasmius cf. confertus var. parvisporus Antonín. Mycotaxon 89: 401. 2004

Type:—KENYA. Central Province, Nairobi District, Thika Fall, 16 March 1968, D.N. Pegler K101 (K(M) 8833).

For a description and illustrations of Madagascan material, refer to Antonín & Buyck (2006). Repeated attempts to

sequence material from collection Buyck 99.424 (PC) were unsuccessful.

34. Marasmius ferruginoides Antonín Mycotaxon 89(2): 399–422. 2004. (Fig. 22, Plate 8)

Type:—DR CONGO. Tshopo Province, Kisangani, forest near Zoo, 2 May 1984, B. Buyck 1615 (BR 11731–91).

Description:—Pileus 7–9 mm diam, campanulate, umbilicate; margin smooth to rugulose; surface dry, glabrous; orange

to reddish orange (8C7). Context thin, orangish red. Lamellae adnexed, close (17–20) with 3 series of lamellulae,

narrow; yellowish white (3A3), non-marginate. Stipe 29–50 × 0.5–1 mm, central, cylindrical, wiry, hollow; surface

glabrous; apex buff to cream (3A3), centrally light brown (5B6), base dark brown (6C4, 6F8). Odor and taste not

distinctive.

Basidiospores 10.4–12.8 × 4–4.8 μm [xm = 11.52 ± 1.07 × 4.48 ± 0.44 µm; Q = 2.17–3.20; Qm = 2.61 ± 0.48, n =5, s

=1], narrowly ellipsoid to oblong, smooth, hyaline, inamyloid, thin-walled. Basidia not observed. Basidioles 17.6–29.6

× 5.6–7.2 μm, clavate to fusoid, hyaline, inamyloid, thin-walled. Cheilocystidia scattered, of Siccus-type broom cells;

main body 17.6–24.8 × 5.6–7.2 μm, clavate to cylindrical, 2–3 lobed, hyaline, inamyloid, thin-walled; apical setulae

0.8–8.8 × 0.8–1.6 μm, cylindrical to conical, seldom branched, hyaline, inamyloid, thin-walled. Pleurocystidia few,

26.4–38.4 × 4.8–7.2 μm, subfusoid to subcylindrical, seldom lobed, often mucronate or capitate, hyaline, inamyloid,

thin-walled. Pileipellis mottled, a hymeniform layer of Siccus-type broom cells; main body 15.2–20 × 5.6–7.2 μm,

clavate to broadly clavate or irregular, light brown, inamyloid, apically thick-walled; apical setulae 1.6–5.6 × 0.8–

1.6 μm, conical, strict, often forked, hyaline, inamyloid, thick-walled. Pileus trama interwoven; hyphae 3.2–10.4

μm diam, cylindrical to inflated, smooth, hyaline, dextrinoid, thick-walled. Lamellar trama regular; hyphae 2.4–6.4

μm diam, cylindrical, smooth, hyaline, inamyloid, thick-walled. Stipe tissue monomitic; cortical hyphae 3.2–5.6 μm

diam, parallel, cylindrical, smooth, brown, dextrinoid, thick-walled; medullary hyphae 1.6–10.4 μm diam, parallel,

cylindrical, smooth, hyaline, inamyloid, thin-walled. Caulocystidia absent. Clamp connections present.

Habit, habitat and known distribution:—Solitary or in gregarious clusters on leaves of Intsia bijuga (Fabaceae).

Africa (Cameroon, DR Congo, Ghana, Kenya, Nigeria), Madagascar.

Material examined:—MADAGASCAR. Region Atsinanana, District Brickaville, Commune Andevoranto,

Andavakimena Forest, elev. 8 m, GPS: 18˚ 53.082’ S, 49˚ 07.559’ E, 30 January 2014, J.E. Shay 209 (SFSU).

