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Entoloma (Agaricales, Basidiomycota) is a species-rich genus with approximately 2000 species known worldwide. In Cen- tral America, however, information about the species of this genus is sparse, despite the generally high biodiversity in this region. Recently, 124 specimens of Entoloma were collected in Panama, Chiriquí Province. In the present publication, the morphology of 20 species represented by more than one specimen is described and depicted with photographs, line drawings, and scanning electron micrographs. Molecular phylograms based on ITS or concatenated ITS and partial nc LSU rDNA sequences are provided. The taxonomic status of these species is evaluated and 17 species of Entoloma are described as new to science. Only one species could be assigned to an already known species, viz. Entoloma belouvense. Nolanea albertinae, described from Brazil, appeared similar and is combined in E. belouvense on varietal level. The identifications of two fur- ther species are uncertain. At least 30 other species, including potentially new species, cannot formally be described due to insufficient material. A preliminary key to the species of the genus Entoloma in Panama is provided. The spatial shape of the polyhedroid basidiospores of Entoloma spp. is discussed based on literature and the micrographs generated for the present study. Our re-evaluations indicate that the type of polyhedroid basidiospore and the structure of its base are not reliable as diagnostic characters for the delimitation of subgenera in Entoloma.
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https://doi.org/10.1007/s11557-021-01752-2
ORIGINAL ARTICLE
Fungal diversity inthetropics: Entoloma spp. inPanama
KaiReschke1 · MachielE.Noordeloos2· CathrinManz1· TinaA.Hofmann3· JoséRodríguez‑Cedeño3·
BálintDima4· MeikePiepenbring1
Received: 30 August 2021 / Revised: 9 October 2021 / Accepted: 12 October 2021
© The Author(s) 2022
Abstract
Entoloma (Agaricales, Basidiomycota) is a species-rich genus with approximately 2000 species known worldwide. In Cen-
tral America, however, information about the species of this genus is sparse, despite the generally high biodiversity in this
region. Recently, 124 specimens of Entoloma were collected in Panama, Chiriquí Province. In the present publication, the
morphology of 20 species represented by more than one specimen is described and depicted with photographs, line drawings,
and scanning electron micrographs. Molecular phylograms based on ITS or concatenated ITS and partial nc LSU rDNA
sequences are provided. The taxonomic status of these species is evaluated and 17 species of Entoloma are described as new
to science. Only one species could be assigned to an already known species, viz. Entoloma belouvense. Nolanea albertinae,
described from Brazil, appeared similar and is combined in E. belouvense on varietal level. The identifications of two fur-
ther species are uncertain. At least 30 other species, including potentially new species, cannot formally be described due to
insufficient material. A preliminary key to the species of the genus Entoloma in Panama is provided. The spatial shape of the
polyhedroid basidiospores of Entoloma spp. is discussed based on literature and the micrographs generated for the present
study. Our re-evaluations indicate that the type of polyhedroid basidiospore and the structure of its base are not reliable as
diagnostic characters for the delimitation of subgenera in Entoloma.
Keywords Neotropic· Entolomataceae· Taxonomy· Phylogeny· New species
Introduction
The genus Entoloma inCentral America
The genus Entoloma (Agaricales, Basidiomycota) includes
species with almost exclusively polyhedroid, faceted
basidiospores, which are angled in outline and pinkish as
seen with a light microscope. It comprises mainly species
with agaricoid basidiocarps; few species develop secotioid
or gasteroid basidiocarps (Co-David etal. 2009; Kinoshita
etal. 2012; Vidal etal. 2016). Currently, approximately 2000
species of Entoloma are known to science (Noordeloos etal.
2018). Two monographic treatments of the genus exist for
two regions of North America (Hesler 1967; Largent 1994).
Such monographs are not available for South America, but
some preliminary and regionally comprehensive treatments
have been published (Horak 1978, 1982; Coimbra etal.
2013; Karstedt and Capelari 2013). Coimbra (2014) pub-
lished a checklist including 271 species of Entolomataceae
recorded for Central and South America. However, almost
nothing is known about Entoloma species in Panama. Only
one species, Entoloma cylindrocapitatum (T.J. Baroni &
Ovrebo) Noordel. & Co-David, has been reported until now
(Ovrebo and Baroni 2007; Hofmann and Piepenbring 2021).
Few Entoloma species are reported from the other Central
American countries, Belize, Costa Rica, El Salvador, Gua-
temala, Honduras, and Nicaragua: six species are reported
Section editor: Zhu-Liang Yang
* Kai Reschke
Reschke@em.uni-frankfurt.de
1 Mycology Research Group, Faculty ofBiological Sciences,
Goethe University Frankfurt Am Main, Max-von-Laue
Straße 13, 60438FrankfurtamMain, Germany
2 Naturalis Biodiversity Center, P.O. Box9517,
2300RALeiden, TheNetherlands
3 Centro de Investigaciones Micológicas (CIMi), Herbario
UCH, Universidad Autónoma de Chiriquí, 0427David,
ChiriquíProvince, Panama
4 Department ofPlant Anatomy, Institute ofBiology, Eötvös
Loránd University, Pázmány Péter sétány 1/c, 1117Budapest,
Hungary
Mycological Progress (2022) 21:93–145
1 3
from Costa Rica (Baroni and Halling 2000; Halling and
Mueller 2005), a single record exists for Belize (Largent
etal. 2008a), and no species are known for El Salvador,
Guatemala, Honduras, and Nicaragua (Coimbra 2014). From
Colombia, the South American neighbouring country of
Panama, six species of Entoloma spp. are reported (Franco-
Molano and Uribe-Calle 2000; Vasco-Palacios and Franco-
Molano 2013; Osorio and Osorio 2016).
