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Mycology
Vol. 1, No. 2, June 2010, 130–153
ISSN 2150-1203 print/ISSN 2150-1211 online
© 2010 Mycological Society of China
DOI: 10.1080/21501203.2010.493531
http://www.informaworld.com
TMYC
Records and new species of Pluteus from Brazil based on morphological and molecular data
Mycology
Nelson Menolli Jr.a,b*, Tatiane Asaia and Marina Capelaria
aInstituto de Botânica, Núcleo de Pesquisa em Micologia, Caixa Postal 3005, 01031-970, São Paulo, SP, Brazil; bInstituto Federal de
Educação, Ciência e Tecnologia de São Paulo, Rua Pedro Vicente 625, 01109-010, São Paulo, SP, Brazil
(Received 31 March 2010)
Ten species of Pluteus are described and illustrated from specimens recently collected at three remnants of Atlantic forest in
São Paulo State, Southeast Brazil. Two new taxa are proposed: Pluteus aureovenatus and P. dominicanus var. hyalinus, as
well as a new status, P. sublaevigatus to P. chrysophlebius subsp. sublaevigatus. The occurrence of P. fuligineovenosus,
P. jamaicensis, and P. riberaltensis var. conquistensis represent the first records from Brazil. Pluteus albostipitatus, P. flu-
minensis, P. harrisii and P. xylophilus are also described and illustrated. In addition, a molecular study was performed
based on parsimony analyses of nLSU and ITS + 5.8S sequences of Brazilian and other Pluteus species. The generated phy-
logenetic trees revealed a well-supported clade with all Pluteus species. In the nLSU analyses, it was possible to identify a
clade with all species of section Pluteus and another including members of both sections Hispidoderma and Celluloderma.
In the ITS + 5.8S analyses, it was possible to identify three major clades, each of them corresponding to the majority repre-
sentatives of sections Celluloderma, Hispidoderma and Pluteus.
Keywords: Agaricales; Basidiomycota; ITS; nLSU; Phylogeny; Pluteaceae; South America; Taxonomy
Introduction
Pluteus Fr. is a large and widely distributed genus of presum-
ably saprophytic and non-ectomycorrhizal species (Singer
1986; Banerjee and Sundberg 1995). The genus as defined by
Singer (1986) is characterised by free lamellae, pinkish spore
print, absence of both annulus and volva, inamyloid basid-
iospores, cystidia metuloid or not and inverse hymenophoral
trama. However, recently Minnis et al. (2006) and Corriol
and Moreau (2007), supported by molecular and morphologi-
cal data, transferred to Pluteus two annulate species previ-
ously classified in the genus Chamaeota (W.G. Sm.) Earle.
Thus, the concept of Pluteus must be amended to also accom-
modate species with partial veil.
The infrageneric classification of Pluteus currently in
use subdivides the genus into three sections (Pluteus, Hispi-
doderma Fayod and Celluloderma Fayod) based on morpho-
logical features such as the structure of the pileipellis and
characteristics of the pleurocystidia (Singer 1958, 1986).
Ahymeniform layer of spheropedunculate cells characterise
the pileipellis of the section Celluloderma, while the other
two sections exhibit pileipellis with elongate, pilose or
hyphous elements. Members of the section Pluteus differ
from those of section Hispidoderma by the presence of met-
uloids and thin-walled pleurocystidia, respectively (Singer
1958, 1986). Singer (1956, 1958, 1986) also considered two
subsections in section Celluloderma: Mixtini Singer and
Eucellulodermini Singer, differed by a pileipellis composed
of uniform or heteromorphic elements, respectively.
Vellinga and Schreurs (1985) also proposed an infra-
generic classification for Pluteus that maintains the section
Pluteus according to Singer’s classification and introduces
the new section Villosi Schreurs & Vellinga to include the
species with pileipellis differentiated as a cutis combined
with thin-walled pleurocystidia. The section Celluloderma
also was maintained but divided into three subsections
according to the shape of the pileipellis cells. The two sub-
sections, Mixtini and Eucellulodermini, proposed by
Singer (1956, 1958, 1986), were maintained and a third sub-
section was proposed, Hispidodermini (Fayod) Vellinga &
Schreurs, which is characterized by a trichodermic
pileipellis or by a hymeniderm with cylindrical to fusiform
elements, grouping mostly members of section Hispido-
derma sensu Singer (Singer 1958, 1986).
The knowledge of Pluteus in Brazil started with
Hennings (1900) with the description of P. scruposus
Henn. from Mato Grosso State. Later, Hennings (1904a,
b) described P. cervinus var. griseoviridis Henn. from
São Paulo State and P. termitum Henn. from Amazonas
State. Most of these early species described from Brazil
were not preserved and may not be Pluteus, as men-
tioned by Saccardo and Trotter (1912), who recognised
that P. termitum was a possible synonym of Collybia
eurhiza (Berk.) Höhn., and by Singer (1958), who estab-
lished P. scruposus as a synonym of Oudemansiella
canarii (Jungh.) Höhn. and P. termitum as a probable
Lepiota sp.
*Corresponding author. Email: menollijr@yahoo.com.br
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Mycology 131
Rick (1907, 1919, 1930, 1938, 1961) also described
and recorded some problematic species. In total, 21
Pluteus taxa from Brazil (Rio Grande do Sul State),
including six species (P. cristatulus Rick, P. fibrillosus
Rick, P. leptonia Rick, P. sensitivus Rick, P. straminellus
Rick, and P. velatus Rick) and two varieties [P. exiguus var.
venosus Rick and P. nanus var. podospileus (Sacc. & Cub.)
Rick] with Brazilian types. However, Singer (1953, 1958)
considered several of Rick’s collections as misidentifica-
tions, nomina dubia, synonyms, or species of other genera.
Another 10 Pluteus taxa described as new from Brazilian
materials were described by Bresadola (1920) and Singer
(1953, 1956, 1958, 1973, 1989): P. amazonicus Singer,
P. cervinus var. brasiliensis Bres., P. fluminensis Singer,
P. hylaeicola Singer, P. melanopotamicus Singer, P. para
ensis Singer, P. riograndensis Singer, P. subfibrillosus
Singer, P. umbrinoalbidus Singer, and P. varzeicola
Singer. Recently, Menolli and Capelari (2010) described
two new species (P. densifibrillosus Menolli & Capelari
and P. puttemansii Menolli & Capelari) from materials
collected in the Atlantic forest in São Paulo State.
Grandi et al. (1984), Singer (1984, 1989), Raithelhuber
(1991), Stijve and Meijer (1993), Stijve (1995), Pegler
(1997), Meijer (2001, 2006, 2008), and Drechsler-Santos
et al. (2007) also contributed to increasing the knowledge
about Pluteus in Brazil. However, these works are just lists
and do not include complete descriptions of the species.
Wartchow et al. (2004, 2006) recorded and provided a
complete description of P. albostipitatus (Dennis) Singer,
P. aquosus Singer, P. beniensis Singer, P. globiger Singer,
P. nigrolineatus Murrill, and P. thomsonii (Berk. & Broome)
Dennis for Rio Grande do Sul State. However, Rodríguez
et al. (2008) considered that the European species P. thom-
sonii recorded by Wartchow et al. (2004) is probably
P. neotropicalis Rodr.-Alcánt., which was recently
described (Rodríguez et al. 2008) based on Mexican mate-
rial previously misidentified as P. thomsonii (Rodríguez
and Guzmán-Dávalos 1999).
Moreover, Meijer (2008) and Menolli and Capelari
(2010) recently provided complete descriptions of one
species [P. xylophilus (Speg.) Singer] from Paraná State
and four [P. densifibrillosus, P. longistriatus (Peck) Peck,
P. puttemansii and P. umbrinoalbidus] from São Paulo
State, respectively. Meijer (2008) also mentioned that 34
species of Pluteus occur in Paraná State, but he did not
mention the names of them. In addition, in his major
papers including Pluteus (Stijve and Meijer 1993, Meijer
2001, 2006, 2008), it was only possible to localize about
24 Pluteus taxa. According to Meijer (2009, personal com-
munication), the remaining 10 taxa mentioned are differ-
ent from any other known Pluteus species and so far they
have not been formally described.
All records of Pluteus from Brazil are very confusing
and incomplete due to poor descriptions, synonyms, misi-
dentifications, the lack of preserved material, and lists
lacking descriptions. So far, 70 epithets have been recorded,
distributed over nine States (Amazonas, Bahia, Mato
Grosso, Pará, Paraná, Rio de Janeiro, Rio Grande do Sul,
Rondônia, and São Paulo), with 23 of them represented by
Brazilian types. However, only 23 are known for certain or
are at least cited with a complete description. The remain-
ders represent misidentifications, synonyms, species of
other genera, unpreserved specimens, or are presented in
lists.
Molecular studies involving Pluteus are restricted and
they do not include South American species. Moncalvo
et al. (2002) and Matheny et al. (2006) conducted a broad
study of Agaricales, wherein the Pluteus clade was super-
ficially discussed. Later, Minnis et al. (2006) added a
Chamaeota sequence to the nLSU analysis done by Mon-
calvo et al. (2002) and proposed a new combination in
Pluteus. Malysheva et al. (2009) and Rodríguez et al.
(2009) performed analyses of the ITS +5.8S region with
the placement of new species or new records from Russia
and Mexico, respectively.
Here, we report, describe and illustrate 10 species of
Pluteus recently collected in the remnants of the Atlantic
forest in Southeast Brazil. We proposed two new taxa, a
new status and three new records for Brazil. Moreover, for
the first time in Pluteus, molecular analyses with sequences
of South American species based on nLSU and ITS + 5.8S
data are presented.
Material and methods
Sampling
The studied materials were collected at the Parque Estadual
da Cantareira, São Paulo city; Parque Estadual das Fontes do
Ipiranga, São Paulo city; and Reserva Biológica de Paranapi-
acaba, Santo André city, all of them are remnants of the
Atlantic forest in São Paulo State, Southeast Brazil. When
necessary, materials from other localities were examined.
Morphological study
The macroscopic descriptions and illustrations of basidi-
omata were based on fresh material. Colour terms are
according to Küppers (1979). For microscopic analyses,
the dried materials were rehydrated in 70% ethanol fol-
lowed by 5% KOH or Melzer’s reagent. All microscopic
illustrations were made with the aid of a drawing tube. The
notation “[a/b/c]” at the beginning of a set of basid-
iospores data is to be read as “a basidiospores were meas-
ured from b basidiomata taken from c collections”. The
basidiospores were measured in lateral view, always 20
basidiospores from each basidioma, and the terms denot-
ing their shapes were according to Bas (1969). Q repre-
sents the range of the length/width quotient for all the
measured spores, Qm represents the average of all
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132 N. Menolli Jr. et al.
computed Q values for all the measured basidiospores and
Lm (Wm) represents the average of all lengths (widths) of
the measured basidiospores. The terms used to character-
ize the pleurocystidia as Cervinus- and Magnus-types are
according to Singer (1958) and Banerjee and Sundberg
(1995). For the pileipellis cells, the length of the pedicel
was included in the measurement of the sphaeropeduncu-
late cells, and its length is also reported separately. A sep-
arate measurement is given when an apical projection was
present. The specimens and the voucher of the sequences
are deposited at the herbarium SP. Generic and infrage-
neric concepts follow Singer (1986); for comparison, the
classification proposed by Vellinga and Schreurs (1985)
was also considered.
Molecular study
DNA extraction
Procedures for DNA extraction were done according to an
adapted protocol by Zolan and Pukkila (1986) using
lyophilised basidiomata previously ground to a fine pow-
der in liquid nitrogen. The sample was resuspended in 50 µl
of TE, incubated at 37 °C for 30 min after the addition of
RNAse (0.01 mg/µl), and stored at −20 °C.
PCR amplification and DNA sequencing
The ITS + 5.8S region was amplified using the primer set
ITS1-F and ITS4 (Vilgalys and Hester 1990; White et al.
1990; Gardes and Bruns 1993), and the nLSU rDNA was
amplified using the primer set LR0R and LR5 (Moncalvo
et al. 2000). The PCR reaction, containing 2.0 U of Plati-
num® Taq DNA Polymerase (Invitrogen), 0.2 mM of each
dNTP, 1.5 mM of MgCl2, and 0.2 µM of each primer of
the selected region in a total of 100 µl, was performed in a
Eppendorf thermocycler, using the following program: 94 °C
for 5 min, 40 cycles at 94 °C for 40 s, 55 °C for 30 s, and
72 °C for 60 s, and then 72 °C for 5 min. Amplification
products were purified using the PureLink PCR Purifica-
tion Kit (Invitrogen). DNA sequencing reactions were per-
formed with the DYEnamic ET Dye Terminator Kit in a
MegaBACE 1000 DNA sequencer (GE Healthcare)
according to the manufacturer’s instructions. The samples
were sequenced in both directions. The sequences were
deposited in GenBank (Table 1).