Notes:—Marasmius ferruginoides is characterized by a relatively small (7–9 mm diam), campanulate, smooth to

wrinkled, orange to reddish orange pileus, close (17–20), non-collariate, non-marginate lamellae, a glabrous stipe lacking

caulocystidia, narrow (4.8–7.2 µm), mucronate pleurocystidia, Siccus-type broom cells, and growth on dicotyledonous

leaves. Antonín (2004) established M. ferruginoides for African material determined by Pegler (1977) as M. gardneri

Singer (= M. ferrugineus (Berk.) Berk. & M.A. Curtis, a different species from Brazil), and subsequently reported the

species from Cameroon, DR Congo, Ghana, Kenya, and Nigeria (Antonín 2007). The Madagascan specimen (JES

209) reported here differs from continental specimens in forming a more reddish orange pileus (rather than yellowish

SHAY ET AL.

orange). Antonín and Buyck (2006) reported M. confertus var. parvisporus from Madagascar, and our material of M.

ferruginioides would key there, but the former has smaller pilei (up to 3 mm diam), fewer lamellae (15), and shorter

basidiospores (8.5–12 µm long). An ITS sequence of JES 209 (KX148983) places M. ferruginoides in an isolated

position in the ITS phylogeny (Fig. 1a).

FIGURE 22. Marasmius ferruginoides (JES 209) a) basidiospores; b) basidioles; c) cheilocystidia; d) pleurocystidia; e) pileipellis cells.

Scale bar = 10 µm. Illustrated by J.E. Shay.

35. Marasmius hinnuleus Berk. & M.A. Curtis, J. Linn. Soc., Bot. 10(45): 297. 1868 (1869). (Fig. 23, Plate 8)

Type:—CUBA. On dead leaves, October, Wright 155 (K).

Description:—Pileus 6–8 mm diam, campanulate to hemispherical, some with a small umbo; margin sulcate; surface

dry, rugulose around disc, glabrous along margin; disc dark brown (6E6), ferruginous to brownish orange or reddish

brown (6E7–D6–7) elsewhere. Context thin (<1 mm), white. Lamellae adnate, distant (15–17), no lamellulae, broad,

not intervenose; cream to buff (4A2–3), edges ferruginous or non-marginate. Stipe 33–52 × 0.5 mm, central, cylindrical,

wiry, hollow; surface glabrous; apex light brown (5D5), base dark brown (6F8). Odor and taste not distinctive.

Basidiospores 10.4–13.6 × 2.4–4 μm [xm = 12.13 ± 0.94 × 3.24 ± 0.49 µm; Q = 3–5; Qm = 3.82 ± 0.45, n = 25, s

=1], fusoid to oblong or narrowly ellipsoid, smooth, hyaline, inamyloid, thin-walled. Basidia not observed. Basidioles

22.4–25.6 × 5.6–8 μm, clavate to fusoid, hyaline, inamyloid, thin-walled. Cheilocystidia of Siccus-type broom cells;

main body 12–23.2 × 4.8–7.2 μm, clavate to cylindrical, hyaline, inamyloid, apically thick-walled; apical setulae 0.8–

7.2 × 0.8–1.6 μm, cylindrical to conical or irregular, seldom branched, hyaline, inamyloid, thick-walled. Pleurocystidia

36–47.2 × 7–10 µm, common, subcylindrical to fusoid, some mucronate, hyaline, inamyloid, refractive, thin-walled.

Pileipellis mottled, a hymeniform layer of Siccus-type broom cells; main body 8–16.8 × 4–7.2 μm, clavate, seldom

2–3 lobed, hyaline, inamyloid, thick-walled; apical setulae 0.8–4.8 × 0.8–1.6, cylindrical to conical, light brown to

brown, inamyloid, thick-walled. Pileus trama interwoven; hyphae, 2.4–8 μm diam, cylindrical, smooth, hyaline,

dextrinoid, thin-walled. Lamellar trama regular; hyphae 2.4–8.8 μm diam, cylindrical to inflated, hyaline, dextrinoid,

thin-walled. Stipe tissue monomitic; cortical hyphae 4.8–7.2 μm diam, parallel, cylindrical, hyaline, dextrinoid, thin-

walled; medullary hyphae 4–7.2 µm, cylindrical, parallel, hyaline, dextrinoid, thin-walled. Caulocystidia absent.

Clamp connections present.

Habit, habitat, and known distribution:—Solitary or in small clusters on bamboo debris in groomed park. Cuba,

Guadeloupe, Madagascar.

PLATE 8, Basidiocarps representing sect. Sicci ser. Haematocephali a) Marasmius haematocephalus (JES 202) b) Marasmius ferruginoides

(JES 209) c) Marasmius hinnuleus (JES 217). Scale bar = 10 mm. Photography by D.S. Newman.