Description ofthestudy region
The province of Chiriquí is located between 8° and 9° north
in the western part of Panama at the border to Costa Rica
(Fig.1). It forms part of the Mesoamerican biodiversity hot-
spot, one of the 25 biodiversity hotspots of the world rec-
ognised by Myers etal. (2000). There are several important
protected areas in Chiriquí including the “Parque Nacional
Volcán Barú” (PNVB) in the northern part of Chiriquí,
which includes the highest mountain in Panama (Volcán
Barú, 3475m asl.), and the “Parque Internacional La Amis-
tad” (PILA), which is listed as a UNESCO World Herit-
age Site and spans from Costa Rica to Panama. These two
parks are mainly covered by montane mixed forests with
Quercus spp. as the dominant trees, while at some locations
Alnus acuminata Kunth is dominant. The “Reserva Forestal
de Fortuna” is a mountainous forest reserve in the north-
eastern part of Chiriquí. It includes forests dominated by
Oreomunnea mexicana (Standl.) J.-F.Leroy in lower mon-
tane sites, while the higher montane forest is dominated by
Quercus spp. South of David, the capital of Chiriquí, there
are extensive mangroves and numerous small islands. The
island of Parida is located about 10km south of the coast and
forms the centre of the “Parque Nacional Marino Golfo de
Chiriquí”. The lowlands of Chiriquí are mainly covered by
cattle pastures and teak plantations. Chiriquí has a tropical
climate, with a rainy season from April/May to December
and a dry season from January to April. The annual rainfall
in David (49m asl.) is approximately 3900mm and the aver-
age temperature 25.2°C, in Boquete (1096m asl.) 3810mm
and 18.4°C, and in Volcán (1412m asl.) 4400mm and
16.9°C, respectively (https:// en. clima te- data. org accessed
10-june-2021). The dry season is less pronounced in the
mountains, with an average precipitation of approximately
60 and 70mm in the driest month, February, in Volcán and
Boquete, respectively, compared to 36mm in David.
Material andmethods
Fieldwork
Specimens of Entoloma spp. were collected by the authors
and collaborators from 2014 to 2018, mainly in montane for-
ests at 1600–2400m asl. dominated by Quercus spp., some-
times by Alnus acuminata. Some collections were made in a
forest dominated by Oreomunnea mexicana at 1200m asl.,
a forest dominated by Calophyllum longifolium Willd. at
40m asl. on Isla Parida, and close to a path in disturbed
secondary vegetation at 120m asl. in Los Algarrobos. Most
of the specimens were photographed in the field. Further
Fig. 1 Study locations in
Chiriquí Province, Panama
longitude (°E)
latitude (°N)
−83.0 −82.5 −82.0 −81.5
8.0 8.5
David
PILA
PNVB
94 Mycological Progress (2022) 21:93–145
1 3
photos were taken in the laboratory and ephemeral charac-
ters as colour, smell, and taste were registered. Colour terms
and codes were determined using Kornerup and Wanscher
(1967). Coordinates of the locations were recorded using
a Garmin GPSmap 62 (Garmin Deutschland, Garching,
Germany). The basidiocarps were dried at 40–45°C on an
electric food dehydrator and frozen for several days before
storage in herbaria. Mainly specimens of species represented
by at least two collections were studied in detail.
Light microscopy
Micromorphological characters of the basidiocarps were
analysed by light microscopy on dried material. The size of
the basidiospores, hymenial structures, and features of the
pileipellis were investigated in hand-cut sections mounted in
tap water or in 5–10% KOH. To measure the long cells of the
hymenophoral trama and stipe surface, sections were soaked
in KOH for 5min, rinsed with deionised water, and stained
with Congo Red. At least 20 basidiospores were measured
from lamellae squash preparations for each collection. For
some specimens, the presence or absence of clamp connec-
tions at the base of basidia was additionally examined using
phase contrast. Spore sizes and Q-values are given in 5th
percentile–mean–95th percentile. Spore sizes are rounded
to the nearest 0.5µm, Q-values to the nearest 0.05. Other
values of measurements were less strictly rounded off to
avoid pseudo-exact indication of sizes.
Scanning electron microscopy (SEM)
Fragments of basidiocarps were prepared for SEM following
a protocol including the use of dioctyl sodium sulfosucci-
nate (DSS), based on Koch etal. (2021) and Erbar (1995)
with several modifications. Small pieces of lamellae were
soaked in a solution of 5% (w/v) DSS in 4:1 water:ethanol
for 24h. The samples were then rinsed three times in 4:1
water:ethanol and two times in 0.1M sodium cacodylate
buffer and thereafter fixed in 4% glutaraldehyde in 0.1M
sodium cacodylate buffer for 20–24h. After fixation, the
samples were rinsed two times in the cadodylate buffer and
then dehydrated in a graded ethanol series of 30, 50, 70, 90,
100, and 100% for 10–20min each. The dehydrated sam-
ples were stored in 100% ethanol until critical point drying
using liquid CO2. Dried samples were sputtered with gold
and studied in a Hitachi (S 4500) scanning electron micro-
scope. The description of basidiospore structures follows
Pegler and Young (1979), except that polyhedron terminol-
ogy is used instead of their spore types. The earlier term
“dièdre basal” (Kühner and Boursier 1929) is used instead of
“dihedral base” to avoid confusion with mathematical terms.
DNA extraction andPCR
Pieces of lamellae of 3–10 mm2 were taken from dry basidi-
ocarps and ground in a MM301 Mixer Mill (Retsch GmbH,
Haan, Germany). DNA was extracted from the resulting
powder using the peqGOLD fungal DNA mini kit (VWR,
Darmstadt, Germany) or the innuPREP Plant DNA Kit (ana-
lytikjena, Jena, Germany) according to the instruction manu-
als. The ITS region was amplified by PCR in a peqSTAR
gradient thermal cycler (PEQLAB, Erlangen, Germany)
using the VWR Taq DNA polymerase (VWR, Darmstadt,
Germany). To obtain the ITS sequences, the forward primer
ITS1 or ITS1F and the reverse primer ITS4 or ITS4B (White
etal. 1990; Gardes and Bruns 1993) were used. Further ITS
sequences were obtained as described by Papp and Dima
(2018). To obtain the D1/D2 region of the nc LSU rDNA,
the primers LR0R together with LR5 (Vilgalys and Hester
1990) or NL1 together with NL4 (O’Donnell 1992) were
used. Both rDNA regions were amplified using the same
PCR protocol: denaturation at 98°C for 4min followed by
35 cycles of 95°C for 45s, 53°C for 30s, and 72°C for
60s, with a final elongation step at 72°C for 5min. A partial
sequence of the second largest subunit of RNA polymer-
ase II (RPB2) was obtained using the primers rpb2-6F and
rpb2-7.1R (Matheny 2005) with a touchdown PCR protocol:
denaturation at 95°C for 4min followed by 14 cycles of
94°C for 45s, 56°C (–0.5°C/cyc) for 60s, and 72°C for
60s, thereafter 40 cycles of 94°C for 30s, 53°C for 40s,
and 72°C for 60s, with a final elongation step at 72°C
for 10min. Success of amplification was checked by gel
electrophoresis using a 1% (w/v) agarose gel. Successfully
amplified products were sent to Microsynth Seqlab (Göt-
tingen, Germany) for purification and Sanger sequencing.