Data analysis
To elucidate the relationship of Pluteus species found in
São Paulo State, Brazil, we conducted phylogenetic analy-
ses using nLSU and ITS + 5.8S sequences determined in
this study with sequences available in GenBank (Table 1).
The sequences were automatically aligned using the
Clustal W version (Thompson et al. 1994) in the BioEdit
version 7.0.5.3 (Hall 1999). Parsimony analysis was per-
formed with PAUP version 4.0b10 (Swofford 2003). Par-
simony trees were obtained by heuristic searches with
simple sequence addition in 1000 replicates, employing
tree-bisection-reconnection (TBR) branch-swapping algo-
rithm. Characters from the extreme 5′ and 3′ ends of the
sequences were deleted from all taxa to obtain individual
datasets that had identical start and end positions, gaps
were treated as missing, all characters were unordered and
equally weighted, and multistate taxa was interpreted as
uncertainty. Starting trees were obtained via stepwise
addition, with one tree held at each step during stepwise
addition and the steepest descent option not in effect. Also,
the initial MaxTrees were set to auto-increase, branches of
zero length were collapsed (creating polytomies), and
MulTrees options were in effect.
Branch and branch node supports were determined
using 1000 BS replicates. Estimated levels of homoplasy
and phylogenetic signal (retention and consistency
indexes) were determined. The generated trees were
rooted to Limacella illinita (Fr.) Maire, Auricularia poly-
tricha (Mont.) Sacc. and Volvariella volvacea (Bull.)
Singer as the outgroup taxa for nLSU analysis following
Minnis et al. (2006), and V. gloiocephala (DC.) Boekhout
& Enderle and V. volvacea for the ITS +5.8S analysis
(Table 1).
Results and discussion
Sequence data from the nLSU region
In the nLSU analysis, 29 sequences of 26 taxa plus three
outgroups were aligned. The alignment dataset consisted of
1304 characters, including gaps. Prior to analysis, 515 char-
acters from 5′ and 3′ ends of the sequences were excluded.
Of the 789 characters included in the analysis, 590 charac-
ters were constant, 109 variable characters were parsimony-
uninformative and 90 were parsimony-informative.
The heuristic searches retained 24 equally most parsi-
monious trees resulting in a bootstrap 50% majority-rule
consensus tree with the following scores: tree length = 380
steps, CI = 0.592, RI = 0.703, RC = 0.416, and HI = 0.408.
The MP tree generated from the nLSU sequence data
(Figure 1) revealed a well-supported clade (89% BS)
including all Pluteus species. Two clades can be distin-
guished with low BS support: one clade (58% BS) cluster-
ing species that belong to section Pluteus, and the other
revealing a polytomy including members of sections
Hispidoderma and Celluloderma (51% BS).
Sequence data from the ITS + 5.8S region
In the ITS + 5.8S analysis, 48 sequences of 31 taxa plus
three outgroups were aligned. The alignment dataset con-
sisted of 1397 characters, including gaps and portions of
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Mycology 133
Table 1. Collection data and GenBank accession number of the Pluteus and outgroups taxa analysed.
Species ITS + 5.8S LSU Voucher/ Strain Country Reference
Auricularia polytricha –AF261554 HN4076 – Moncalvo et al. 2002
Limacella illinita – AF261439 VT8.9.96 – Moncalvo et al. 2002
Pluteus admirabilis – AF261577 DAOM193532 – Moncalvo et al. 2002
– AF261578 DAOM197226 – Moncalvo et al. 2002
P. albostipitatus FJ816656 – NMJ128 Brazil This paper
FJ816661 FJ816647 FK782 Brazil This paper
P. albostipitatus var. poliobasis FJ375244 Isolate 113 Mexico Rodríguez et al. 2009
P. atromarginatus DQ494687 – HKAS 31573 China Matheny et al. 2006
EF530926 – UBC F16254 Canada Denis et al. Not published
FJ774075 LE246081 – Malysheva et al. 2009
– AF261572 JB97/14 – Moncalvo et al. 2002
P. aurantiorugosus – AF261579 DAOM197369 – Moncalvo et al. 2002
FJ774077 – LE216873 Russia Malysheva et al. 2009
FJ375248 – Isolate 160 Spain Rodríguez et al. 2009
FJ375249 – Isolate 216 Mexico Rodríguez et al. 2009
P. aureovenatus FJ816663 FJ816649 FK1045 Brazil This paper
P. brunneoradiatus – AF261567 JB97/3 – Moncalvo et al. 2002
P. cervinus – AF261570 JB97/19 – Moncalvo et al. 2002
FJ774088 – LE217575 Russia Malysheva et al. 2009
FJ375241 – Isolate 6 Mexico Rodríguez et al. 2009
P. cervinus var. cervinus EU486448 – UBC F16293 Canada Denis and Berbee. Not published
P. chrysophlebius – AF261581 DAOM190194 – Moncalvo et al. 2002
P. diverticulatus FJ375247 – Isolate 242 France Rodríguez et al. 2009
P. dominicanus var. hyalinus FJ816665 FJ816651 FK1058 Brazil This paper
P. ephebeus – AF261574 JB97/23 – Moncalvo et al. 2002
FJ774074 – LE227554 Russia Malysheva et al. 2009
P. exiguus FJ774083 – LE226543 Russia Malysheva et al. 2009
P. fenzlii FJ774082 – LE246083 Russia Malysheva et al. 2009
P. fluminensis FJ816655 – NMJ127 Brazil This paper
FJ816664 FJ816650 FK1046 Brazil This paper
P. fuligineovenosus FJ816662 – FK826 Brazil This paper
P. granulatus FJ774086 – LE212990 Russia Malysheva et al. 2009
P. harrisii FJ816654 FJ816644 NMJ122 Brazil This paper
FJ816666 FJ816652 FK1066 Brazil This paper
P. horakianus FJ375250 – Isolate 58 Mexico Rodríguez et al. 2009
FJ375251 – Isolate 60 Mexico Rodríguez et al. 2009
P. jamaicensis FJ816657 – NMJ130 Brazil This paper
P. leoninus FJ774079 – LE212090 Russia Malysheva et al. 2009
P. mammillatus –DQ451549ASM 7916 – Minnis et al. 2006
P. nanus FJ774081 – LE213093 Russia Malysheva et al. 2009
P. nigrolineatus FJ375245 – Isolate 114 Mexico Rodríguez et al. 2009
P. pallidus* – AF261569 JB90/27 – Moncalvo et al. 2002
P. pellitus – AF261571 JB93/3 – Moncalvo et al. 2002
FJ774078 – LE217548 Russia Malysheva et al. 2009
FJ375243 – Isolate 222 Mexico Rodríguez et al. 2009
P. petasatus – AF042611 JB91/21 – Moncalvo et al. 2000
AF085495 – CBS441.85 Czech Republic Lim and Jung 1998
FJ375242 – Isolate 100 Mexico Rodríguez et al. 2009
P. plautus FJ774076 – LE213024 Russia Malysheva et al. 2009
P. podospileus FJ774085 – LE213015 Russia Malysheva et al. 2009
P. pouzarianus – AF261568 JB94/26 – Moncalvo et al. 2002
P. primus – AF042610 JB94/24 – Moncalvo et al. 2000
P. riberaltensis var. conquistensis – FJ816648 FK1043 Brazil This paper
P. romellii – AF261575 JB97/26 – Moncalvo et al. 2002
AY854065 – ECV 3201 USA Matheny et al. 2006
FJ774073 – LE217944 Russia Malysheva et al. 2009
FJ375246 – Isolate 119 Mexico Rodríguez et al. 2009
P. salicinus – AF261573 JB97/6 – Moncalvo et al. 2002
FJ774087 – LE215427 – Malysheva et al. 2009
(Continued)
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134 N. Menolli Jr. et al.
nSSU and/or nLSU. Prior to analysis, 898 characters from
the 5′ and 3′ ends of the sequences were excluded. Out of
the 499 characters included in the analysis, 152 characters
were constant, 103 variable characters were parsimony-
uninformative and 244 were parsimony informative.
The heuristic searches retained two equally most parsi-
monious trees resulting in a bootstrap 50% majority-rule
consensus tree with the following scores: tree length = 1425
steps, CI = 0.451, RI = 0.711, RC = 0.321, and HI = 0.549.
The MP tree generated from the ITS +5.8S sequence
data also revealed a well-supported clade (82% BS),
including all Pluteus species, divided into three major
clades (Figure 2). Species from sections Pluteus and
Celluloderma are clustered into two sister-clades with 85
and 73% BS support, respectively. In an external branch
of these clades there is a third clade that groups the
majority species from section Hispidoderma with 64%
BS support.
Phylogenetic analyses from the rDNA sequence data
It was possible to confirm the monophyletic origin of Plu-
teus from both analyses (nLSU and ITS + 5.8S), as also
shown by Moncalvo et al. (2002) and Matheny et al.
(2006). Moreover, especially in the ITS + 5.8S (Figure 2)
it was possible to recognize three major clades, each one
corresponding to the sections proposed by Singer (1958,
1986).
All species from section Pluteus clustered in the same
clade (Figures 1 and 2), forming a well-supported group in
accordance with the infrageneric classification of Singer
(1958, 1986).
The clustering of species from section Celluloderma in
one clade was observed in both analyses (Figures 1 and 2).
The nLSU analysis for section Celluloderma (Figure 1)
showed a polytomy within a clade with low BS support
(51%) including members of section Hispidoderma, such
as P. umbrosus (Pers.) P. Kumm. and P. riberaltensis var.
conquistensis Singer, and also including members of the
section Villosi, viz. P. ephebeus (Fr.) Gillet, and species
with partial veil traditionally classified in the genus
Chamaeota, viz. P. mammillatus (Longyear) Minnis,
Sundb. & Methven. The relationship between members of
section Celluloderma and section Hispidoderma was also
demonstrated by Moncalvo et al. (2002) in a well-supported
clade (92% BS).
In the ITS + 5.8S analysis (Figure 2), the majority
members of the section Celluloderma are clustered in a
branch connected to section Pluteus clade, while the mem-
bers of the section Hispidoderma are mostly in an external
branch from the clades with members of the sections
Pluteus and Celluloderma. Some species formerly classi-
fied in the section Hispidoderma, such as P. ephebeus,
P. exiguus (Pat.) Sacc. and P. fenzlii (Schulzer) Corriol &
P.-A. Moreau (an annulated species), formed a clade with
all species of section Celluloderma.
For the members of the section Hispidoderma (includ-
ing Villosi and Chamaeota members), it will be necessary
to perform a detailed study to elucidate the evolutionary
history of the group and to find characters that are shared
with other members of section Celluloderma (or also of
section Pluteus, if the position of P. albostipitatus and P.
nigrolineatus are considered, as discussed below). Until
now, it has not been possible to recognise the monophyly
of section Hispidoderma as proposed by Singer (1958,
1986), and perhaps this represents an artificial group
delimited by morphological characters. However, one
should not disregard the possibility that the section Hispi-
doderma is a subsection of Celluloderma, as proposed by
Vellinga and Schreurs (1985).
Table 1. (Continued)
Species ITS + 5.8S LSU Voucher/ Strain Country Reference
P. semibulbosus FJ774080 – LE227534 Russia Malysheva et al. 2009
P. sublaevigatus FJ816667 FJ816653 FK1085 Brazil This paper
P. thomsonii FJ774084 – LE234787 Russia Malysheva et al. 2009
FJ375252 – Isolate 85 France Rodríguez et al. 2009
FJ375253 – Isolate 155 Spain Rodríguez et al. 2009
P. umbrosus – AF261580 DAOM197235 – Moncalvo et al. 2002
P. xylophilus FJ816659 FJ816645 NMJ143 Brazil This paper
FJ816660 FJ816646 NMJ147 Brazil This paper
Pluteus sp. – AF261576 JMCR.124 – Moncalvo et al. 2002
–AF042612JM96/28 –Moncalvo et al. 2000
FJ816658 – NMJ132 Brazil This paper
Volvariella gloiocephala FJ375254 – – Rodríguez et al. 2009
DQ494701 – PMB2272 USA Matheny et al. 2006
Volvariella volvacea – AF261531 Jmleg.SRL – Moncalvo et al. 2002
AY636050 – OE-140 – Singh et al. 2003
*Voucher material misidentified as P. pallidus (LAU!), see discussion in the text.
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Mycology 135
In the ITS +5.8S analysis (Figure 2), it is possible to
observe the presence of two species currently classified in
section Hispidoderma (P. albostipitatus and P. nigrolineatus)
but positioned in the section Pluteus clade. Rodríguez et al.