Material examined:—MADAGASCAR. Region Analamanga, City of Antananarivo, Parc Botanique de Zoologique

(P.B.Z.T.), near the garden of Crops Wild and Relatives (CWR), elev. 1270 m, GPS: 18˚ 55.530’ S, 47˚ 31.350’ E, 8

February 2014, J.E. Shay 217 (TAN).

Notes:—Marasmius hinnuleus, described originally from Cuba, is characterized by a small (<10 mm diam), sulcate,

brownish orange to reddish brown pileus, distant (15–17), non-marginate lamellae, a non-insititious, glabrous stipe

lacking caulocystidia, refractive, often mucronate pleurocystidia 7–10 µm diam, Siccus-type broom cells, and growth

on dead leaves. The Madagascan specimen matches quite closely material reported from the Caribbean (Singer 1976,

Pegler 1983). The species shows similarities to M. hypophaeus Berk. & M.A. Curtis, M. confertus Berk. & Broome, M.

suthepensis, and M. ferrugineus (Berk.) Berk. & M.A. Curtis. Marasmius hypophaeus forms fewer lamellae (11–13)

with brownish orange edges, a smaller stipe (23–35 mm long), larger basidiospores (x = 17.2 × 4.0 µm) and strangulate

pleurocystidia (Desjardin et al. 2000, Wannathes et al. 2009a). Marasmius confertus and M. suthepensis have nearly

smooth pilei and more numerous lamellae with multiple series of lamellulae (Antonín 2007, Wannathes et al. 2009a).

Marasmius ferrugineus forms a paler pileus, has fewer lamellae (8–10) and larger basidiospores (x = 17.0 × 4.0 µm)

(Singer 1976, Desjardin et al. 2000). An ITS sequence of JES 217 (KX148988) places M. hinnuleus in a clade with M.

hypophaeus and M. grandisetulosus with 89% BS and 1.0 PP support (Fig. 1a).

SHAY ET AL.

FIGURE 23. Marasmius hinnuleus (JES 217) a) basidiospores; b) basidioles; c) cheilocystidia; d) pleurocystidia; e) pileipellis cells. Scale

bar = 10 µm. Illustrated by J.E. Shay.

Discussion

As of 2006, only 19 species of Marasmius sensu stricto were known from Madagascar. Our fieldwork in 2014

revealed another 16 species for the region, of which 11 are new distribution reports and 5 represent new species. A

nearly doubling of the number of Marasmius species from only a month of fieldwork in 2014 suggests that continued

explorations would undoubtedly yield numerous more species. Documenting the biodiversity of Marasmius and allied

litter-decomposing species from Madagascar is in its infancy and much more work is needed.

Over 1700 epithets have been published to date in Marasmius, representing over 600 distinct species. As this

project highlights, there are numerous species awaiting discovery through intensive sampling of underexplored habitats.

Circumscribing species of Marasmius requires a combination of morphological, ecological and molecular characters.

Historically, morphology alone was used to develop infrageneric groups. All species with collariate lamellae and

insititious stipes were recognized as belonging to sect. Marasmius, and within this group, those with Rotalis-type

broom cells were grouped in subsect. Marasmius, while those with Siccus-type broom cells belonged to subsect.

Sicciformes. If a species lacked a collarium but the stipe was insititious, the species was accepted in sect. Leveilleani.

All species lacking a stipe or with a very rudimentary lateral stipe were placed in sect. Neosessiles. By far the most

diverse group of Marasmius are those species with non-collariate lamellae and non-insititious stipes, historically placed

in sections Globulares (with smooth Globulares-type cells in the pileipellis) and Sicci (with Siccus-type broom cells in

the pileipellis). Recent molecular studies by Tan et al. (2009) and Wannathes et al. (2009a) have shown that these two

sections are not monophyletic, and the current trend is to accept members of the two groups in a single section, sect.

Globulares (Antonín & Noordeloos 2010). Within this lineage, historical infrageneric classifications have distinguished

groups based on the presence or absence of setae, pleurocystidia, and simple cylindrical caulocystidia. For example,

species with setae on the pileus, hymenium and/or stipe surface were grouped in ser. Spinulosi; those with simple

cylindrical caulocystidia were placed in ser. Atrorubentes; those lacking setae and simple cylindrical caulocystidia

but with distinct pleurocystidia were accepted in ser. Haematocephali, while those lacking setae, caulocystidia and

pleurocystidia were recognized in ser. Leonini.