Phylogenetic analyses
Sequences were edited with Geneious 2019.2.1 (Biomat-
ters Ltd., Auckland, New Zealand) and aligned with Mafft
(Katoh and Standley 2013) using the E-INS-i algorithm. The
ends of the resulting alignments as well as unreliable termi-
nal parts of sequences were manually pruned in AliView
(Larsson 2014). For the phylogenetic analysis of sequences
of subg. Cyanula, aligned ITS and LSU sequences were
concatenated and a piece of 61 nucleotides at the 5ʹ end
of the LSU, which was not available for many sequences,
excluded from the final alignment. Maximum likelihood
trees were built using RAxML 8.2.11 (Stamatakis 2014).
The GTRGAMMA model was used for alignments of less
than 50 sequences, the GTR CAT model with 45 per site
rate categories for 50 sequences, and more, along with a
bootstrap analysis (Felsenstein 1985) with 500 repetitions.
Resultant phylogenetic trees were visualised using FigTree
95Mycological Progress (2022) 21:93–145
1 3
(Rambaut 2014). All sequences generated are deposited in
GenBank (Table1).
Results
A total of 124 specimens of Entoloma spp. were collected.
These specimens represent more than 50 species, with at
least 22 species being represented by more than one speci-
men. Twenty of these species were studied in detail and are
described below. Only one species could be assigned to
an already described species and two species were identi-
fied with considerable uncertainty. Seventeen species are
unknown to science and are described as new species below.
The most frequently represented subclade was the subgenus
Cyanula, with 35 specimens belonging to 18 species. Speci-
mens of the subgenera Alboleptonia, Cubospora Entoloma,
and Nolanea were frequently found as well (Table2).
In 2017 and 2018, most specimens were collected in mon-
tane forests with abundant to dominating trees of Quercus
spp. The highest number of Entoloma specimens for a sin-
gle location, i.e. 22 specimens found during three visits,
was collected in the PILA, in a montane forest dominated
by Alnus acuminata, with some Quercus spp. and at least
one individual of Podocarpus sp. On the contrary, only two
specimens, both of subg. Cyanula, were collected in lowland
habitats, one in a forest close to mangrove forest on Island
Parida and the other one at a path in disturbed secondary
vegetation (Table3). This path in Los Algarrobos was the
most often visited location in these years. The specimens
collected in the years before 2017 were mainly collected
during a local inventory project and are thus less informative
for comparisons of diversity of habitats.
Shape ofbasidiospores
Polyhedroid basidiospores of 17 species of Entoloma from
Panama were analysed by SEM. The basidiospores displayed
a considerable degree of variability, increasing with the
number of facets. Apart from the cuboid basidiospores, the
number of angles of the respective facets was often variable
and thus not determined. The adaxial facet is always a single
facet and consistent in its position above the apiculus. The
spatial formation of facets can abaxially be variable. The
basal facet can either be next to the abaxial facet or sepa-
rated from it by lateral facets (Fig.2e). When two abaxial
facets are formed, they can be next to each other or sepa-
rated by lateral facets (Figs.2i, 26d). Shape and a consistent
base type of the basidiospores of two species of subg. Ento-
loma (E. nuboocculatum and E. mediorobustum, described
below) could not be determined because of considerable
variability of these characters. Five species have hendeca-
hedroid spores, three of subg. Cyanula and two of subg.
Nolanea. Two species each have heptahedroid, octahedroid,
or decahedroid basidiospores, respectively, while one spe-
cies each presents cuboid, enneahedroid, dodecahedroid, or
tetrahedroid basidiospores. Reduced forms frequently occur,
mainly concerning reduction or lack of the apical facet and/
or reduction of the pair of adaxial facets to a single adaxial
facet. Basidiospore shapes in subgenera are not consistent.
The largest variability was observed in subg. Alboleptonia,
including one species with mainly octahedroid, one species
with enneahedroid, and another species with tetradecahe-
droid basidiospores. A simple base is formed in four spe-
cies, two in subg. Entoloma and two in subg. Nolanea,
while basidiospores with a dièdre basal are present in all
subgenera, exclusively so in Alboleptonia, Cubospora, and
Cyanula. A base built by three facets was found in the newly
described species E. mediorobustum (Fig.2l).
Taxonomy
The traditional infrageneric classification of Entoloma is
based on morphological characters, viz., habit, lamellae
attachment, pileipellis structure, and basidiospore shape
(Romagnesi 1974; Romagnesi and Gilles 1979). This clas-
sification has been continuously revised and emended
(Noordeloos 1981, 1992, 2004; Largent 1994). Co-David
etal. (2009) demonstrated that a high number of subgroups
in this classification were not monophyletic based on molec-
ular phylogenetic results. With the incorporation of DNA
sequence data, several subgenera have since been revised,
viz., Pouzarella by He etal. (2013), Leptonia by Morozova
etal. (2014a), Entoloma (as “Rhodopolia”) by Kokkonen
(2015), and Claudopus by He etal. (2019). The section
Cyanula was elevated to the rank of subgenus by Noordeloos
and Gates (2012) and two new subgenera, Cubospora and
Cuboeccilia, were described by Karstedt etal. (2019). He
etal. (2019) recognised 10 major clades, largely referring
to the studies just cited. Baroni etal. (2011) described the
genus Entocybe for a few aberrant species of the large, basal
clade of Entoloma. This new genus, however, causes the
basal clade to be paraphyletic and was, therefore, not recog-
nised by He etal. (2019).