(2009) had already demonstrated the affinities between sec-
tion Pluteus and section Hispidoderma (89% BS) due to the
presence of these species in the same clade with other repre-
sentative species of section Pluteus. Morphological charac-
ters, such as the presence of a slightly thick-walled
pleurocystidia and sometimes with poorly developed prongs
and/or a repent epicutis, could be used to make assumptions
about the position of P. albostipitatus and P. nigrolineatus
in the section Pluteus (Figure 2) and to consider that these
species probably represent a transition between sections
Hispidoderma and Pluteus (see discussion under P. albostipi-
tatus). However, it is not possible to confirm this assumption
with ITS + 5.8S analysis because the internal structure of the
section Pluteus is unresolved. A molecular study is necessary
Figure 1. Phylogenetic tree generated by MP analysis of partial LSU sequences. The tree is rooted using Auricularia polytricha, Lima-
cella illinita and Volvariella volvacea. The BS numbers are shown before the nodels. *Voucher material misidentified as P. pallidus, see
discussion in the text.
Limacella illinita AF261439
Volvariella volvacea AF261531
Auricularia polytricha AF261554
Pluteus atromarginatus AF261572
P. pellitus AF261571
P. salicinus AF261573
P. albostipitatus FJ816647
P. harrisii FJ816644
P. harrisii FJ816652
P. petasatus AF042611
P. xylophilus FJ816646
P. xylophilus FJ816645
P. pouzarianus AF261568
P. primus AF042610
P. brunneoradiatus AF261567
P. pallidus AF261569*
P. cervinus AF261570
P. umbrosus AF261580
P. dominicanus var. hyalinus FJ816651
P. riberaltensis var. conquistensis FJ816648
P. mammillatus DQ451549
P. ephebeus AF261574
P. admirabilis AF261577
P. admirabilis AF261578
P. chrysophlebius AF261581
P. fluminensis FJ816650
P. aurantiorugosus AF261579
P. romellii AF261575
P. sublaevigatus FJ816653
P. aureovenatus FJ816649
Pluteus sp. AF042612
Pluteus sp. AF261576
87
99
82
65
71
52
56
66
85
99
57 60
60
81
51
89
58
Section Pluteus clade
Section Celluloderma +
Hispidoderma clade
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136 N. Menolli Jr. et al.
which would show a well-resolved topology for section Plu-
teus, placing these species as sister group of all other species
in section Pluteus, to confirm this phylogenetic relation
(Justo 2009, personal communication).
In both analyses, it is possible to recognize some species
or species-complexes that need revision to confirm the
scope and/or the monophyly of the section. The nLSU ana-
lysis showed the presence of P. pallidus Homola, a species
of section Celluloderma, in the section Pluteus clade (Fig-
ure 1). This fact was also observed by Moncalvo et al.
(2002) but the authors did not comment the position of
P. pallidus and considered the section Pluteus as a well-
supported group. However, a re-examination of the voucher
material, demonstrated that it actually represents a species
of section Pluteus because it has the typical horned and met-
uloidal pleurocystidia. This was confirmed by Bonnard
Figure 2. Phylogenetic tree generated by MP analysis of ITS + 5.8S region. The tree is rooted using Volvariella gloiocephala and
V. volvacea. The BS numbers are shown before the nodels. *Voucher materials that need revision to confirm the identification, see dis-
cussion in the text.
Volvariella volvacea AY636050
Volvariella gloiocephala FJ375254
Volvariella gloiocephala DQ494701
P. atromarginatus EF530926
P. atromarginatus DQ494687
P. atromarginatus FJ774075
P. nigrolineatus FJ375245
P. pellitus FJ774078
P. pellitus FJ375243*
P. salicinus FJ774087
P. albostipitatus var. poliobasis FJ375244
P. albostipitatus FJ816661
P. albostipitatus FJ816656
P. harrisii FJ816654
P. harrisii FJ816666
P. petasatus AF085495
P. petasatus FJ375241*
P. xylophilus FJ816659
P. xylophilus FJ816660
P. cervinus FJ375241*
P. cervinus FJ774088
P. cervinus var. cervinus EU486448
P. fluminensis FJ816655
P. fluminensis FJ816664
P. podospileus FJ774085
P. jamaicensis FJ816657
P. fuligineovenosus FJ816662
P. aurantiorugosus FJ375248
P. aurantiorugosus FJ375249
P. aurantiorugosus FJ774077*
P. semibulbosus FJ774080
P. plautus FJ774076
P. granulatus FJ774086
P. horakianus FJ375251
P. horakianus FJ375250
P. nanus FJ774081
P. fenzlii FJ774082
P. ephebeus FJ774074
P. exiguus FJ774083
P. thomsonii FJ375253
P. thomsonii FJ375252
P. thomsonii FJ774084
P. leoninus FJ774079
P. diverticulatus FJ375247
P. dominicanus var. hyalinus FJ816665
P. romellii FJ774073*
P. romellii FJ375246*
P. romellii AY854065*
P. sublaevigatus FJ816667
P. aureovenatus FJ816663
Pluteus sp. FJ816658
84
86
Section Pluteus clade
Section Celluloderma clade
Section Hispidoderma clade
79 90
90
100
60
91
99
68
100
51
90
56
73
85
69
82
93
54
61
53
91
91
99
70
54
99
76
94
96
52
100
64
97
99
54
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Mycology 137
(2001), who cited the voucher material (JB 90/27) in the
original description of P. albineus Bonnard.
In the clade including members of the Cervinus group
(section Pluteus clade), based on the ITS + 5.8S analysis
(Figure 2), there is also some species for which identifica-
tion must be revised. Two sequences of P. cervinus
(Schaeff.) P. Kumm., representative (EU486448 and
FJ774088) of different countries (USA and Russia, respec-
tively), clustered in a well-supported clade (84% BS),
while the Mexican material of P. cervinus (FJ375241) also
clustered in a well-supported clade (90% BS), but with
sequences of P. pellitus (Pers.) P. Kumm. and P. petasatus
(Fr.) Gillet, both from Mexico (FJ375243 and FJ375242,
respectively), and with another sequence of P. petasatus
from the Czech Republic (AF085495, origin of voucher
strain CBS441.85 obtained at http://www.cbs.knaw.nl/
databases/). Rodríguez et al. (2009) already reported
molecular affinities (96% BS) between their sequences of
P. cervinus, P. pellitus and P. petasatus, and also men-
tioned macroscopic similarities between them. Previous
morphological relations between P. pellitus and P. petasa-
tus were indicated by Vellinga and Schreurs (1985) and
Rodríguez and Guzmán-Dávalos (2000). The sequence of
P. pellitus from Russia (FJ774078), which probably repre-
sents this species, formed a separate branch from the Mex-
ican sequence of P. pellitus. The sequences of P. petasatus
clustered in a well-supported clade (90% BS), despite their
origins being from different regions of the world (Mexico
and Czech Republic).
Considering these data, it is probable that the Mexican
materials forming the P. cervinus/pellitus/petasatus com-
plex could represent only one species, most likely P.
petasatus. Thus, the collections of all P. cervinus, P. pelli-
tus and P. petasatus present in the analysis must be revised
because sequences from different collections around the
world and with the same identification are not grouped
together or species with different names clustered in the
same clade. Moreover, many species of the Cervinus
group most likely represent a complex of species with
morphological similarities, which means that different
worldwide species are erroneously named with the same
epithet, e.g. P. cervinus.
For the Celluloderma and Hispidoderma clades (Figure 2)
some collections of P. romellii (Britzelm.) Sacc. and
P. aurantiorugosus (Trog) Sacc. also need revision to
confirm their GenBank identification. The sequences of
P. romellii (FJ774073, FJ375246 and AY854065) from
different countries (Russia, Mexico and USA, respec-
tively) were grouped in the same clade with P. auranti-
orugosus, P. aureovenatus Menolli & Capelari and
P. sublaevigatus (Singer) Menolli & Capelari (both of
the latter herein described) but in different branches. The
three sequences of P. romelli need revision, especially
the material from the USA (California), which segre-
gated from the other P. romelli sequences and clustered,
despite the low support (53% BS), with the morphologi-
cally different P. aureovenatus and P. sublaevigatus (see
discussion under P. aureovenatus and P. sublaevigatus).
The Russian collection of P. aurantiorugosus
(FJ774077) also needs revision because it grouped with
100% BS with P. leoninus (Schaeff.) P. Kumm. in the sec-
tion Hispidoderma clade (Figure 2). Both species, despite
some ecological and macromorphological similarities
pointed out by Malysheva et al. (2009), have different
pileipellis structures and are consequently traditionally
classified in different sections, Hispidoderma and Cellulo-
derma, respectively. Malysheva et al. (2009) obtained the
same support (100% BS) for these species groupings.
However, considering this high support and that the other
sequences of P. aurantiorugosus (FJ375249 and FJ375246)
from different countries clustered in another clade with all
species of the section Celluloderma, it is likely that the
Russian collection named as P. aurantiorugosus in the
GenBank actually represents P. leoninus.
The presence of P. semibulbosus (Lasch) Quél. in the
section Hispidoderma clade and the clustering with
P. plautus (Weinm.) Gillet (52% BS) should also be
regarded with caution, because Singer (1986) considered
the first as a member of section Hispidoderma, while
Orton (1986) classified it in the section Celluloderma.
These data are in accordance with Malysheva et al.
(2009), whom already reported molecular affinities
(96% BS) between them, and with Vellinga and Schreurs
(1985), whom considered P. semibulbosus sensu Singer
as synonymous of P. plautus, a member of section
Hispidoderma. In addition, Vellinga and Schreurs (1985)
considered P. semibulbosus sensu Orton as a P. inquili-
nus Romagn., which in fact is a member of section
Celluloderma.
Taxonomy
Pluteus albostipitatus (Dennis) Singer, Lloydia 21: 240
(1959) (Figure 3).
≡ Pluteus spilopus var. albostipitatus Dennis, Bull. Soc.
Mycol. Fr. 69: 195, 1953.
Pileus 34–37 mm diam, plane, slightly to deeply
umbonate, light brown to greyish brown (N60A50M40 to
N80A40M50), darker with coffee shades at the centre
(N90A50M60), covered by appressedly and radially arranged
fibrils, more concentrated at the centre and dissolving out-
side of the central disc, finely pruinose at the centre, margin
sulcate-striate. Lamellae free, white then pinkish, crowded,
with lamellulae. Stipe 35–54 × 2(apex)–4(base) mm,
slightly tapering towards the apex, central, occasionally
flexuous, white to pale cream, with greyish to slightly sil-
ver base, slightly longitudinally striate, with scanty basal
mycelium. Odour, taste, and flesh colour not recorded.
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138 N. Menolli Jr. et al.
Basidiospores [40/2/2] (6.2–)7.5–10.0(−11.2) × 6.2–7.5
(−8.7) µm [Q = 1.00–1.29(−1.40); Qm = 1.14; Lm = 8.4
µm; Wm = 7.4 µm], globose to broadly ellipsoid, occa-
sionally subglobose or ellipsoid, widely variable in form,
especially among different collections, inamyloid, hyaline,
smooth, thick-walled, sometimes guttulate. Basidia (20–)
25–31(−35) × 8.7–12.5 µm, versiform to clavate, thin-
walled, four-spored, sometimes with small scattered guttules.
Pleurocystidia (39–)50–83(−92) × (11.2–)15.0–24 µm,
slightly ventricose with a rounded to subcapitate apex or
slightly strangulated at the apex, thin- to relatively thick-
walled, sometimes confused with a metuloid with a evenly
thickened wall up to 1 µm wide, colourless, hyaline and
moderately numerous. In some cases a second and rare
type of pleurocystidia is observed, with two short lateral
prongs, almost similar in size to the most common type, but
rarely shorter in length with approximately 27 µm high.
Cheilocystidia (25–)31–52(−55) × (6.2–)8.7–13.7(−15.0)
µm, clavate, rarely slightly ventricose, sometimes with a
well developed pedicel at the base, colourless, hyaline,
thin-walled, abundant. Lamellar edge sterile. Lamellar
trama inverse, up to 62 µm wide, composed of thin or
slightly thick-walled hyphae, 2.5–10.0(−12.5) µm diam,
hyaline. Pileus context undifferentiated, thin, approxi-
mately 38 µm thick, composed of thin- or slightly thick-
walled hyphae, 3.7–10(−18.7) µm diam, hyaline. Pileipel-
lis a repent epicutis up to 62 µm thick, composed of thin or
slightly thick-walled hyphae, 2.5–6.2 µm diam, elongated,
septate, with brown vacuolar content. Clamp connections
absent in all parts examined.