In most historically recognized infrageneric groups of Marasmius, species may be nearly indistinguishable in

morphology but very different molecularly. This is particularly true in sect. Marasmius, where most species are small,

have plicate pilei, distant collariate lamellae lacking lamellulae, black, wiry glabrous stipes, basidiospores in a limited

size range, lack pleurocystidia and caulocystidia, and are generally character-poor. For these taxa, molecular sequences

are invaluable in delimiting species. As in previous studies based on ITS sequences (Tan et al. 2009, Wannathes et

al. 2009a), the phylogenetic analyses presented here indicate that the historical infrageneric classification based on

morphology does not represent monophyletic lineages. The ITS phylogeny (Fig. 1) clearly indicates that setae (ser.

Spinulosi) have evolved independently numerous times, as have pleurocystidia (ser. Haematocephali), simple cylindrical

caulocystidia (ser. Atrorubentes), pileipelli with smooth Globulares-type cells (sect. Globulares), and collariate species

with Siccus-type broom cells (subsect. Sicciformes). Morphology alone does not indicate phylogenetic relationships.

Molecular datasets are needed to better understand relationships in this diverse and widespread genus.

The genus Marasmius is over 90 million years old, worldwide in distribution, and megadiverse. With such a

long evolutionary history, a quickly evolving gene region such as the ITS region shows extensive variability amongst

species. This creates problems in accurately aligning sequences for phylogenetic analyses, especially among species

from geographically distant populations, and provides low resolution at the deeper nodes. Sequences from additional

gene regions, particularly RPB2 and EF1-α, need to be analyzed to develop an infrageneric classification that better

reflects the phylogeny of Marasmius.

Acknowledgments

The authors would like to recognize San Francisco State University Department of Biology, Mycological Society of

San Francisco (MSSF), Sonoma County Mycological Association (SOMA), and the generous donors from a Kickstarter

campaign who made this project possible. We are grateful to Rokiman Letsara, Emile Randrianjohany, and their

colleagues at CNRE in Antananarivo for their valuable field assistance. We also wish to thank Dr. Vladimír Antonín

(Czech Republic), Dr. Bart Buyck (France), and Taylor Lockwood for providing samples from their collections from

Madagascar.

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Spider Fungi: New species of Marasmius and Pusillomyces in the aerial rhizomorph web-maker guild in Amazonia

Article

Full-text available

Feb 2024
FUSE

Rhizomorphs are hair- or wire-like melanized structures with structural differentiation analogous to plant roots that help fungi spread over an area and find food resources. Some species of multiple groups of the Ascomycota and the Basidiomycota produce different types of rhizomorphs. In the Agaricales, the structures are largely found in Marasmiineae, particularly in the Omphalotaceae, Marasmiaceae, and Physalacriaceae. Many rhizomorph-forming species spread over the forest floor (ground level), while others form aerial litter-trapping rhizomorph systems that hang on living branches of the forest understorey. Here, we describe three new species that form aerial systems, two in Marasmius and one in Pusillomyces, all of which inhabit areas of the Amazon rainforest in Amazonas State, Brazil. Support for these new species is based on evidence derived from morphological and molecular phylogenetic analyses using nuclear ribosomal internal transcribed spacer region (nrITS) and nuclear ribosomal large subunit (nrLSU) markers. Marasmius foliauceps sp. nov., with a pileipellis made up of Rotalis-type broom cells, is different from all other species described to date and branched along with species of the monophyletic Marasmius sect. Marasmius subsect. Marasmius, being conspecific with specimens from Guyana. Also different from all other described Marasmius species, Marasmius arachnotropus sp. nov. has a pileipellis mostly composed of Siccus-type but in transition to Rotalis-type broom cells and branched along with species of the Marasmius crinis-equi species complex. Pusillomyces cuieirasensis is morphologically very similar to Pusillomyces manuripioides to which it is sister, but it has a non-umbonate pileus and smaller basidiospores. They all form aerial litter-trapping rhizomorph systems that mimic spider webs, particularly those hanging on the understorey. Whether made of mycelia or silk, such webs serve to capture food. The aerial rhizomorph-forming ‘spider fungi’ make up a guild with hidden species diversity in tropical systems.