Subgenus Entoloma Fig.3
As in most other subgenera, species of subgenus Entoloma
are hard to identify on macroscopical characters in the field,
as they are pretty uniform in colour and stature. Kühner and
Romagnesi (1953, 1954) first considered the structure and
pigmentation of the covering layer of the pileus as potential
tools to distinguish species, besides size and shape of basidi-
ospores. Noordeloos (1981, 1992, 2004) expanded the con-
cept, resulting in an overview of sect. Rhodopolia in Europe
96 Mycological Progress (2022) 21:93–145
1 3
Table 1 Data corresponding to the sequences used for phylogenetic analyses, including GenBank accessions, if not stated otherwise
Species (annotation) Specimen voucher Origin Accession Reference
ITS LSU
subgenus Alboleptonia
Alboleptonia sericella Lueck1 Germany KP965765 Karich etal. 2015
Entoloma amistadosericellum KaiR490 Panama MZ611648 This study
Entoloma amistadosericellum KaiR492 Panama MZ611649 This study
Entoloma amistadosericellum KaiR495 Panama MZ611650 This study
Entoloma amistadosericellum KaiR501 Panama MZ611651 This study
Entoloma amistadosericellum KaiR502 Panama MZ611652 This study
Entoloma amistadosericellum KaiR612, holotype Panama MZ611663 This study
Entoloma amistadosericellum KaiR672 Panama MZ611673 This study
Entoloma amistadosericellum KaiR675 Panama MZ611674 This study
Entoloma confusissimum KaiR489 Panama MZ611647 This study
Entoloma confusissimum KaiR670 Panama MZ611672 This study
Entoloma niveum PDD104142 New Zealand MZ611695 This study
Entoloma nubilosilvae KaiR429 Panama MZ611643 This study
Entoloma nubilosilvae KaiR430 Panama MZ611644 This study
Entoloma nubilosilvae KaiR465, holotype Panama MZ611645 This study
Entoloma nubilosilvae KaiR474 Panama MZ611646 This study
Entoloma sericellum KaiR1203 Sweden MZ611636 This study
Entoloma sericellum KaiR1214 Sweden MZ611637 This study
Entoloma sericellum LE254362 Russia KC898453 Morozova etal. 2014a
Entoloma serrulatum LE254361 Russia KC898447 Morozova etal. 2014a
Subgenus Cubospora
Entoloma aurantiovirescens KaiR623, holotype Panama MZ611665 This study
Entoloma aurantiovirescens PAN419 Panama MZ611691 This study
Entoloma luteum GDGM27698 China JQ281486 He etal. 2012
Entoloma murrayi MHHNU30602 China MK250917 Zhang and Chen,
unpublished
Entoloma murrayi QI1001 China KJ658967 He etal. 2015b
Entoloma murrayi QI1002 China KJ658968 He etal. 2015b
Entoloma murrayi SDR NAMA 2017–160 USA, Wisconsin MK575459 Russell, unpublished
Entoloma quadratum LE254355 Russia KC898452 Morozova etal. 2014a
Entoloma quadratum MHHNU30632 China MK250921 Zhang and Chen,
unpublished
Entoloma quadratum MHHNU82 China KU518319 He and Chen, unpub-
lished
Entoloma quadratum PAN241 Panama MZ611690 This study
Entoloma semilanceatum NS2283 Cameroon MN069544 Largent etal. 2019
Entoloma virescens MHHNU30619 China MK250919 Zhang and Chen,
unpublished
Inocephalus murrayi ECO-TA-HO 7874 Mexico MF156254 Arias-Hernandez
etal., unpublished
Subgenus Cyanula
Entoloma aff. griseocyaneum NL5097 USA MZ869020 MK277993 This study/Varga etal.
2019
Entoloma aff. griseocyaneum PO6 Canada, Ontario KY706188 Hay etal. 2018
Entoloma aff. necopinatum KaiR646 Panama MZ611670 MZ678748 This study
Entoloma arcanum KaiR614 Panama This study
Entoloma arcanum KaiR488, holotype Panama MZ678738 This study
+ RPB2: MZ605445
97Mycological Progress (2022) 21:93–145
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Table 1 (continued)
Entoloma caeruleomarginatum CME3 Panama MZ611627 This study
Entoloma caeruleomarginatum KaiR535, holotype Panama MZ611658 This study
Entoloma caeruleomarginatum CME11 Panama MZ611622 This study
Entoloma cf. catalaunicum E163 Estonia Unite: UDB011680 Liiv, unpublished
Entoloma cf. indutoides TU106924 Estonia Unite:
UDB015261
Liiv, unpublished
Entoloma cf. largentii OSC144006 USA KX574458 Gordon, unpublished
Entoloma cf. pseudoturci Cro16 Croatia MZ611633 This study
Entoloma cf. unicolor PBM3995 USA KY777373 Matheny etal.,
unpublished
Entoloma cf. violaceobrunneum KaiR632 Panama MZ678740 This study
Entoloma cf. violaceobrunneum CME2 Panama This study
Entoloma coracis O-F-256850, holotype Norway MW934571 MW934251 Crous etal. 2021
Entoloma fuscosquamosum KY744158 USA KY744158 Matheny etal.,
unpublished
Entoloma glaucobasis Roth 16–6-95/NL-2704 Germany/Sweden MZ869021 MK277991 This study/Varga etal.
2019
Entoloma griseocaeruleum CME8 Panama MZ611631 This study
Entoloma griseocaeruleum CME13 Panama MZ611624 This study
Entoloma griseocaeruleum KaiR536 Panama MZ611659 This study
Entoloma griseocaeruleum KaiR534, holotype Panama MZ611657 This study
Entoloma griseocyaneum KaiR997 Germany MZ611684 This study
Entoloma holmvassdalenense O-F-304575 Norway MZ869018 MZ678746 This study
Entoloma holmvassdalenense O-F75311, holotype Norway KM610321 Weholt etal. 2014
Entoloma incanum KaiR990 Germany MZ611683 This study
Entoloma indutoides WU19603 Austria MZ869022 This study/Varga
etal. 2019
Entoloma inocephalum LE262922 Vietnam KC898449 MH259311 Morozova etal.
2014a/Karstedt
etal. 2019
Entoloma isborscianum KaiR1004 Germany MW934565 Crous etal. 2021/This
study
Entoloma longistriatum PBM4018 USA KY744164 Matheny etal.,
unpublished
Entoloma mediterraneense Cro26 Croatia MZ611634 This study
Entoloma melleosquamulosum KaiR638 Panama MZ611669 This study
Entoloma melleosquamulosum CME16, holotype Panama MZ611626 This study
Entoloma microserrulatum KaiR413, holotype Panama MZ611642 This study
Entoloma microserrulatum KaiR664 Panama MZ611671 This study
Entoloma montanum O-F-293389/N03-09–2010 Norway/Norway MW340878 MZ678747 Noordeloos etal.
2021/This study
Entoloma montanum GB191635, holotype Sweden MW340896 Noordeloos etal.
2021
Entoloma mougeotii LE254352 Russia, Caucasus KC898446 Morozova etal. 2014a
Entoloma nigrovelutinum LE295077, holotype Vietnam MF898426 MF898427 Crous etal. 2017
Entoloma nipponicum TNS-F70747 Japan MK693223 MK696392 Crous etal. 2019
Entoloma norlandicum O-F-76176, holotype Norway MW340899 Noordeloos etal.