Habitat and habit: Solitary, on wood.
Material examined: BRAZIL, São Paulo State, São
Paulo city, Parque Estadual da Cantareira, Núcleo Engor-
dador, 24 Apr. 2007, Menolli Jr. et al. NMJ128 (SP);
23 Oct. 2006, F. Karstedt & M. Capelari FK782 (SP).
Comments: Pluteus albostipitatus was described from
Trinidad (Dennis 1953 as P. spilopus var. albostipitatus
Dennis), and since then, it was also reported for several
countries: Argentina (Singer 1956, 1958, Horak 1964),
Bolivia (Singer 1958), Martinique (Pegler 1983), Galapa-
gos Islands (Reid et al. 1981) and the French Guyana
(Courtecuisse 1991). For Brazil, this species was already
recorded from Paraná (Stijve and Meijer 1993 as P. cf.
albostipitatus; Meijer 2001, 2006), Rio Grande do Sul
(Wartchow et al. 2006) and São Paulo States (Pegler 1997).
The two materials herein studied differ themselves in
basidiospore, pleuro- and cheilocystidia shapes (Figure 3).
The collection Menolli Jr. et al. NMJ128 has predomi-
nantly globose basidiospores, pleurocystidia with evenly
thickened walls up to 1.0 µm and rarely with two subacute
lateral prongs, and cheilocystidia usually clavate and with-
out a long pedicel at the base. On the other hand, the col-
lection F. Karstedt & M. Capelari FK782 has usually
slightly ellipsoid basidiospores, pleurocystidia that some-
times have two rounded lateral prongs and with thin to
slightly thickened walls but that are not distinctly thick,
and cheilocystidia frequently with a well developed pedi-
cel at the base. These morphological differences observed
in our two collections probably represent intraspecific var-
iations, and it will be necessary to study more materials to
determine the frequency of these characters and to or not
attribute a varietal status to them.
Pluteus albostipitatus is probably a taxon with wide
micromorphological variations, especially regarding the
basidiospore, pleuro- and cheilocystidia shapes. Singer
(1958) observed a variation in the basidiospore shape from
subglobose to broad ellipsoid, but never globose, and
Figure 3. Pluteus albostipitatus. A. Basidioma (Menolli Jr.
et al. NMJ128). B. Basidiospores; B1. F. Karstedt & M. Capelari
FK782; B2. Menolli Jr. et al. NMJ128. C. Basidia (Menolli Jr.
et al. NMJ128). D. Pleurocystidia; D1. F. Karstedt & M. Cape-
lari FK782; D2. Menolli Jr. et al. NMJ128. E. Cheilocystidia; E1.
F. Karstedt & M. Capelari FK782; E2. Menolli Jr. et al.
NMJ128. Bars (A) = 1 cm; (B–E) = 10 µm.
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Mycology 139
recorded the presence of short obtuse or subacute small
prongs in the pleurocystidia, in accordance with our mate-
rial, except for the absence of globose basidiospores.
Pegler (1983) also observed a variation in basidiospore
shape from subglobose to ellipsoid and recorded the pres-
ence of a narrow pedicelate base in the cheilocystidia, but
observed pleurocystidia without prongs. Horak (1964) and
Courtecuisse (1991) described pleurocystidia with rounded
or truncate apex, without mention of the presence or
absence of prongs and cheilocystidia with a long basal
pedicel. However, Horak (1964) recorded basidiospores as
round-oval to almost globose as shown in the illustrations
presented, like those observed in our collections. Courte-
cuisse (1991) recorded small basidiospores (5.5–6.5 ×
4.5–5.2 µm), but had performed their measurements in an
immature material.
Moreover, the materials herein analysed have minor
differences when compared to the Brazilian collections
studied by Wartchow et al. (2006). They recorded
P. albostipitatus with a larger pileus (50–55 mm) and
basidiospores varying from subglobose to broad ellipsoid,
but never globose or ellipsoid, as observed in our collections.
The pleuro- and cheilocystidia are also moderately longer
[56.7–97.6(−113) × 12–24 µm and 35.2–71.2 × 8–18.4 µm,
respectively], and for the pleurocystidia, there is no men-
tion of the two short lateral prongs and either the wall is
slightly thickened and not relatively thick (nearly to metu-
loid), as observed in our materials. Wartchow et al. (2006)
did not describe, but illustrated a different shape of pleuro-
cystidia, with round and small several finger-like protu-
berances at the apex.
Singer (1973) described P. albostipitatus var. poliobasis
from Mexico (Veracruz State), which differs from the type
variety only by the pearl-gray colour (griseo) of the stipe
base. Regarding this variety characterization, Singer
(1956, 1958) described the stipe of P. albostipitatus as
“white, towards the base mostly pale sordid greyish” or
“white, finally becoming somewhat silvery-white to
glassy-cinereous”, respectively. The stipe colour of our
material resembles the descriptions of Singer (1956,
1958), and the stipe base of P. albostipitatus var. poliobasis
probably has a darker grayish shade, as mentioned by
Rodríguez and Guzmán-Dávalos (2000, 2001) from mate-
rial recollected in Jalisco State, Mexico.
Pluteus albostipitatus is close to P. melanopotamicus
Singer described from Amazonas State, Brazil (Singer
1989). The latter species is macroscopically the same as
P. albostipitatus, and according to the protologue, the
main differences between them are the dimorphic basid-
iospores of P. melanopotamicus, which for us is irrelevant,
because type 2 are only 0.5 µm longer than type 1 and
there can be a gradual variation in size as observed in
P. albostipitatus. The pleurocystidia of P. melanopotami-
cus were described (Singer 1989) as non-metuloidal, with
wall thicknesses varying from 0.3 to 0.6 µm, and with
round and small several finger-like protuberances at the
apex. In our collections of P. albostipitatus, a relatively
thickened wall, up to 1.0 µm, was observed which could
be compared to those of P. melanopotamicus. On the other
hand, the pleurocystidia with apical ornamentation of P.
melanopotamicus were not found, but they were observed
in the P. albostipitatus collections studied by Wartchow et
al. (2006). Probably, either P. melanopotapicus is a later
synonym of P. albostipitatus or the material identified by
Wartchow et al. (2006) as P. albostipitatus is actually P.
melanopotamicus. These materials should be revised to
solve this question.
In the ITS + 5.8S analysis (Figure 2), P. albostipitatus
clustered (68% BS) in a clade with P. nigrolineatus,
P. albostipitatus var. poliobasis, P. harrisii and Pluteus
sp. (section Pluteus). The molecular relation of P.
albostipitatus with P. nigrolineatus and P. harrisii may
indicate the transitional position of P. albostipitatus
between section Hispidoderma and section Pluteus,
because P. nigrolineatus belongs to section Hispidoderma,
and P. harrisii, although classified in section Pluteus, has
pleurocystidia with poorly developed prongs. Pegler
(1983) also observed this intermediary position for
P. albostipitatus, due to the unusual pleurocystidia shape as
reported by Singer (1958), which could indicate an interme-
diate relationship with the metuloidal pleurocystidia of the
section Pluteus.
In the nLSU analysis (Figure 1), P. albostipitatus clus-
tered (82% BS) with P. salicinus. Despite the results of the
molecular analyses, P. albostipitatus is considered distinct
from all these species because, among other characteris-
tics, P. harrissi has a glabrous pileus without appressedly
fibrils and metuloidal pleurocystidia (Murrill 1911; Singer
1956, 1958; Pegler 1983; Banerjee and Sundberg 1995),
P. nigrolineatus is a clamped species (Singer 1956, 1961;
Wartchow et al 2006), and P. salicinus is also clamped
and has characteristically metuloid pleurocystidia (Singer
1956; Orton 1986; Banerjee and Sundberg 1995).
Pluteus aureovenatus Menolli & Capelari, sp. nov. (Figure 4)
MycoBank: MB 513078
Etym.: The name refers to the colour and ornamenta-
tion of the pileus surface.
Pileus 22 mm diam, conicus vel convexus, aureofla-
vus, hygrophanus, venosus ad rugulosum, venae rubro-
luteae, margo leviter sulcata, striata et leviter erosa.
Lamellae subliberae, luteae. Stipes 30 × 1(apex)–3(basis)
mm, curvus, leviter compressus, subbulformis in basi, cen-
tralis, luteus. Basidiosporae 6.2(−7.5) × (5.0–)6.2(−7.5) µm,
globosae, hyalinae, inamiloydeae, laeves, crassis parietibus.
Pleurocystidia (50–)56–68(−64) × (12.5–)15.0–22(−31)
µm, clavata, hyalina, cum parietibus tenuibus. Cheilo-
cystidia (25–)31–49(−61) × (8.7–)10.0–17.0(−21) µm, cla-
vate vel versiformi-clavata, interdum leviter ventricosa,
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140 N. Menolli Jr. et al.
hyalina, cum parietibus tenuibus. Pileipellis cellularis a
strato unico composita ex cellulis subglobosis, 54–70 ×
(36–)46–58 µm, hyalineis, incoloratis, absque pedicellis,
interdum papilla apicali munitis. Fibulae absentes.
Holotypus: BRAZIL, São Paulo State, São Paulo city,
Parque Estadual das Fontes do Ipiranga, 24.I.2008, F.
Karstedt & L.A.S. Ramos FK1045 (SP).
Pileus 22 mm diam, conic to convex, deep yellowish-
orange, hygrophanous, veined-rugulose all over except at
the margin, vein reddish-orange, margin slightly sulcate,
translucently striate and slightly eroded. Lamellae sub-
free, yellow, crowded, with lamellulae. Stipe 30 × 1(apex)–
3(base) mm, curved, slightly compressed, with a subbul-
bous base, central, yellow, smooth, slightly longitudinally
striate. Odour, taste, and flesh colour not recorded. Basid-
iospores [20/1/1] 6.2(−7.5) × (5.0–)6.2(−7.5) µm [Q =
1.00(−1.24); Qm = 1.03; Lm = 6.4 µm; Wm = 6.2 µm]
globose, rarely broadly ellipsoid, inamyloid, hyaline,
smooth, thick-walled, guttulate. Basidia 27–36 × 7.5–8.7
µm, clavate, slim, thin-walled, four-spored. Pleurocystidia
(50–)56–64(−68) × (12.5–)15–22(−31) µm, clavate, some-
times slightly fusoid, frequently with apical mucilage in
KOH, colourless, hyaline, thin-walled, moderately abun-
dant. Cheilocystidia (25–)31–49(−61) × (8.7–)10–17(−21)
µm, clavate to versiform-clavate, sometimes slightly ven-
tricose, colourless, hyaline, thin-walled, moderately abun-
dant. Lamellar edge heteromorphous. Lamellar trama
inverse, up to 44 µm wide, composed of thin-walled
hyphae, some commonly inflated, 2.5–10.0(–18.7) µm
diam, hyaline, septate, sometimes with oleiferous hyphae up
to 3.7 µm diam. Pileus context undifferentiated, up to 230
µm thick, composed of thin-walled hyphae, 3.7–8.7 µm
diam, hyaline, septate. Pileipellis cellular approximately
56 µm, composed of one layer of subglobose cells, 54–70 ×
(36–) 46–58 µm, non-pedicelate, thin-walled, colourless,
hyaline, occasionally with an apical extension digitate to
papillate up to 20 µm long. Clamp connections absent in
all parts examined.
Habitat and habit: Solitary, on wood.
Material examined: BRAZIL, São Paulo State, São
Paulo city, Parque Estadual das Fontes do Ipiranga, 24 Jan.
2008, F. Karstedt & L.A.S. Ramos FK1045 (SP—holotype).
Comments: Pluteus aureovenatus is characterised by a
deep yellowish-orange and veined pileus, conic to convex
and margin translucently striate, sub-free and yellow lamel-
lae, globose basidiospores, large non-pedicelate pileipellis
cells, and frequently clavate pleuro- and cheilocystidia.
This species is closely related to P. aurantiacus Murrill,
P. aurantiorugosus, and P. laetifrons (Berk. & Curt.) Sacc.
However, according to the type description (Murrill, 1917)
and Smith and Stuntz (1958), P. aurantiacus is different
because the pileus is convex umbonate, the basidiospores
are subglobose to broadly ellipsoid, the pleuro- and cheilo-
cystidia are similar and slightly shorter [28–38 (−50) × 9–14 ×
5–10 µm] when compared to P. aureovenatus, and the cells
of the pileipellis are clavate-pedicellate and much smaller
(15–33 µm). Singer (1956) considered P. aurantiacus as a
synonym for P. aurantiorugosus. However, Smith and
Stuntz (1958) recognised them as distinct species, since P.
aurantiorugosus has an orange stipe and slightly longer and
more elongated basidiospores. Regardless, P. aurantiorugo-
sus differs from P. aureovenatus because, according to
Homola (1972), P. aurantiorugosus has a whitish to yel-
lowish stipe at first and then orange to red tints at the base, a
orange to yellowish-orange pileus in the dried specimens
instead of beige-straw for P. aureovenatus, basidiospores that
are not predominantly globose (6–7 × 4.5–5 µm), and cells of
the pileipellis that are vesiculose, clavate, or occasionally
Figure 4. Pluteus aureovenatus (holotype). A. Basidioma.