Morphological, phylogenetic, and genomic evidence reveals the causal agent of thread blight disease of cacao in Peru is a new species of Marasmius in the section Neosessiles, Marasmius infestans sp. nov

Article

Full-text available

Jan 2024

The thread blight disease (TBD) of cacao ( Theobroma cacao ) in the department of Amazonas, Peru was recently reported to be caused by Marasmius tenuissimus (sect. Neosessiles). This same species is known to be the main causal agent of TBD in West Africa. However, some morphological characteristics, such as the presence of rhizomorphs, the almost exclusively white color, and pileus sizes less than 5 mm, among others, differ to the description of M. tenuissimus. Therefore, we aimed to conduct a taxonomic revision of the cacao-TBD causal agent in Peru, by using thorough micro and macro morphological, phylogenetic, and nuclear and mitochondrial genomic approaches. We showed that the causal agent of TBD of cacao in Amazonas, Peru, belongs to a new species, Marasmius infestans sp. nov. This study enriches our knowledge of species in the sect. Neosessiles, and strongly suggests that the M. tenuissimus species complex is highly diverse.

Marasmius australotrichotus (Marasmiaceae), a new setose species from Australia, and an intriguing range extension for M. paratrichotus

Article

Full-text available

Oct 2023

Article

Oct 2023

The thread blight disease (TBD) of cacao ( Theobroma cacao ) in the department of Amazonas, Peru was recently reported to be caused by Marasmius tenuissimus (Sect. Neosessiles). This same species is known to be the main causal agent of TBD in West Africa. However, some morphological characteristics, such as the presence of rhizomorphs, the almost exclusively white color, and pileus sizes less than 5 mm, among others, differ to the description of M. tenuissimus. Therefore, we aimed to conduct a taxonomic revision of the cacao-TBD causal agent in Peru, by using thorough micro and macro morphological, phylogenetic, and nuclear and mitochondrial genomic approaches. We showed that the causal agent of TBD of cacao in Amazonas, Peru, belongs to a new species, Marasmius infestans sp. nov. This study enriches our knowledge of species in the Sect. Neosessiles, and strongly suggests that the M. tenuissimus species complex is highly diverse.

Catalogue of fungi in China 5. Preliminary survey of macrofungi in Medog, Southwest China

Article

Full-text available

Apr 2025

DNA-based fungal diversity in Madagascar and arrival of the ectomycorrhizal fungi to the island

Article

Full-text available

Jul 2023
BIOTROPICA

Madagascar is known for its high diversity and endemism of fauna and flora. Fungi, however, have been largely overlooked in diversity and evolution studies on the island, and whether fungi exhibit the same patterns as animals and plants has yet to be further examined. We collected fungal sporocarps and ectomycorrhizal (EcM) roots during three opportunistic surveys in five forests in Madagascar and generated a dataset of fungal Internal Transcribed Spacer (ITS) DNA sequences. We analyzed them together with all publicly available fungal ITS DNA sequences and identified 620 Operational Taxonomic Units (OTUs) from Madagascar, 10% of which contained only sequences from our surveys. Two hundred and ninety-two OTUs belonged to EcM species with /russula-lactarius, /boletus, /tomentella-telephora, /cortinarius, and /amanita being the most abundant EcM lineages. Overall, 60% of all fungi and 81% of the EcM species from Madagascar appear to be endemic. We conducted a phylogenetic analysis using all the OTUs in Amanitaceae, Boletaceae, and Russulaceae families to elucidate their relative timing of arrival in Madagascar. We found that most EcM species from Madagascar in the three families diverged less than 22 million years ago (mya), long after the separation of India and Madagascar (88 mya), which is consistent with a dispersal mediated process of arrival to the island. Our study provides the first comprehensive view of the overall DNA-based fungal diversity in Madagascar and the current state of knowledge of EcM fungi based on DNA sequences, useful for further ecological and evolutionary studies. Abstract in Malagasy is available with online material.