2021
Entoloma ochromicaceum TUF120040/DMS-9201008 Estonia/Denmark Unite:
UDB023715
MZ678743 Liiv, unpublished/
This study
Entoloma odoratum DMS-166826 Denmark MZ869017 MZ678745 This study
Entoloma querquedula TUR Finland LN850627 Kokkonen 2015
Entoloma roseotinctum JL-26–19 Norway MZ869019 This study
98 Mycological Progress (2022) 21:93–145
1 3
Table 1 (continued)
Entoloma sarcitulum K378, holotype Great Britain LN850561 Kokkonen 2015
Entoloma serrulatum O-F-158208/DMS-730296 Norway/Denmark MZ869016 This study
Entoloma sp. PO2 Canada, Ontario KY706185 Hay etal. 2018
Entoloma sp. CM13_233 New Caledonia KY774214 Carriconde etal.,
unpublished
Entoloma sp. MES-534 Chile KY462681 Truong etal. 2017
Entoloma sp. KaiR511 Panama MZ611655 This study
Entoloma sp. OTA61944 New Zealand KP191935 Lebel and Cooper,
unpublished
Entoloma sp. CY13_140_1 New Caledonia KY774215 Carriconde etal.,
unpublished
Entoloma subcaesiocinctum GDGM31059 China KY972699 He etal. 2017
Entoloma subcoracis LE312483, holotype Russia MW934593 MW934255 Crous etal. 2021
Entoloma subfarinaceum SAT1518702 USA KY777374 Matheny etal.,
unpublished
Entoloma subserrulatum MGW1490 USA KY744177 Matheny etal.,
unpublished
Entoloma subserrulatum EL9 Canada KY706167 Hay etal. 2018
Entoloma turci WPR004 Germany MZ611693 This study
Entoloma viiduense G1602 Estonia UDB015211 MK278008 Liiv, unpublished/
Varga etal. 2019
Entoloma violaceoserrulatum TUR JV 8329F, isotype Finland MF476913 MF487803 Morozova etal. 2017
Entoloma yanacolor QCAM6312, holotype Ecuador MG947210 Crous etal. 2018
Subgenus Entoloma
Entoloma aff. alpicola O:F7195 Norway MZ868961 This study
Entoloma aff. rhodopolium 68–88 Finland LN850500 Kokkonen 2015
Entoloma aff. sinuatum TRTC156546 Canada JN021019 Dentinger etal. 2011
Entoloma aff. sordidulum PGL110817 Norway MZ869007 This study
Entoloma alnobetulae K53-59, G262083,
holotype
France LN850537 Kokkonen 2015
Entoloma alpicola O:F-61507 Norway MZ868967 This study
Entoloma atrosericeum GG160811 Sweden MZ868968 This study
Entoloma atrosericeum K69-310, G262084,
holotype
France LN850607 Kokkonen 2015
Entoloma aurorae-borealis O-F-254651, holotype Norway MH234486 Noordeloos etal. 2021
Entoloma baeosporum holotype France MZ868973 This study
Entoloma bisporigerum 2047, K108895, holotype United Kingdom LN850536 Kokkonen 2015
Entoloma bisporigerum OW-E20-14 Norway MZ868976 This study
Entoloma boreale 106/09, TUR, holotype Finland LN850624 Kokkonen 2015
Entoloma borgenii TB79.120, C5933 Greenland LN850524 Kokkonen 2015
Entoloma brunneipes TENN9029, holotype USA LN850621 Kokkonen 2015
Entoloma brunneocinereum TENN12881, holotype USA LN850615 Kokkonen 2015
Entoloma brunneorugulosum KaiR583 Panama MZ611660 This study
Entoloma brunneorugulosum KaiR691, holotype Panama MZ611676 This study
Entoloma brunneorugulosum KaiR692 Panama MZ611677 This study
Entoloma brunneorugulosum KaiR695 Panama MZ611679 This study
Entoloma brunneorugulosum PAN532 Panama MZ611692 This study
Entoloma bryorum 117/07 Finland LN850539 Kokkonen 2015
Entoloma bryorum O:F-303879 Norway MZ868978 This study
Entoloma caccabus “Podoba”, G262081 France LN850540 Kokkonen 2015
Entoloma caccabus C157, 4226 The Netherlands MZ868979 This study
Entoloma caccabus C158, 4001 The Netherlands MZ868981 This study
99Mycological Progress (2022) 21:93–145
1 3
Table 1 (continued)
Entoloma carolinianum TENN21855, holotype USA LN850617 Kokkonen 2015
Entoloma cf. alpicola 23, N1F31, 2 NA HQ445607 Bjorbækmo etal.
2010
Entoloma cf. alpicola O:F-260165 Norway MZ869001 This study
Entoloma cf. bryorum O:TEB 395–18 Norway MZ868985 This study
Entoloma cf. carolinianum IBUG:Montanez222 Mexico MZ869010 This study
Entoloma cf. rhodopolium DB03-06–2018-3 Hungary MZ868960 This study
Entoloma cf. spadiceum TENN070791 USA, Tennessee MF686497 Matheny and
Sanchez-Garcia,
unpublished
Entoloma cf. venosum WU19672 MZ868991 This study
Entoloma cistophilum Trim2006, isotype France MZ868999 This study
Entoloma dulciodorans IBUG:LGD7580 Mexico MZ869011 This study
Entoloma dulciodorans IBUG:Rodriguez322,
paratype
Mexico MZ869012 This study
Entoloma eminens 417/12, TUR, holotype Finland LN850584 Kokkonen 2015
Entoloma eminens O:F-248409 Norway MZ868982 This study
Entoloma eminens TEB699-17 Norway MZ868983 This study
Entoloma fasciculatum TENN29376, holotype USA LN850614 Kokkonen 2015
Entoloma festivum holotype The Netherlands MZ869009 This study
Entoloma gerriae holotype The Netherlands MZ868980 This study
Entoloma gerriae WU28035 Austria MZ868990 This study
Entoloma griseopruinatum JLC_030924-8, holotype France MZ868984 This study
Entoloma griseopruinatum JLC030924-8, isotype France LN850556 Kokkonen 2015
Entoloma griseorugulosum RFS-020921–01, isotype Spain LN850591 Kokkonen 2015
Entoloma indutoides 1617, K108968, holotype United Kingdom LN850608 Kokkonen 2015
Entoloma inusitatum holotype Germany MZ868969 This study
Entoloma jubatum PAM13-26 Sweden MZ868975 This study
Entoloma lactarioides Liiv217, holotype Russia, Karelia MZ869000 This study
Entoloma lacus KUB110, holotype Japan LC088049 Kondo etal. 2017
Entoloma leucocarpum MEN701, holotype The Netherlands MZ868970 This study
Entoloma lividoalbum GE12-025 France MZ868957 This study
Entoloma lividomurinum TENN28208, holotype USA LN850616 Kokkonen 2015
Entoloma lupinum 13/14, TUR, holotype Finland LN850570 Kokkonen 2015
Entoloma majaloides 1168, K69959, holotype United Kingdom LN850489 Kokkonen 2015
Entoloma majaloides GG160812 Sweden MZ868966 This study
Entoloma mediorobustum JR064, holotype Panama MZ611635 This study
Entoloma mediorobustum PA728 Panama MZ611688 This study
Entoloma melenosmum JHP-338, C6229, holo-
type
Greenland LN850508 Kokkonen 2015
Entoloma myriadophyllum 366VN10 MZ869004 This study
Entoloma myrmecophilum IBUG:Quezada 1 Mexico MZ869014 This study
Entoloma myrmecophilum IBUG:SantosBarba 5 Mexico MZ869013 This study
Entoloma myrmecophilum var.