B. Basidiospores. C. Basidia. D. Pleurocystidia. E. Cheilo-
cystidia. F. Pileipellis cells. Bars (A) = 1 cm; (B–F) = 10 µm.
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Mycology 141
subfusoid-ventricose and much shorter (22–39 × 22–31 µm)
than P. aureovenatus.
Pluteus laetifrons is another species with orange-red
pileus related to P. aureovenatus. However, Dennis (1953)
and Singer (1958) described P. laetifrons with slightly
smaller basidiospores (spheric with 5–6 µm and 4.5–5.5 ×
4.3–5.3 µm, respectively) and according to Dennis (1953)
much smaller cells of the pileipellis (20–35 µm). Singer
(1958) described a new variety of P. laetifrons from
Bolivia, viz. P. laetifrons var. bolivianus Singer, that dif-
fers from the type variety due to its larger and almost geo-
metrically globose basidiospores (5.3–6.8 × 5.3–6.8 µm)
and its darker, deeply coloured spots on the pileus that
consist of pigmented pileipellis cells. Nevertheless, P.
aureovenatus has pileipellis cells that are always colour-
less and basidiospores that are occasionally broadly ellip-
soid and slightly longer than those described for
P. laetifrons var. bolivianus. Moreover, Singer (1958) did
not mention the size of pileipellis cells for P. laetifrons
var. bolivianus, which are approximately 20–35 µm, as
described in the type variety and they are, therefore, much
shorter than those observed for P. aureovenatus.
The relationship of P. aureovenatus with P. aurantiacus,
P. aurantiorugosus and P. laetifrons is probably comparable
to the relation between P. chrysophlebius and P. sublaevigatus
(see discussion under P. sublaevigatus), where these species
are separated by molecular data and have some micro-
morphological differences such as pileipellis cell size and
basidiospores that differ slightly in size and shape.
In the molecular analyses (Figures 1 and 2), P. aureo-
vanatus clustered in the clade with all species of section
Celulloderma and together with P. romellii (from USA –
California) and P. sublaevigatus. However, P. romellii is
described to have a different pileus colour (date-brown,
umber or snuff-brown), much longer pleuro- and cheilo-
cystidia [52–100 × (16–)20–40 µm and 30–80 × 12–40(−53)
µm, respectively] and pedicelate pileipellis cells with brown-
ish vacuolar contents (Orton 1986). Pluteus sublaevigatus
differs from P. aureovenatus because it has a yellow pileus
that is slightly rugulose just at the centre and without orange
tints or reddish-orange veins all over, a sterile lamellar edge
and shorter [(35–)44–51(−55) × (22–) 29–34 µm] pedicelate
pileipellis cells (see description under P. sublaevigatus). Plu-
teus aureovenatus belongs to section Celluloderma.
Pluteus dominicanus var. hyalinus Menolli & Capelari,
var. nov. (Figure 5)
MycoBank: MB 513080
Etym.: The varietal name refers to the hyaline and col-
ourless pileipellis cells.
Similis Pluteus dominicanus var. dominicanus Singer
sed cellulis hyalinis et achromaticis pileipellis differt.
Holotypus: BRAZIL, São Paulo State, São Paulo city,
Parque Estadual da Cantareira, Núcleo Engordador,
2.I.2008, F. Karstedt et al. FK1058 (SP).
Pileus 31 mm diam, plane-convex, slightly umbonate,
beige to light brown, slightly rugose at the centre, margin
translucently striate, slightly sulcate or sometimes eroded.
Lamellae free, pinkish, subdistant, with lamellulae. Stipe
54 × 2(apex)–5(base) mm, tapering towards the apex,
central, whitish to beige, smooth. Odour, taste, and flesh
colour not recorded. Basidiospores [20/1/1] 6.2–7.5(−8.7)
× 5.0–6.2(−7.5) µm [Q = 1.16–1.24(−1.40); Qm = 1.22;
Lm = 7.3 µm; Wm = 6.0 µm], broadly ellipsoid, rarely
ellipsoid, inamyloid, hyaline, smooth, thick-walled, some-
times guttulate. Basidia 20–29(−34) × 7.5–8.7 µm, clavate
to versiform, thin-walled, four-spored. Pleurocystidia
(35–)42–61(−69) × (10.0–)11.2–16.2(−17.5) µm, slightly
ventricose, lancelate or sometimes fusiform-clavate, col-
ourless, hyaline, sparse and few seen, thin-walled. Cheilo-
cystidia, like the pleurocystidia, but smaller and very rare,
(20–)29–43(−45) × (8.7–)10.0–14.0 µm. Lamellar edge
heteromorphous. Lamellar trama inverse, up to 50 µm
wide, composed of thin- or slightly thick-walled hyphae,
2.5–7.5(−15.0) µm diam, hyaline. Pileus context undiffer-
entiated, approximately 280 µm thick, composed of thin-
or slightly thick-walled hyphae, 3.7–7.5(−18.7) µm diam,
hyaline. Pileipellis cellular up to 44 µm thick, composed
of one layer of clavate to spheropedunculate cells, (26–)
32–47(−53) × (13.7–)16.2–21(−25) µm, with a short to
Figure 5. Pluteus dominicanus var. hyalinus (holotype).
A. Basidioma. B. Basidiospores. C. Basidia. D. Pleurocystidia. E.
Cheilocystidia. F. Pileipellis cells. Bars (A) = 1 cm; (B–F) = 10 µm.
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142 N. Menolli Jr. et al.
long pedicel (6.2–25 µm long), thin-walled, colourless,
hyaline. Clamp connections absent in all parts examined.
Habitat and habit: Solitary, on wood.
Material examined: BRAZIL, São Paulo State, São
Paulo city, Parque Estadual da Cantareira, Núcleo Engordador,
02 Jan. 2008, F. Karstedt et al. FK1058 (SP—holotype).
Comments: Pluteus dominicanus var. hyalinus is pro-
posed as a variety of P. dominicanus especially because of
the presence of hyaline and colourless pileipellis cells.
P. dominicanus is known only from its type locality (Ven-
ezuela) in its original description (Singer 1961) and was
subsequently recorded by Dennis (1970) based on the
same material. Singer (1961) established P. dominicanus
as “cheilocystidia not seen” and having pileipellis cells
“with deep brownish-melleous dissolved intracellular pig-
ment”, instead of the Brazilian material in which the
cheilocystidia are rarely seen and the pileipellis cells are
hyaline and colourless. The re-examination of the type of
P. dominicanus (K!) demonstrated that the major differ-
ence of the new variety is, in fact, regarding the colour of
the pileipellis cells. The type of P. dominicanus (Figure 6)
has broadly ellipsoid or rarely subglobose basidiospores
[20/1/1] 7.5–8.7(−10.0) × 6.2–7.5(−8.7) µm [Q = (1.15–)
1.16(−1.21); Qm = 1.17; Lm = 8.7 µm; Wm = 7.4 µm];
slightly lancelate to ventricose pleurocystidia, 44–55 ×
15.0–19.0 µm; cheilocystidia present, 36–55 × 12.5–
20(−24) µm, like the pleurocystidia, but that are very rare and
only possible to see in tangential view; and pileipellis com-
posed of vesiculose to espheropedunculate cells, 37–48(−57)
× 22–34(−39) µm, with a short to medium pedicel (2.5–10.0
µm long) and condensed or dissolved brownish pigment.
Pluteus dominicanus var. hyalinus is also related to the
North American species P. pallidus and P. umbrinodiscus,
but it differs from these especially in the pileus colour and
the frequency of pleuro- and cheilocystidia.
Pluteus dominicanus belongs to section Celluloderma
and clustered with all other species from section Cellulo-
derma in both molecular analyses (Figures 1 and 2)
Pluteus fluminensis Singer, Lloydia 21: 292 (1959).
(Figure 7)
Pileus 27–31 mm diam, plane-convex, eventually
slightly depressed at the centre or concave, not
umbonate, deep sepia (N80A90M60) to porphyry brown,
darker brown and rugose to venose at the centre, slightly
paler towards the margin, margin sulcate or finely crack-
ing and showing the white flesh. Lamellae free to sub-
free, white then pinkish, close, with lamellulae. Stipe 30–
38 × 2–3(apex)–5(base) mm, tapering towards the apex,
with or without a small bulb, central, hollow, pale cream
slightly greyish or translucent, finely pruinose and with finely
brownish fibrils at base, slightly longitudinally striate, with
scanty basal mycelium. Odour, taste, and flesh colour not
recorded. Basidiospores [80/4/4] 6.2–7.5 × 5.0–6.2 µm
(Q = 1.21–1.24; Qm = 1.23; Lm = 6.8 µm; Wm = 5.5
µm), broadly ellipsoid, inamyloid, hyaline, smooth,
thick-walled, guttulate. Basidia 21–37 × 7.5–8.7(−10.0) µm,
clavate to versiform, sometimes more slender, thin-
walled, four-spored. Pleurocystidia (41–)47–65(−69) ×
12.5–21(−25) µm, ventricose or slightly ventricose to
lageniform, rarely vesiculose-saccate, colourless, hya-
line, thin-walled, sparse, moderately abundant to scat-
tered. Cheilocystidia (25–)32–50(−56) × (10.0–)11.2–
18.7(−22)µm, like the pleurocystidia or rarely subclavate
to clavate, difficult to see, moderately numerous. Lamel-
lar edge heteromorphous. Lamellar trama inverse, up to 62
µm wide, composed of thin- or slightly thick-walled hyphae,
Figure 6. Pluteus dominicanus (holotype). A. Basidiospores.
B. Basidia. C. Pleurocystidia. D. Cheilocystidia. E. Pileipellis
cells. Bars = 10 µm.
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Mycology 143
1.2–8.7(−15.0) µm diam, hyaline, septate. Pileus context
undifferentiated, up to 125 µm thick, composed of thin or
slightly thick-walled hyphae, 3.7–5.0 µm diam, hyaline, sep-
tated, sometimes with oleiferous up to 3.7 µm diam. Pileipel-
lis cellular approximately 45 µm thick, composed of one
layer of subglobose, vesiculose or subclavate cells, (20–)
25–46(−52) × (17.5–) 20–32(−37) µm, frequently non-pedi-
celate, often with a short to medium pedicel (1.2–7.5 µm
long), thin-walled, with condensed or dissolved brown
pigment. Clamp connections absent in all parts examined.
Habitat and habit: Solitary or in pairs, on wood.
Material examined: BRAZIL, São Paulo State,
Cananéia city, Ilha do Cardoso, between Morro Três
Irmãos and Sítio Grande, 23 Oct. 1984, M. Capelari
MC164 (SP); near Morro Três Irmãos, 18 Dec. 1984, M.
Capelari & R. Maziero MC212 (SP); São Paulo city,
Parque Estadual da Cantareira, Núcleo Engordador, 24
Apr. 2007, Menolli Jr. et al. NMJ127 (SP); 24 Jan. 2008,
F. Karstedt & L.A.S. Ramos FK1046 (SP).
Additional specimen examined: BRAZIL, Rio de
Janeiro State, Angra dos Reis, 31 Sep. 1953, R. Singer
B432 (F—holotype).
Comments: Pluteus fluminensis was described from
Brazil (Rio de Janeiro State) and was also reported for
Bolivia and USA (Florida) in its original description
(Singer 1958). Later, Pegler (1997) reported it for São
Paulo State and Stijve and Meijer (1993) and Meijer
(2006) for Paraná State as P. cf. fluminensis. The basid-
iospores of P. fluminensis were described in the proto-
logue (Singer, 1958) as “subglobose or rarely a few
piriform, (4.5–)5–6.3(−7) × (4–)4.3–5.3(−5.8) µm”; how-
ever, upon re-examination of the type material (F!), it was
demonstrated that the basidiospores are broadly ellipsoid,
[20/1/1] 6.2–7.5 × 5.0–6.2 µm (Q = 1.21–1.24; Qm = 1.23;
Lm = 6.8 µm; Wm = 5.5 µm). The pleuro- and cheilocystidia
of the type are probably ventricose, but they had collapsed
and it was not possible to confirm their size and shape. The
pileipellis cells of the type are subglobose to vesiculose and
slightly longer [(37–)41–57(−61) × (29–) 31–38(−42) µm]
than those of the other collections herein studied.