Mushroom Exploration in Tulung Agung Village, East Lampung and Its Utilization Potential

Article

Full-text available

Jan 2023

Mushrooms an be found in various places, such as litter, rotten wood, trees, and soil around residential areas. Tulung Agung Village in East Lampung Regency has various kinds of vegetation and several other abiotic factors that support the growth of mushroom. The study of mushroom's exploration could elaborate the knowledge about mushroom diversity in resdential areas. This study aims to explore the types of mushrooms based on the description of their fruiting bodies from the residential location of Tulung Agung Village, District Pekalongan, East Lampung Regency, Lampung. The methods used were purposive sampling technique in a shady area or under the shade of a tree, documenting the mushroom fruiting bodies hat were found, and presenting mushrooms with fruiting bodies characters. After the exploration, 18 species from 14 genera of mushrooms were found, including Auricularia auricula-judae, Collybia sp., Conocybe sp., Daldinia sp., Gymnopilus sp., Marasmius sp., Marasmius haematocephalus, Mycena sp., Lepiota sp., Parasola sp., Pycnoporus sp., Trametes sp., Tremellodendron sp., and Xylaria sp. The various potentials of each type of mushrooms that were found are for food sources, traditional medicines, biodegradation of contaminants, sources of bioluminescence, and sources of various bioactive compounds. The Mushroom that were found need molecular identification to get the correct species so that the potential of Mushroom can be utilized optimally by residents based on more specific type information.

Untangling horsehair fungi in Australia: Marasmius crinis-equi (Marasmiaceae) and related taxa

Article

Full-text available

Sep 2024

Fungi with prominent hairlike rhizomorphs, aerial habit, sparse small sporophores, and usually saprotrophic biology are commonly known as ‘horsehair or spider fungi’. The earliest descriptions of them, from the mid-late 1800s, were published from Australian material under the names Marasmius crinis-equi and M. equicrinis . For the original ‘horsehair fungus’, we review this early historical material, briefly explore the changes in the species concept over time, and investigate the nomenclatural tangle of potentially competing names. Our analysis of morphological and molecular data for over 60 collections across eastern Australia shows that material labelled as M. crinis-equi forms part of a complex of at least three closely related species and that this name has been misapplied both within Australia and internationally. An epitype is nominated for an updated concept of M. crinis-equi , and the closely related taxa, M. tropicus sp. nov . and M. kabakada sp. nov . , are described. Two more distantly related Australian taxa to which the name M. crinis-equi has been misapplied, M. perumbilicatus sp. nov . and M. argillaceus sp. nov., are also described to further stabilise the concept of the authentic horsehair fungus M. crinis-equi.

Taxonomic revision of Marasmius Fr. and Marasmiaceae Roze ex Kühner based on multigene phylogenetics and morphological evidence

Article

Jul 2024
FUNGAL DIVERS

Many taxonomic and systematic rearrangements were proposed to Marasmius Fr. since its original concept in 1835, and since 1980 when it became the type of Marasmiaceae. These were based on morphological and/or more recently molecular phylogenetic studies. This study conducted a comprehensive taxonomic and systematic evaluation of Marasmius that benefits the whole family, implementing multilocus (SSU, LSU, ITS, rpb2 and ef1-α) phylogenetic analyses integrated with morphological and other features. The resulting trees support (1) a Marasmiaceae clade-based circumscription within Marasmiineae, (2) a Marasmius clade-based circumscription within Marasmiaceae, and (3) a subgenus-section-subsection-series system. Two subgenera are proposed: Globulares and Marasmius. Marasmius auton. subgen. includes Crinis-eques sect. nov., sect. Marasmius, Sanguirotales sect. nov., Variabilispori sect. nov., and Sicciformes sect. nov., while Globulares subg. nov. groups sect. Globulares and sect. Sicci. Four subsections are proposed in sect. Globulares and three in sect. Marasmius and sect. Sicciformes. Seventeen series were defined in sect. Globulares and three in sect. Sicci. Selected traits were assessed for their phylogenetic signals within Marasmius, providing a robust framework for a natural system. Based on this analysis, Marasmiaceae includes Chaetocalathus, Crinipellis, Marasmius and Moniliophthora/Paramarasmius, and Campanellaceae fam. nov. includes Brunneocorticium, Campanella/Tetrapyrgos, Neocampanella, and Marasmiellus sect. Candidi. New species, names, combinations and epitypes are also proposed.