coalescens holotype Germany MZ868958 This study
Entoloma nidorosum GG160815 Sweden MZ868997 This study
Entoloma nidorosum s. MEN FR2016708 France MZ868995 This study
Entoloma nidorosum s. MEN MEN2012111 The Netherlands MZ868996 This study
Entoloma nigrobrunneum TENN9100, holotype USA LN850618 Kokkonen 2015
Entoloma noordeloosii WU18780, holotype Austria MZ868962 This study
Entoloma noordeloosii WU39739 MZ868963 This study
Entoloma nuboocculatatum KaiR588 Panama MZ611661 This study
100 Mycological Progress (2022) 21:93–145
1 3
Table 1 (continued)
Entoloma nuboocculatatum KaiR687, holotype Panama MZ611675 This study
Entoloma nubooccultatum PA165 Panama MZ611685 This study
Entoloma nubooccultatum PA696 Panama MZ611686 This study
Entoloma nubooccultatum PA727 Panama MZ611687 This study
Entoloma nubooccultatum PA864 Panama MZ611689 This study
Entoloma olorinum G:00,126,192, holotype France MZ868998 This study
Entoloma paludicola K178125, holotype United Kingdom LN850517 Kokkonen 2015
Entoloma palustre 101/14, TUR, holotype Finland LN850592 Kokkonen 2015
Entoloma paragaudatum 383/08, TUR, holotype Finland LN850530 Kokkonen 2015
Entoloma philocistus WU18871, paratype Portugal LN850600 Kokkonen 2015
Entoloma platophylloides MD2017-10 Italy MZ868959 This study
Entoloma politum 289/09 Finland LN850511 Kokkonen 2015
Entoloma politum O:F61875 Norway MZ869002 This study
Entoloma politum s. Gulden O:F-73841 Norway MZ869006 This study
Entoloma pruinosum CME12 Panama MZ611623 This study
Entoloma pruinosum CME14 Panama MZ611625 This study
Entoloma pruinosum KaiR533, holotype Panama MZ611656 This study
Entoloma pseudorhodopolium KUB102, holotype Japan LC088042 Kondo etal. 2017
Entoloma pusillulum C-F-5917 Greenland MZ611694 This study
Entoloma radicipes 42/14, TUR, holotype Finland LN850585 Kokkonen 2015
Entoloma rhodopolium 213/14, TUR, neotype Sweden LN850494 Kokkonen 2015
Entoloma rhodopolium Montanez149-150 Mexico MZ869015 This study
Entoloma rhodopolium O:F-254019 Norway MZ868993 This study
Entoloma rhodopolium OF254444 Norway MZ868992 This study
Entoloma rigidulum PRM153709, holotype Czech Republic LN850629 Kokkonen 2015
Entoloma rivulare 703/12, TUR, holotype Finland LN850544 Kokkonen 2015
Entoloma roseoalbum ARN6209, holotype The Netherlands MZ868972 This study
Entoloma rubrobasis 24,101, TURA Finland LN850580 Kokkonen 2015
Entoloma rubrobasis O:F-304601 Norway MZ868987 This study
Entoloma sarcopum KUB205 Japan LC088067 Kondo etal. 2017
Entoloma saussetiense PC0085778, holotype France LN850594 Kokkonen 2015
Entoloma sericatum 237/11 Finland LN850445 Kokkonen 2015
Entoloma sericatum 358/08 Finland LN850444 Kokkonen 2015
Entoloma serpens 410/09, TUR, holotype Finland LN850526 Kokkonen 2015
Entoloma silvae-frondosae L-DB6568 Hungary: Vas, Apatist-
vanfalva
MH790432 Crous etal. 2018
Entoloma sinuatum J.Vauras 8181F Finland KC710116 Morgado etal. 2013
Entoloma sinuatum J.Wisman, 2003–09-19 NA KC710109 Morgado etal. 2013
Entoloma sordidulum Co-David 2003 NA KC710062 Morgado etal. 2013
Entoloma sordidulum s. MEN ALV8406 Norway MZ869008 This study
Entoloma sp. 46-18A MZ868994 This study
Entoloma sp. EL145-18 Sweden MZ869005 This study
Entoloma sp. JO160811 Sweden MZ868971 This study
Entoloma sp. KBEB83, 17 Norway MZ868988 This study
Entoloma sp. MEN2012125 The Netherlands MZ868977 This study
Entoloma sp. N128 Irmgard6540 USA MZ868964 This study
Entoloma sp. O:F304982 Norway MZ869003 This study
Entoloma sp. UBC, F-32218 Canada, BC MF955123 Berbee etal., unpub-
lished
Entoloma subarcticum JHP-392, isotype Greenland MZ868974 This study
101Mycological Progress (2022) 21:93–145
1 3
Table 1 (continued)
Entoloma subradiatum “rhosse”, G262076,
lectotype
France LN850596 Kokkonen 2015
Entoloma subrhodopolium KUB1, holotype Japan LC088033 Kondo etal. 2017
Entoloma svalbardense GG310/86, O74756,
holotype
Svalbard LN850610 Kokkonen 2015
Entoloma svalbardense MENJG228 Norway, Svalbard JF304386 Geml etal. 2012
Entoloma tiliae LE254179 Russia, Saint Petersburg MH790420 Crous etal. 2018
Entoloma tiliae O:F-251963 Norway MZ868986 This study
Entoloma transvenosum MEN1215, holotype Denmark MZ868965 This study
Entoloma venosum MD2012-15 Germany MZ868989 This study
Rhodophyllus myrmecophilus holotype France LN850462 Kokkonen 2015
uncultured Entoloma NA Mexico KY574307 Garibay-Orijel and
Garcia-Guzman,
unpublished
uncultured fungus NA USA EU807287 Hanson etal., unpub-
lished
subgenus Nolanea
Entoloma belouvense var. bel-
ouvense WU27132, holotype France, Reunion MZ611698 This study
Entoloma belouvense var.