Pluteus jamaicensis is closely related to P. fluminensis,
but, according to Singer (1958), the first has epicuticular
elements with dissolved internal pigment instead of the
pigmented condensations observed in P. fluminensis. The
occurrence of pileipellis cells with condensed brown pig-
ment was confirmed in the re-examination of the type and
in additional collections of P. fluminensis herein exam-
ined. Singer (1958) also considered a future possibility of
a subspecies status for P. fluminensis in P. jamaicensis,
but herein they are considered as two distinct species.
Figure 7. Pluteus fluminensis. A. Basidioma; A
1. Menolli Jr. et al. NMJ127; A2. F. Karstedt & L.A.S. Ramos FK1046. B–F. F.
Karstedt & L.A.S. Ramos FK1046. B. Basidiospores. C. Basidia. D. Pleurocystidia. E. Cheilocystidia. F. Pileipellis cells. Bars (A) = 1 cm;
(B–F) = 10 µm.
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144 N. Menolli Jr. et al.
Pluteus fluminensis belongs to section Celluloderma
and clustered with all other species from section Cellulo-
derma in both molecular analyses (Figures 1 and 2).
Pluteus fuligineovenosus Horak, Nova Hedwigia 8: 190
(1964). (Figure 8)
Pileus 14–38 mm diam, plane to slightly conic, appar-
ently subumbonate, dark brown, slightly paler towards the
margin and darker to blackish at the centre, veined all over
especially around the centre then radially arranged towards
the margin, vein darker, margin apparently neither sulcate nor
striate. Lamellae free to slightly sinuate, white then pinkish,
subdistant, with few lamellulae. Stipe 8–31 × 1–4 mm,
slightly tapering towards the apex, central to slightly excen-
tric, slightly flexuous, apparently hollow, translucent white,
slightly longitudinally striate, with scanty basal mycelium.
Odour, taste, and flesh colour not recorded. Basidiospores
[20/1/1] 6.2–7.5 × (5.0–)6.2–7.5 µm [Q = 1.00(−1.24); Qm =
1.06; Lm = 6.5 µm; Wm = 6.2 µm], globose, occasionally
broadly ellipsoid, inamyloid, hyaline, smooth, thick-walled,
guttulate. Basidia 18.7–21(−25) × 6.2–7.5 µm, clavate to ver-
siform, thin-walled, 4-spored. Pleurocystidia (51–)57–77 ×
11.2–18.7 µm, fusoid-ventricose to lancelate, colourless, hya-
line, thin-walled, sparse and moderately abundant. Cheilo-
cystidia (37–)45–63 × (12.5–)15.0–20 µm, clavate to
elongate-clavate, colourless, hyaline, thin-walled, not numer-
ous. Lamellar edge apparently sterile. Lamellar trama
inverse, up to 31 µm wide, composed of thin- or slightly
thick-walled hyphae, 3.7–11.2 µm diam, hyaline, few septa
seen, with oleiferous hyphae up to 5.0 µm diam. Pileus con-
text undifferentiated, approximately 47 µm thick, composed
of thin or slightly thick-walled hyphae, 5.0–7.5 µm diam,
hyaline, few septa seen. Pileipellis cellular up to 100 µm
thick, composed of one layer of vesiculose to spheropeduncu-
late cells, (32–) 39–50(−62) × (17.5–)20–32 (−39) µm, with a
short to long pedicel (2.5–23 µm long), thin-walled, with con-
densed or dissolved brown to chestnut-brown pigment.
Clamp conncetions absent in all parts examined.
Habitat and habit: Gregarious to disperse, on wood.
Material examined: BRAZIL, São Paulo State, São
Paulo city, Parque Estadual da Cantareira, Núcleo Engordador,
12 Dec. 2006, F. Karstedt & M. Capelari MC826 (SP).
Comments: Pluteus fuligineovenosus is characterised
by a brown and veined pileus, with darker to blackish cen-
tre and veins, globose basidiospores, and spheropeduncu-
late cells of the pileipellis with condensed or dissolved
brown to chestnut-brown pigment.
This species is known from Chile where it was origi-
nally described (Horak 1964), and later from Argentina
(Singer 1969). The Brazilian collection has basidiospores
that are slightly bigger than those mentioned in the original
description (5.6–6.6 × 5.2–5.6 µm), but more similar in size
[5.8–7(−7.8) × 5−6.2 µm] to those from the Argentinean
collections (Singer 1969), pleurocystidia that are more fre-
quently fusoid ventricose, and stipe that is apparently hol-
low and with scanty mycelium at the base, instead of solid
stipe and a lack of basal micelium as described from the
Chilean material (Horak 1964). This work reports for the
first time P. fuligineovenosus from Brazil, and it is the
second collection since its description.
In the ITS + 5.8S analyses (Figure 2), P. fuligineovenosus
clustered in the clade with other species from section
Celluloderma, forming a well-supported clade (99% BS)
with P. fluminensis and P. jamaicensis. However, P. fulig-
ineovenosus can be distinguished from P. fluminensis and
P. jamaicensis by the basidiospore shape (preponderantly
globose in P. fuligineovenosus instead of mostly broadly
ellipsoid in P. fluminensis and P. jamaicensis) and the
Figure 8. Pluteus fuligineovenosus (F. Karstedt & M. Capelari
FK826). A. Basidioma. B. Basidiospores. C. Basidia. D. Pleuro-
cystidia. E. Cheilocystidia. F. Pileipellis cells. Bars (A) = 1 cm;
(B–F) = 10 µm.
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Mycology 145
pileus ornamentation, because P. fuligineovenosus is char-
acterised by a pileus with a darker to blackish centre and
veins all over, especially around the centre and then radi-
ally arranged towards the margin.
Pluteus harrisii Murrill, Mycologia 3:277(1911).
(Figure 9)
Pileus 20–33(−55) mm diam, conic to convex, slightly
umbonate, hazel (N60A80M70) to brown or with coffee
shades, slightly darker at the centre, with or without finely
white punctuation especially at the centre, margin finely sul-
cate to sulcate-striate. Lamellae free, white then pinkish,
attachment partially unequal, crowded, with lamellulae. Stipe
30–48(−62) × 2(apex)–5(base) mm, subequal, central, occa-
sionally flexuous, apparently hollow, white to whitish grey,
slightly pruinose at the apex, longitudinally striate, with basal
mycelium. Odour, taste, and flesh colour not recorded.
Basidiospores [80/4/4] 7.5–8.7(−10.0) × (5.0–)6.2–7.5
(−8.7) µm [Q = (1.15–)1.16–1.21(−1.50); Qm = 1.23; Lm =
8.2 µm; Wm = 6.7 µm], broadly ellipsoid, occasionally ellip-
soid or rarely subglobose, inamyloid, hyaline, slightly pinkish
in KOH, smooth, thick-walled, sometimes guttulate. Basidia
24–30 × 7.5–10.0 µm, versiform to clavate, thin-walled, four-
spored, sometimes with small scattered guttules. Pleuro-
cystidia (40–)52–82(−91) × 15.0–24 µm, fusoid-ventricose,
colourless, hyaline, sparse, moderately numerous to abundant,
moderately thick-walled, wall usually uniform up to 1.2 µm
Figure 9. Pluteus harrisii. A–C. Menolli Jr. et al. NMJ122. A. Basidioma. B. Basidiospores. C. Basidia. D. Pleurocystidia; D1. Menolli
Jr. et al. NMJ122; D2. F. Karstedt et al. FK1066. E. Cheilocystidia; E1. Menolli Jr. et al. NMJ122; E2. F. Karstedt et al. FK1066. Bars
(A) = 1 cm; (B–E) = 10 µm.
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146 N. Menolli Jr. et al.
wide or sometimes thinner towards the base and thickening
up to 2.5 µm wide at the apex, apices usually with two to four
versiform lateral prongs (up to 5.0 µm long), prongs rarely
with a secondary bifurcation, occasionally without prongs
and with a rounded or deformed apex, sometimes a colourless
internal condensed content is present. Cheilocystidia (31–)36–
59 × 8.7–15(−17.5) µm, clavate to versiform-clavate, usually
with a moderately long pedicel, colourless, hyaline, moder-
ately abundant, thin-walled. Lamellar edge sterile. Lamellar
trama inverse, up to 62 µm wide, composed of thin or
slightly thick-walled hyphae, 2.5–13.7 µm diam, sometimes
branched or with terminal elements slightly inflated up to
17.5 µm diam, hyaline, septate. Pileus context undifferenti-
ated, up to 200 µm thick, composed of thin- or slightly thick-
walled hyphae, 3.7–8.7 µm diam, sometimes inflated up to 25
µm diam, hyaline, septate. Pileipellis a repent epicutis, up to
100 µm thick, composed of thin- to slightly thick-walled
hyphae, 5.0–12.5 µm diam, elongated, septate, with light to
dark brown content, sometimes with terminal elements slightly
inflated. Clamp connections absent in all parts examined.
Habitat and habit: Solitary, on leaf of palm tree or
decaying wood.
Material examined: BRAZIL, São Paulo State, São
Miguel Arcanjo city, Parque Estadual Carlos Botelho, 25
Apr. 1986, M. Capelari & V.L.R. Bononi MC10 (SP);
Santo André city, Reserva Biológica de Paranapiacaba, 11
Apr. 1990, M. Capelari et al. MC3282 (SP); 22 Mar.
2007, Menolli Jr. et al. NMJ122 (SP); São Paulo city,
Parque Estadual da Cantareira, Núcleo Engordador, 31
Jan. 2008, F. Karstedt et al. FK1066 (SP).
Comments: Pluteus harrisii is characterised by a dark
brown pileus, a white stipe with greyish tones, and metu-
loids with poorly developed apical prongs. This species
was originally described from Jamaica by Murrill (1911),
who established P. harrisii without cystidia. However,
Singer (1956, 1958), Pegler (1983) and Banerjee and Sun-
dberg (1995) found thick-walled pleurocystidia with short
apical prongs in the holotype.
Besides the type locality, P. harrisii was also found in
Cuba (Murrill 1911), Trinidad (Baker and Dale 1951 as
“P. cervinus var. bambusinus”), USA – Florida (Singer
1956, 1958), Guadeloupe (Pegler 1983), Mexico (Vargas
1993; Rodríguez and Guzmán-Dávalos 2001), and Brazil
(Pegler 1997; Meijer 2006 as P. aff. harrisii).
The Brazilian materials herein studied has basid-
iospores moderately larger than those found by Singer
(1956, 1958) from the Central and North American materi-
als (6.0–9.8 × 5.0–6.5 µm) and pleurocystidia sometimes
with a colourless amorphous substance, as recorded by
Banerjee and Sundberg (1995) in the type revision. Moreo-
ver, it has a stipe that is apparently hollow instead of a solid
stipe as recorded by Singer (1956, 1958) and Pegler (1983).
Two of the studied collections, Menolli Jr. et al.
NMJ122 and F. Karstedt et al. FK1066, have the same
nLSU sequence and differ only in two base pairs in the ITS +
5.8S sequence. However, Menolli Jr. et al. NMJ122 differs
from F. Karstedt et al. FK1066 by the presence of a colour-
less internal condensed content in the pleurocystidia and
fewer cheilocystidia that are usually versiform. It will be
necessary to study more collections to determine whether
these characters are constant in successive fruitings to verify
if they constitute intraspecific variations or if is sufficient to
consider them as independent taxa.
The molecular analyses (Figures 1 and 2) show the
sequence of P. harrisii clustered with all other species of the
section Pluteus and forming a low-supported clade (68%
BS), albeit in a separate branch, with P. albostipitatus.
However, P. albostipitatus is currently classified in the sec-
tion Hispidoderma (see discussion under P. albostipitatus)
and differs from P. harrisii as it has a densely fibrillose
pileus and pleurocystidia that are not distinctly metuloid and
frequently without prongs. Pluteus harrisii belongs to sec-
tion Pluteus.