Taxonomic and phylogenetic evidence reveal a new species and a new record of the genus Marasmius (Marasmiaceae) from Pakistan

Article

Full-text available

Apr 2024

This research reveals a significant contribution to fungal taxonomy through the integration of morphological and phylogenetic analyses, utilizing ITS and 28S sequences. Focused on the Marasmius genus in Pakistan, the combined approach has successfully identified a new species, namely M. pakistanicus, and a new record, M. paratrichotus. The new species is characterized by its small orange pileus with a grooved surface, creamy lamellae, lacrymoid to subcylindrical basidiospores, broadly clavate cheilocystidia, and a hymeniderm pileipellis composed of inconspicuous or broadly clavate elements. This study enhances our knowledge of fungal diversity by scrutinizing both morphological traits and genetic profiles. The morphological and phylogenetic analyses based on the combined ITS-28S dataset confirmed that both species belong to the Marasmius sect. Sicci. These findings not only broaden our understanding of local fungal biodiversity but also underscore the efficacy of a comprehensive methodology. They emphasize the synergy between morphology and phylogenetics as instrumental in advancing our knowledge of fungal communities.

Marasmius sensu stricto in Peninsular Malaysia

Article

Full-text available

Jul 2009

A total of 43 species of Marasmius sensu stricto belonging to six sections (Globulares, Hygrometrici, Leveilleani, Marasmius, Neosessiles and Sicci) are reported from Peninsular Malaysia. Macro- and micromorphological characters in combination with ITS sequence data were used to circumscribe species. Of the 43 species reported herein, 9 represent new species, viz., M. angustilamellatus, M. diminutivus, M. distantifolius, M. iras, M. kanchingensis, M. kuthubutheenii, M. musicolor, M. ochropoides, and M. olivascens. An additional 19 species are new records for Peninsular Malaysia. Phylogenetic analyses of 36 Malaysian Marasmius species were conducted based oil sequence data from nrDNA ITS-1-5.8S-ITS-2 gene regions. The ITS data support the circumscription of species based oil morphological characters and are useful in aiding species delimitations. The ITS data do not, however, Support the current sectional delimitation of Marasmius wherein most sections are not monophyletic. Comprehensive descriptions, illustrations and photographs of each species, a key to aid in diagnosis, comparisons with allied species, and a preliminary phylogenetic construct are provided.

Nuclear ribosomal internal transcribed spacer (ITS) region as a universal DNA barcode marker for Fungi

Article

Full-text available

Jan 2012

Six DNA regions were evaluated as potential DNA barcodes for Fungi, the second largest kingdom of eukaryotic life, by a multinational, multilaboratory consortium. The region of the mitochondrial cytochrome c oxidase subunit 1 used as the animal barcode was excluded as a potential marker, because it is difficult to amplify in fungi, often includes large introns, and can be insufficiently variable. Three subunits from the nuclear ribosomal RNA cistron were compared together with regions of three representative protein-coding genes (largest subunit of RNA polymerase II, second largest subunit of RNA polymerase II, and minichromosome maintenance protein). Although the protein-coding gene regions often had a higher percent of correct identification compared with ribosomal markers, low PCR amplification and sequencing success eliminated them as candidates for a universal fungal barcode. Among the regions of the ribosomal cistron, the internal transcribed spacer (ITS) region has the highest probability of successful identification for the broadest range of fungi, with the most clearly defined barcode gap between inter- and intraspecific variation. The nuclear ribosomal large subunit, a popular phylogenetic marker in certain groups, had superior species resolution in some taxonomic groups, such as the early diverging lineages and the ascomycete yeasts, but was otherwise slightly inferior to the ITS. The nuclear ribosomal small subunit has poor species-level resolution in fungi. ITS will be formally proposed for adoption as the primary fungal barcode marker to the Consortium for the Barcode of Life, with the possibility that supplementary barcodes may be developed for particular narrowly circumscribed taxonomic groups.

Marasmius (Basidiomycotina, Marasmiaceae) in Madagascar and the Mascarenes.