albertinae CME5 Panama MZ611628 This study
Entoloma belouvense var.
albertinae KaiR630 Panama MZ611668 This study
Entoloma cetratum LE235480 Russia KC898450 Morozova etal. 2014a
Entoloma clandestinum KaiR1282 Sweden MZ611639 This study
Entoloma conferendum KaiR1342 Austria MZ611640 This study
Entoloma conferendum KaiR1358 Germany MZ611641 This study
Entoloma conferendum KaiR978 Germany MZ611680 This study
Entoloma cremeostriatum CME10 Panama MZ611621 This study
Entoloma cremeostriatum KaiR508 Panama MZ611654 This study
Entoloma cremeostriatum KaiR613, holotype Panama MZ611664 This study
Entoloma euchroum KR-M-0032474 Germany KC898421 Morozova etal. 2014a
Entoloma flavoconicum KaiR609 Panama MZ611662 This study
Entoloma flavoconicum KaiR628, holotype Panama MZ611667 This study
Entoloma infula KaiR988 Germany MZ611681 This study
Entoloma infula KaiR989 Germany MZ611682 This study
Entoloma luteofuscum K-M-188309, holotype India KF698730 Raj etal. 2014
Entoloma olivaceohebes WU17841, holotype Italy JX454804 Vila etal. 2013
Entoloma olivaceohebes WU20798 Italy JX454803 Vila etal. 2013
Entoloma paraconferendum CME6, holotype Panama MZ611629 This study
Entoloma paraconferendum CME7 Panama MZ611630 This study
Entoloma piceinum LE254131, holotype Russia KM262035 Morozova etal. 2014b
Entoloma pulchellum HMLD1300 China KC257436 Wang and Bau, unpub-
lished
Entoloma pulchellum KA12-1242 South Korea KR673495 Kim etal. 2015
Entoloma readiae PDD71294 New Zealand MZ611696 This study
Entoloma readiae PDD87270 New Zealand MZ611697 This study
Entoloma sericeum KaiR1267 Sweden MZ611638 This study
Entoloma sp. KaiR626 Panama MZ611666 This study
Entoloma sp. KaiR693 Panama MZ611678 This study
Entoloma subelegans WU32902, holotype Mauritius MZ611699 This study
Entolomatransitionisporum CME9 Panama MZ611632 This study
102 Mycological Progress (2022) 21:93–145
1 3
with species distinguished by a combination of macro- and
microscopical characters. Since an ectomycorrhizal lifestyle
was discovered for species of subg. Entoloma (Antibus etal.
1981; Loree etal. 1989; Agerer 1997), ecological charac-
ters are also considered to delimit species. Kokkonen (2015)
used ITS sequence data as a new tool to tackle concepts of
species in subg. Entoloma in Northern Europe. New species
were defined based on a combination of morphological and
ecological characteristics combined with an ITS phylogeny.
As a result, nine species were described as new to science
and many species were synonymised. Ongoing studies in
Europe (Brandrud etal. 2018; Noordeloos etal. 2018) show
that there are still many questions as how to interpret clas-
sical species, as well as how to deal with apparently cryptic
speciation in this clade. Montañez etal. (2016) presented
six species of subg. Entoloma collected in Quercus-Pinus
forests in central Mexico. Also, Largent (1994) described
species of subg. Entoloma from Quercus-dominated habi-
tats. As Quercus spp. migrated from North America south-
wards and reached Central America approximately 10 mil-
lion years ago (Kappelle etal. 1992; Hooghiemstra 2006),
there should be links to species described from Mexico or
southern USA. Unfortunately, hardly any molecular data of
the species described from these regions are available for
comparison and we must rely on morphological comparisons
for several species.
Entoloma brunneorugulosum Reschke, Noordel. &
Lotz-Winter, sp. nov., Figs.4 and 5
Mycobank number: MB840935.
Typification: PANAMA. CHIRIQUÍ: near Volcán, Paso
Ancho, PNVB, forest trail towards summit of volcano,
Table 1 (continued)
Entolomatransitionisporum KaiR503, holotype Panama MZ611653 This study
Entoloma vernum LE312418 Russia MF476911 Morozova etal. 2017
Nolanea albertinae FK0912 Brazil KF679349 Karstedt etal. 2020
Nolanea albertinae FK0935, holotype Brazil KF679348 Karstedt etal. 2020
Nolanea albertinae FK1731 Brazil KF679350 Karstedt etal. 2020
Nolanea albertinae FK1732 Brazil KF679351 Karstedt etal. 2020
Nolanea atropapillata FK0898, holotype Brazil KF679354 Karstedt etal. 2020
Nolanea pallidosalmonea FK0891 Brazil KF738923 Karstedt etal. 2020
Nolanea parvispora FK1140 Brazil KF679353 Karstedt etal. 2020
Nolanea sp. FLAS-F-61537 USA MH211959 Kaminsky etal.,
unpublished
Nolanea tricholomatoidea FK1049, holotype Brazil KF679352 Karstedt etal. 2020
Table 2 Infrageneric positions of the specimens collected between
2014 and 2018
Subclade Specimens Number
of spe-
cies
Subg. Cyanula 35 18
Subg. Entoloma 23 5
Subg. Alboleptonia 17 3
Subg. Nolanea 15 7
Subg. Cubospora 11 6
Subg. Pouzarella 5 5
Subg. Inocephalus 3 3
Sect. Calliderma 3 1
Subg. Claudopus 1 1
Not determined 11 > 5
Table 3 Specimens per habitat
in the years 2017 and 2018 Habitat Specimens
Montane forest with abundant to dominant Quercus spp. 47
Montane forest dominated by Alnus acuminata 22
Montane forest with abundant Quercus spp. and Comarostaphylis arbutoides 9
Transition of submontane tropical forest dominated by Oreomunnea mexicana to montane
Quercus-dominated forest
2
Montane tropical forest 1
Disturbed lower montane forest with Quercus spp. and Alnus acuminata 1
Lowland forest close to mangrove 1
Lowland, path in disturbed secondary vegetation 1
Total 84
103Mycological Progress (2022) 21:93–145
1 3
approximately 2000m asl., montane forest with abundant
Quercus spp. and Comarostaphylis arbutoides, 28 June
2017, B. Wergen, KaiR691, holotype (M).