Pluteus jamaicensis Murrill, Mycologia 3:278 (1911)
(Figure 10)
Pileus 17–34 mm diam, campanulate-convex, some-
times slightly umbonate, dark beige to dark brown
(N70A80M70), slightly paler towards the margin, finely
rivulose all over, rugose to rivulose at the centre, margin
slightly sulcate and sometimes finely cracking and showing
the white flesh. Lamellae free to sinuate-remote, white then
pinkish, with numerous lamellulae. Stipe 20–30 × 1–2 mm,
equal, central, hollow, cream to translucent white, slightly
greyish at base. Odour, taste, and flesh colour not
recorded. Basidiospores [20/1/1] 6.2(−7.5) × 5.0(−6.2) µm
[Q = (1.21–)1.24; Qm = 1.24; Lm = 6.4 µm; Wm = 5.2
µm], broadly ellipsoid, inamyloid, hyaline, smooth, thick-
walled, sometimes guttulate. Basidia 25–29(−31) ×
6.2–8.7 µm, clavate to versiform, thin-walled, four-
spored. Pleurocystidia (46–)50–56 × 10.0–13.7(−17.5) µm,
fusoid-ventricose to lancelate, colourless, hyaline, thin-
walled, sparse and not abundant. Cheilocystidia (25–)31–
45(−50) × (7.5–)10.0–12.5 µm, clavate to elongate-clavate,
colourless, hyaline, thin-walled, numerous. Lamellar edge
heteromorphous with few basidia. Lamellar trama inverse,
up to 62 µm wide, composed of thin- or slightly thick-
walled hyphae, 3.7–13.7 µm diam, hyaline. Pileus context
undifferentiated, approximately 62 µm thick, composed of
thin- or slightly thick-walled hyphae, hyaline, 3.7–10.0 µm
diam. Pileipellis cellular up to 125 µm, composed of one or
more layers of spheropedunculate cells, (37–)45–54 × (31–)37–
42 µm, with a short pedicel (5.0–10.0 µm long), thin-walled,
with dissolved brown cytoplasmic content. Clamp connec-
tions absent in all parts examined.
Habitat and habit: Gregarious, on wood.
Material examined: BRAZIL, São Paulo State, São
Paulo city, Parque Estadual da Cantareira, Núcleo Engor-
dador, 24 Apr. 2007, Menolli Jr. et al. NMJ130 (SP).
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Mycology 147
Comments: Pluteus jamaicensis was described from
Jamaica (Murrill 1911), and since then, it was reported
from Trinidad (Dennis 1953 as P. aethalus var. jamaicensis;
Pegler 1983), Argentina (Singer and Digilio 1951 as “P.
phlebophorus var. ?”; Singer 1956, 1958), Venezuela ?
(Dennis 1970), and Martinique and Guadalupe (Pegler
1983). Moreover, according to Singer (1958), P. phlebo-
phorus (Ditmar) P. Kumm. sensu Rick (1938) reported
from São Leopoldo, Rio Grande do Sul State, Brazil, is
perhaps P. jamaicensis. This species was originally
described without cystidia (Murrill 1911), but posterior
revisions of the type confirmed the presence of marginal
and facial cystidia (Singer 1958; Smith and Stuntz 1958;
Pegler 1983; Banerjee and Sundberg 1993). The Brazilian
collection has a slightly sulcate pileus margin, as reported
by Singer (1956, 1958) from Argentinean materials; prepon-
derantly clavate cheilocystidia, as described by Pegler (1983);
and broadly ellipsoid basidiospores that are slightly longer
than those described by Smith and Stuntz (1958) in the revi-
sion of the type specimen (subglobose, 5–6 × 5–5.5 µm).
In the ITS + 5.8S analysis (Figure 2), P. jamaicensis
clustered with P. fluminensis and P. fuligineovenosus.
However, they are morphologically distinct (see discussion
under P. fluminensis and P. fuligineovenosus). This work
reports P. jamaicensis for the first time from Brazil. As sup-
ported by the molecular (Figure 2) and the morphological
data, Pluteus jamaicensis belongs to section Celluloderma.
Pluteus riberaltensis var. conquistensis Singer, Lloydia
21: 255 (1959). (Figure 11)
Pileus 27–53 mm diam, campanulate when young,
then convex, umbonate, dark brown, slightly darker in the
centre, slightly radially coloured, finely pruinose all over
and finely cracking especially around the centre, margin
not striate and showing the whitish flesh between the
fibrils. Lamellae free, pinkish, moderately crowded, with
lamellulae. Stipe 53–65 × 4–9 mm, slightly tapering
towards the apex, base subbulbous, central to slightly
eccentric, white-cream with brown fibrils over the surface
especially when young, then brownish and slightly longi-
tudinally striate, with scanty mycelium at base. Odour,
taste, and flesh colour not recorded. Basidiospores [20/1/1]
6.2–7.5 × 6.2 µm (Q = 1.00–1.21; Qm = 1.08; Lm = 6.7 µm;
Figure 10. Pluteus jamaicensis (Menolli Jr. et al. NMJ130). A. Basidioma. B. Basidiospores. C. Basidia. D. Pleurocystidia. E. Cheilo-
cystidia. F. Pileipellis cells. Bars (A) = 1 cm; (B–F) = 10 µm.
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148 N. Menolli Jr. et al.
Wm = 6.2 µm), globose or occasionally broadly ellipsoid,
inamyloid, hyaline, smooth, thick-walled, guttulate.
Basidia 21–27(−31) × (7.5–)8.7–10.0 µm, clavate, thin-
walled, four-spored, with small scattered guttules. Pleuro-
cystidia (44–)48–69(−78) × (15.0–)17.5–28(−35) µm,
slightly ventricose to lageniform, sparse and not abundant.
Cheilocystidia (27–)30–47(−57) × (12.5–)17.7–27 µm,
like pleurocystidia, but very rooting on the hymenium and
hard to see. Lamellar edge heteromorphous with abundant
basidia. Lamellar trama inverse, up to 37 µm wide, com-
posed of thin- or thick-walled hyphae, 3.7–10.0 µm diam,
hyaline, few septa seen, with oleiferous hyphae up to 3.7 µm
diam. Pileus context undifferentiated, approximately 125
µm thick, composed of thin- or thick-walled hyphae, 3.7–
7.5 µm diam, hyaline, septate. Pileipellis a cutis, approxi-
mately 100 µm thick, composed of thin-walled hyphae,
10–20 µm diam, elongated, septate, with dissolved brown
vacuolar content, sometimes with the terminal elements
slightly inflated and often ascendant. Clamp connections
absent in all parts examined.
Habitat and habit: In pairs on wood.
Material examined: BRAZIL, São Paulo State, São
Paulo city, Parque Estadual das Fontes do Ipiranga, 24 Jan.
2008, F. Karstedt & L.A.S. Ramos FK1043 (SP).
Comments: This variety was described from Bolivia
(Singer 1958) and differs from the type variety almost
exclusively by the umber-grey fibrils on the stipe. The
Brazilian specimen is characterised by brown fibrils over
the stipe surface, especially when young, as described for
this variety. Also, there are some cavities on the pileus
surface showing the flesh, probably caused by insects.
However, our material has moderately longer pleuro-
cystidia than those described for P. riberaltensis var.
conquistensis (38–41 × 14–22.5 µm), and these are more
similar to those of the type variety (41–86 x 12.5–39 µm),
as described in the protologue (Singer 1958).
Singer (1961) described another variety, viz. P. riber-
altensis var. missionensis Singer, based on an Argen-
tinean specimen that has a white stipe similar to the type
variety but that differs from the latter by the conspicuous
incrustation of the cystidia, and the missing differentia-
tion in size of cheilo- and pleurocystidia. Pluteus riber-
altensis and its varieties were described based on single
collections and no other records have been reported. This
work is the first to mention P. riberaltensis var. con-
quistensis from Brazil and reports the first collection
since its description.
Pluteus riberaltensis var. conquistensis, as well as other
species formerly classified in the section Hipidoderma
(P. ephebeus, P. exiguus, P. umbrosus), clustered in the
clade including members of the section Celluloderma. As
discussed above, these species probably share morphological
similarities, and the scope of the section Hispidoderma must
be revised. For now, Pluteus riberaltensis var. conquistensis
has been classified in the section Hispidoderma.
Pluteus sublaevigatus (Singer) Menolli & Capelari, stat.
nov. (Figure 12)
MycoBank: MB 513081
Basionym: Pluteus chrysophlebius subsp. sublaevigatus
Singer, Lloydia 21: 278 (1959).
Pileus 11–28 mm diam, campanulate when young,
then convex to plane, not umbonate, sometimes slightly
depressed at the centre, yellow (N00A30M40 to
N00A80M30), hygrophanous, slightly rugulose at the cen-
tre, margin translucently striate, sometimes eroded.
Lamellae free to subfree, close, yellowish, moderately
crowded, with lamellulae. Stipe 28–33 × 1–4 mm, curved,
cylindric with a subbulbous base, central, whitish yellow,
more yellowish at the base, surface smooth, sometimes
with white punctuations and white pruinose mycelium at
the base. Odour, taste, and flesh colour not recorded.
Basidiospores [40/2/1] (5.0–)6.2–7.5 × 5.0–6.2(−7.5) µm
(Q = 1.00–1.21(−1.24); Qm = 1.08; Lm = 6.5 µm; Wm =
6.1 µm), preponderantly globose, sometimes broadly ellip-
soid, inamyloid, hyaline, smooth, thick-walled, guttulate,
sometimes with small scattered guttules. Basidia (20–)22–27
(−30) × 7.5–8.7(−10.0) µm, clavate, thin-walled, four-
spored. Pleurocystidia (34–)36–50(−65) × 12.5–17.5(−25)
µm, clavate to vesiculose, sometimes slightly ventricose,
very translucent and difficult to see in KOH, sparse and
very rare, thin-walled. Cheilocystidia (22–)31–49(−54)
Figure 11. Pluteus riberaltensis var. conquistensis (F. Karstedt
& L.A.S. Ramos FK1043). A. Basidioma. B. Basidiospores.
C. Basidia. D. Pleurocystidia. E. Cheilocystidia. Bars (A) = 1 cm;
(B–E) = 10 µm.
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Mycology 149
× (7.5–)10.0–21(−26) µm, like the pleurocystidia, clavate
to vesiculose, very translucent, abundant, thin-walled.
Lamellar edge sterile. Lamellar trama inverse, up to 62 µm
wide, composed of thin- or slightly thick-walled hyphae,
3.7–10 µm diam, hyaline, septate. Pileus context undiffer-
entiated, up to 187 µm thick, composed of thin- or slightly
thick-walled hyphae, 2.5–6.2(−16.2) µm diam, hyaline,
septate. Pileipellis cellular up to 50 µm thick, composed of
one layer of clavate, vesiculose or subglobose cells, (35–)
44–51(−55) × (22–)29–34 µm, with a short pedicel (2.5–
7.5 µm long), thin-walled, colourless, hyaline. Clamp con-
nections absent in all parts examined.
Habitat and habit: Gregarious on decaying wood.
Material examined: BRAZIL, São Paulo State, São
Paulo city, Parque Estadual da Cantareira, Núcleo Engor-
dador, 19 Feb. 2008, F. Karstedt et al. FK1085 (SP).
Additional specimen examined: BOLIVIA, Dpto La
Paz, Prov. Nor-Yungas, Charobamba, 13 Feb. 1956,
R. Singer 1147 (LIL—holotype).
Comments: The frequently globose basidiospores and
the large subglobose cells of the pileipellis, along with the
results of the molecular analyses, are the basis to propose
the new status for P. chrysophlebius ssp. sublaevigatus.
Singer (1958) used the frequently globose basidiospores
as the distinctive character to propose P. chrysophlebius
ssp. sublaevigatus based on material collected in Bolivia.
The Brazilian collection is slightly different because it has a
pileus slightly rugose at the centre and pleurocystidia pre-
dominantly clavate to vesiculose and slightly longer than
those described for P. chrysophlebius ssp. sublaevigatus,
which has a pileus with “the disc seemingly smooth when
wet, but in dry or dried condition becoming rugulose” and
cystidia “about 35 × 15 µm if vesiculose, about 53 × 9.5 µm
if ampullaceous” (Singer, 1958). These are minor differ-
ences that we consider intraspecific variations between the
two collections. Moreover, the re-examination of the type of
P. sublaevigatus (LIL!) demonstrated that it has basid-
iospores that are preponderantly globose and rarely broadly
ellipsoid, [20/1/1] 6.2–7.5 × 6.2–7.5 µm [Q = 1.00(−1.21);
Qm = 1.01; Lm =7.0 µm; Wm = 7.0 µm], as well as a
pileipellis composed of vesiculose or subglobose cells,
35–54 × 21–37 µm, characterised as thin-walled, colourless,
hyaline and by have a short pedicel (2.5–7.5 µm long). The
pleuro- and cheilocystidia of the type are collapsed and it
was not possible to confirm their size and shape.
Pluteus sublaevigatus is close to P. chrysophlebius and
P. admirabilis in morphology, but the molecular analysis
of the nLSU gene (Figure 1) showed that they clustered in
separated clades, with P. chrysophlebius and P. admirabilis
in one clade and P. sublaevigatus in another clade with the
Brazilian yellow species P. aureovenatus.