Article

Full-text available

Oct 2006
FUNGAL DIVERS

Antonín, V. and Buyck, B. (2006). Marasmius (Basidiomycota, Marasmiaceae) in Madagascar and the Mascarenes. Fungal Diversity 23: 17-50. Twenty six collections representing 19 taxa of the genus Marasmius from Madagascar, Mauritius and Réunion were studied. The following new taxa are described: Marasmius andasibensis, Marasmius andasibensis var. obscurostipitatus, M. brunneoaurantiacus and M. curreyi var. bicystidiatus in sect. Marasmius; M. cecropiformis and Marasmius neosessiliformis (introduced as a nom. prov. because of the absence of the macroscopic description) in sect. Neosessiles; M. pseudocyphella in sect. Hygrometrici and M. eyssartieri in sect. Sicci.

Phylogenetic relationships in the gymnopoid and marasmioid fungi (Basidiomycetes, euagarics clade)

Article

May 2005

Three distinct lineages of gymnopoid and marasmioid fungi are recognized in parsimony and Bayesian analyses of nLSU rDNA sequences. One lineage contains the genera Lentinula, Rhodocollybia, Tetrapyrgos, a resurrected and redefined Mycetinis, and two unresolved clades designated /marasmiellus and /gymnopus. /marasmiellus includes the type species of Marasmiellus and is dominated by members of Gymnopus sect. Vestipedes. /gymnopus includes the type species of Gymnopus, Micromphale and Setulipes, and members of Gymnopus sect. Levipedes. A second lineage includes the genera Marasmius s.s. and Crinipellis and represents a redefined /marasmiaceae. A third lineage includes the genera Cylindrobasidium, Flammulina, Gloiocephala, Physalacria, Strobilurus, Xerula and Marasmius sect. Epiphylli and represents /physalacriaceae. One new combination in Rhodocollybia and four new combinations in Mycetinis are proposed. A discussion of the taxonomic implications resulting from the phylogenetic reconstruction is presented.

Monograph of Marasmius, Gloiocephala, Palaeocephala and Setulipes in tropical Africa

Article

Jan 2007

Vladimír Antonín

New species of marasmioid genera (Basidiomycetes, Tricholomataceae) from tropical Africa III. Marasmius sect. Sicci

Article

Apr 2004

Vladimír Antonín

Twelve new tropical African taxa of Marasmius sect. Sicci (M. confertus var. tenuicystidiatus, M. confertus var. parvisporus, M. ferruginacies, M. ferruginoides, M. longistipitatus, M. pseudotorquescens, M. rammelooi, M. robertsii, M. rubro-stipitatus, M. striaepileus, M. tanougouensis, M. xeslocephaloides) are described and briefly discussed.

New species of marasmioid genera (Basidiomycetes, Tricholomataceae) from tropical Africa IV. Four new taxa of the genus Marasmius and one new combination

Article

Apr 2004

Vladimír Antonín

Four new African tropical taxa of the genus Marasmius, M. aurantiostipitatus, M. curreyi var. distantifolius, M. rotalis van latisporus (sect. Marasmius) and M. subalbidulus (sect. Hygrometrici) are described and one new combination, Setulipes congolensis, is proposed.

Monographic studies on South American Basidiomycetes, especially those of the east slope of the Andes and Brazil 2. The genus Marasmius in South America

Article

Jan 1965
SYDOWIA

R. Singer

Marasmieae (Basidiomycetes - Tricholomataceae)

Article

Jan 1976

R. Singer

New species of Marasmius (Basidiomycetes, Tricholomataceae) from tropical Africa - I. Sect. Epiphylli, Fusicystides, Globulares, Hygrometrici and Neosessiles

Article

Jan 2003

Vladimír Antonín

Twelve new taxa (11 species and 1 variety) of the genus Marasmius s. str. (Basidiomycetes, Tricholomataceae) from tropical Africa are published. Six species (M. albidocremeus, M. camerunensis, M. lacteoides, M. muramwyanensis, M. kigwenensis, M. tshopoensis) belong to sect. Globulares, two species (and one variety) to sect. Hygrometrici (M. minutoides, M. minutoides var. angustisporus, M. nyikae), and one species each to sect. Epiphylli (M. foliiphilus), sect. Fusicystides (M. longicystidiatus) and sect. Neosessiles (M. bururiensis). Detailed macroscopic and microscopic description, drawings and a short discussion is given for each species. These studies represent a part of a larger taxonomic project dealing with African Marasmii.

Biodiversity and phylogeny of Marasmius (Agaricales, Basidiomycota) from Madagascar

Abstract

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