The morphological re-examination of the three collec-
tions (DAOM!) of P. admirabilis and P. chrysophlebius
used in the molecular analyses showed some differences in
microcharacters among themselves and when compared to
P. sublaevigatus (F. Karstedt et al. FK1085), especially in
the pileipellis cells (Figures 13–15). The pileipellis cells
from these collections are usually clavate-vesiculose to
subglobose, ranging in size and in the presence or absence of
apical projections (Table 2). Moreover, the size and shape of
basidiospores are a constant character; the pleurocystidia are
constantly slightly ventricose with few differences in size;
and the cheilocystidia are frequently clavate-vesiculose to
slightly ventricose (Table 2). These microcharacters partly
agree with the type study results presented by Banerjee
and Sundberg (1993) and Singer (1956) for P. admirabilis
and P. chrysophlebius, and they differ from those
observed in the Brazilian collection of P. sublaevigatus
(Table 2).
These differences are probably due to intraspecific
variations because in our analysis, the nLSU sequences of
P. chrysophlebius and P. admirabilis from GenBank clus-
tered in a clade with 99% BS. This relationship between
the sequences was already reported by Moncalvo et al.
Figure 12. Pluteus sublaevigatus (F. Karstedt et al. FK1085).
A. Basidioma. B. Basidiospores. C. Basidia. D. Pleurocystidia.
E. Cheilocystidia. F. Pileipellis cells. Bars (A) = 1 cm; (B–F) =
10 µm.
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150 N. Menolli Jr. et al.
(2002) and Minnis et al. (2006). Then, if these collections
really are representatives of P. chrysophlebius and
P. admirabilis, according to the morphological and molec-
ular data presented, they cannot be considered independent
species. This fact is in accordance with Murrill (1917) and
Singer (1956), who already considered them as synonyms.
The study of the voucher collections from the GenBank
sequences was important for establishing the morphological
Figures 13–15. 13. Pluteus admirabilis (DAOM193532). 14. P. admirabilis (DAOM197226). 15. P. chrysophlebius (DAOM190194).
A. Basidiospores. B. Basidia. C. Pleurocystidia. D. Cheilocystidia. E. Pileipellis cells. Bars = 10 µm.
Table 2. Micromorphological comparison between collections of Pluteus admirabillis, P. chrysophlebius and P. sublaevigatus.
Characters
Species/Collection
P. admirabillis
DAOM193532
P. admirabillis
DAOM197226
P. chrysophlebius
DAOM190194
P. sublaevigatus
F. Karstedt et al. 1085
Basidiospores 7.5 × 6.2 µm 7.5 × 6.2 µm 7.5 × 6.2 µm 6.2–7.5 × 5.0–6.2 µm
broadly ellipsoid broadly ellipsoid broadly ellipsoid preponderantly
globose
Pleurocystidia 47–71 × 16.2–21 µm 50–81 × 17.5–25 µm 41–52 × 10.0–18.7 µm 36–50 × 12.5–17.5 µm
slightly ventricose, with
apical mucilage
slightly ventricose slightly ventricose clavate to vesiculose
Cheilocystidia 21–38 × 7.5–12.5 µm 26.2–50 × 10.0–18.7 µm 30–40 × 10.0–11.2 µm 31–49 × 10.0–21 µm
clavate-vesiculose to
slightly ventricose
clavate-vesiculose to
slightly ventricose
clavate-vesiculose to
slightly ventricose
clavate to vesiculose
Pileipellis cells 16.2–51 × 12.5–25 µm 35–49 × 13.7–25 µm 26–37.5 × 12.5–17.5 µm 44–51 × 29–34 µm
with or without a medium
pedicel up to 12.5 µm long,
frequently with a short to
long apical projection
approximately
5.0–25 µm long
with a short to
medium pedicel up to
13.7 µm long, without
apical projections, and
sometimes with a
broadly obtuse apex
with a short to medium
pedicel approximately
2.5–15.0 µm long, and
rarely subventricose
with a broadly obtuse
apex up to 5.0 µm long
with a short pedicel
approximately
2.5–7.5 µm long
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Mycology 151
differences between them and the Brazilian collections, as
well as for supporting the results obtained with the molecular
analysis. Pluteus sublaevigatus differs from the vouchers of
P. chrysophlebius and P. admirabilis because it has close and
not remote lamellae, basidiospores that are frequently glo-
bose, larger pileipellis cells, and similar pleuro- and cheilo-
cystidia that are predominantly clavate-vesiculose (Table 2).
In the ITS + 5.8 analyses (Figure 2), P. sublaevigatus
clustered with low BS support (53%) together with
P. aureovenatus and P. romellii. However, as to P. aureo-
venatus, P. romellii can be distinguished from P. sublaevi-
gatus by the pileus colour and the size of the pleuro- and
cheilocystidia. Pluteus aureovenatus can also be separated
from P. sublaevigatus as discussed under the description
of P. aureovenatus.
Stijve and Meijer (1993) and Meijer (2006) recorded
in a checklist the occurrence of P. chrysophlebius subsp.
bruchii Singer from Paraná State, Brazil, which probably
represents P. sublaevigatus. The records of P. chrysophle-
bius for Martinique and Guadeloupe by Pegler (1983) also
probably represent P. sublaevigatus, due to the description
and illustrations presented. Supported by the molecular
(Figures 1 and 2) and the morphological data, Pluteus sub-
laevigatus belongs to section Celluloderma.
Pluteus xylophilus (Speg.) Singer, Lilloa 22: 405
(1951). (Figure 16)
≡ Entoloma xylophilum Speg., Bol. Ac. Nac. Cord. 28:
305, 1926.
Pileus 30–70(−110) mm diam, plane-convex, occa-
sionally slightly concave, sometimes slightly umbonate,
light brown (N60A60M50 to N
60A40M40), darker at the
centre (N80A50M50), paler and discolouring radially
towards the margin, margin sometimes splitting and
apparently not sulcate or striate. Lamellae remote, slightly
pinkish to pinkish, crowded, with lamellulae. Stipe 35–
60(−120) × 3–10(apex)–9–20(base) mm, tapering towards
the apex, rarely equal with 3 mm diam, central, sometimes
flexuous, cream with brown fibrils over the surface and
especially at base, longitudinally striate, with scanty myc-
elium at base. Odour, taste, and flesh colour not recorded.
Basidiospores [160/8/7] (5.0–)6.2–7.5(−8.7) × 3.7–6.2 µm
[Q = (1.21–)1.24–1.68(−1.74); Qm = 1.38; Lm = 6.7 µm;
Wm = 4.9 µm], broadly ellipsoid to ellipsoid, occasionally
elongate, inamyloid, hyaline, smooth, thick-walled, guttu-
late. Basidia (18.7–)21–26 × 6.2–8.7 µm, clavate or versi-
form, thin-walled, four-spored, with small scattered
guttules. Pleurocystidia of three types: (I) the normal
Cervinus-type, (46–)52–75(−87) × 12.5–23(−25) µm,
fusoid-ventricose, colourless, hyaline, apices usually with
two to six lateral prongs, moderately thick-walled espe-
cially in the apical region and tapering towards the base,
sparse, and abundant to very abundant, frequently with
adhered basidiospores on many pleurocystidial apices; (II)
the modified Cervinus-type as found in P. harrisii, usually
without prongs and subcapitate or slightly strangulated at
the apex, or sometimes with two short lateral prongs, size
similar to type I; (III) preponderantly of the Magnus-type,
usually shorter especially in length, 38–62 × (8.7–)11.0–28
µm, lageniform to fusoid, with acute apex or rarely with a
rounded apex, colourless, hyaline, thin- to moderately
thick-walled, rare. Cheilocystidia dimorphic, not abun-
dant, very rooting on hymenium, and hard to see: (I) short
vesiculose, clavate to subclavate, 21–37 × 8.7–15 µm; (II)
long-cylindrical-clavate, sometimes with a moderately
long pedicel, 40–69 × 10.0–15.0 µm; sometimes transitions
between the two extreme forms are found that are usually
slightly ventricose or versiform and moderately deformed.
Lamellar edge heteromorphous. Lamellar trama inverse, up
to 62 µm wide, composed of thin or slightly thick-walled
hyphae, sometimes moderately inflated, 1.2–13.7 µm diam,
hyaline, septate. Pileus context undifferentiated, approxi-
mately 187 µm thick, occasionally very thick more than
1,000 µm thick, composed of slightly thick-walled hyphae,
sometimes inflated, 3.7–21 µm diam, hyaline, septate.
Pileipellis a repent epicutis up to 125 µm thick, composed
of thin- to slightly thick-walled hyphae, 3.7–6.2 µm diam,
elongated, septate, with dissolved brown vacuolar content.
Clamp connections absent in all parts examined.
Figure 16. Pluteus xylophilus. A. Basidioma; A
1. Menolli Jr.
et al. NMJ138 ; A2. Menolli Jr. et al. NMJ143. B, C, D. Menolli
Jr. et al. NMJ143. B. Basidiospores. C. Basidia. D. Pleuro-
cystidia; D1. Normal Cervinus-type; D2. Modified Cervinus-
type; D3. Magnus-type. E. Cheilocystidia (Menolli Jr. et al.
NMJ138); E1. Short vesiculose; E2. Long-cylindrical-clavate.
Bars (A) = 1 cm; (B–E) = 10 µm.
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152 N. Menolli Jr. et al.
Habitat and habit: Solitary to scattered on wood.
Material examined: BRAZIL, São Paulo State, São
Paulo city, Parque Estadual das Fontes do Ipiranga,
10 Nov. 1982, G. Guzmán 22986 (SP); 16 Jan. 1987,
Pegler et al. 3712 (SP); 20 June 2006, F. Karstedt et al.
FK683 (SP); 30 May 2007, Menolli Jr. et al. NMJ138
(SP); 18 Oct. 2007, Menolli Jr. & F. Karstedt NMJ150
(SP); Parque Estadual da Cantareira, Núcleo Engordador,
21 Aug. 2007, Menolli Jr. et al. NMJ143 (SP); 23 Oct.
2008, M. Capelari & L.A.S. Ramos MC4397 (SP).
Comments: Pluteus xylophilus is characterised by a
moderately large pileus, three types of pleurocystidia, and
two types of cheilocystidia. The specimens herein studied
have surfaces usually with some cavities, probably caused
by insects, showing the white flesh.
Singer (1958) described two varieties, viz. P. xylophilus
var. tucumanensis Singer and P. xylophilus var. major
Singer, based on the size and pigmentation differences of the
pileus and the predominance of each cheilocystidia type.
However, he emphasises that it is necessary to check
whether each of these characters remains constant in succes-
sive fruitings and to verify if these morphological variations
constitute a forma, varieties, or independent microspecies.
Singer’s varieties are not considered in this work because
considerable morphological variations were not observed.
Singer (1956, 1958) accepted two varieties of P. cervi-
nus, viz. P. cervinus var. brasiliensis and P. cervinus var.
tucumanensis Singer, as synonyms of P. xylophilus, and
also considered P. cervinus sensu Rick (1938), which was
recorded for Brazil, as a probable P. xylophilus.
The geographic distribution of P. xylophilus includes
Argentina (Spegazzini 1925 as Entoloma xylophilum;
Singer and Digilio 1951 as P. cervinus var. tucumanensis
and P. cervinus var. brasiliensis; Singer 1958), Bolivia
(Singer 1958), Brazil (Bresadola 1920 as P. cervinus var.
brasiliensis; Grandi et al. 1984; Stijve and Meijer 1993 as
P. xylophilus and P. xylophilus var. tucumanensis; Pegler
1997; Meijer 2006, 2008) and Peru (Singer 1958).
The molecular analyses (Figures 1 and 2) confirm the
position of P. xylophilus in the section Pluteus and relate it
with the other species of the Cervinus-group, forming a
well-supported clade (86% BS, Figure 2).
Acknowledgements
The authors thank Dr Alfredo Justo (Clark University) for critical
comments and presubmission review of the manuscript;
Dr Tarciso S. Filgueiras (Reserva Biológica do IBGE) and Fran-
cisco Kuhar (FCEN Universidad de Buenos Aires) who kindly
revised the Latin diagnoses; the curators of DAOM, F, K, LAU
and LIL for the loan of collections; Dr Maria Helena Pelegrinelli
Fungaro (Universidade Estadual de Londrina) for some DNA
sequencing; Fernanda Karstedt (Instituto de Botânica) for col-
lecting some specimens; Klei R. Sousa for preparing the illustra-
tions; the Conselho Nacional de Desenvolvimento Científico e
Tecnológico (CNPq) for the support and grant to the first author;
and the Fundação de Amparo à Pesquisa do Estado de São Paulo
(FAPESP grant 04/04319-2) for financial support.
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