Etymology: brunneus (Latin) = brown, rugosus
(Latin) = rugose; refers to colour and structure of the pileal
surface.
Description: Basidiocarps tricholomatoid. Pileus
3.0–5.0cm diameter, convex when young, becoming
plane to depressed in old basidiocarps, usually with an
obtuse umbo, margin initially deflexed, then straight,
blackish brown to dark yellowish brown (5F8) in the
centre, dark brown to light brown (6D7, 6E7, 6F7, 6F8),
olive brown (4F8), yellowish brown (5E7, 5E8) to grey-
ish yellow (4C6, 4C7) towards the margin, pileal surface
glabrous, with a waxy appearance, rugulose, slightly
translucently striate at the margin up to a quarter of the
radius in fresh basidiocarps. Lamellae adnate to sinuate,
segmentiform to ventricose, rather distant, relatively thick
and broad, initially whitish with greyish to cream tinge,
later greyish pink, with smooth, concolorous edges. Stipe
3.5–4.0 × 0.6–0.9cm, cylindrical, solid in young basidi-
ocarps, becoming hollow, basically yellowish brown (5E5,
5E6, 5E7) to olive brown (4E6, 4E7, 4E8, 4F6, 4F7),
overlaid by whitish fibrils, paler, almost white, towards
base, whitish fibrillose-flocculose at the apex when young.
Basal mycelium white. Odour and taste farinaceous to fari-
naceous rancid.
Basidiospores 8.5–9.5–11.0 × 7.0–7.9–9.0 µm,
Q = 1.05–1.20–1.35 (n = 85 spores of 4 specimens), deca-
hedroid, composed of an adaxial facet (D), a pair of apico-
adaxial facets (AD), an apical facet (A), an abaxial facet
(B), two pairs of lateral facets (L1–L2), and a basal facet
(Ba) forming a simple base, mainly subisodiametrical to
broadly heterodiametrical, with 5–7, predominantly 6, pro-
nounced angles in outline, weakly pigmented yellowish
pink, somewhat thick-walled. Basidia 43–61
×
12–15µm,
clavate, hyaline, predominantly 4-spored, with up to 6 µm
long sterigmata, rarely 2-spored. Lamellar edge fertile,
hymenial cystidia absent. Hymenophoral trama regular,
formed by cylindrical to fusiform cells, 55–135 × 6–25µm.
Pileipellis a cutis formed by rather long, narrow, and cylin-
drical cells, 55–190 × 6.0–12.0µm, with some cylindrical
to clavate terminal cells, cells shorter and wider down-
wards, gradually passing into the pileitrama, composed of
Fig. 2 Basidiospore shapes of
Entoloma spp. as seen by SEM
a cuboid with dièdre basal
(KaiR623), b heptahedroid
with simple base (CME6), c
octahedroid with dièdre basal
(KaiR465), d enneahedroid
with dièdre basal (KaiR612),
e decahedroid with simple
base (KaiR691), f hendeca-
hedroid with dièdre basal
(KaiR628), g hendecahedroid
with dièdre basal (KaiR534), h
dodecahedroid with dièdre basal
(CME16), i tetradecahedroid
with dièdre basal (KaiR670),
j, k indetermined shape with
frequent small intercalary
facets and more than two apico-
adaxial facets (KaiR687), l
indetermined shape with differ-
ent base types, including a base
formed by three facets, upper
right (JR64); D: adaxial facet,
AD: apico-adaxial facets, sAD:
single apico-adaxial facet, A:
apical facet, Ba: basal facet, B:
abaxial facet, L: lateral facets;
bars = 5µm
104 Mycological Progress (2022) 21:93–145
1 3
inflated to fusiform cells, up to 22µm wide. Pigment intra-
cellular, brown, mainly in dense clusters of brown granules
in the upper pileipellis. Stipitipellis a cutis composed of
cylindrical cells, 75–300 × 3.0–9.5µm, with intracellular,
brown, mainly granular pigment, like the pigment of the
pileipellis. Caulocystidia absent. Oleiferous hyphae present
in the trama. Clamp connections abundant in all parts of
the basidiocarp.
0.05
WU28035_Entoloma_gerriae_AUT
17283_Entoloma_nidorosum_GG160815_SWE
PA864_Entoloma_nubooccultatum_PAN
S585_Entoloma_cistophilum_isotype_FRA
LN850600_Entoloma_philocistus_paratype_WU18871_PRT
N90_Entoloma_venosum_MD2012_15_DEU
LN850511_Entoloma_politum_289_09_FIN
LN850494_Entoloma_rhodopolium_neotype_213_14_TUR_SWE
MH790432_Entoloma_silvae_frondosae_L_DB6568_holotype_HUN
T628_Entoloma_bryorum_cf_TEB395_18_NOR
S492_Entoloma_olorinum_holotype_FRA
ALV17780_Entoloma_sp
ALV14241_Entoloma_sordidulum_aff_NOR
KaiR692_Entoloma_brunneorugulosum_PAN
KC710062_Entoloma_sordidulum_Co_David_2003
KaiR588_Entoloma_nuboocculta tum_PAN
LN850500_Entoloma_aff_rhodopolium_68_88_68_88_FIN
LN850624_Entoloma_boreale_holotype_106_09_TUR_FIN
MH790420_Entoloma_tiliae_LE254179_holotype_RUS
Entoloma_politum_s_Gulden_NOBAS472617_NOR
19259_Entoloma_sp_EL145_18_SWE
4003828308_MEN2012111_Entoloma_nidorosum_NDL
KaiR687_Entoloma_nubooccultatum_holotype_PAN
NOBAS580018_Entoloma_rhodopolium_OF254444_NOR
KaiR691_Entoloma_brunneorugulosum_holotype_PAN
LN850580_Entoloma_rubrobasis_24101_TURA_FIN
S606_Entoloma_festivum_holotype_NLD
KaiR695_Entoloma_brunneorugulosum_PAN
Entoloma_sordidulum_s_MEN_ALV8406_NOR
S602_Entoloma_lactarioides_holotype_RUS
LN850530_Entoloma_paragaudatum_holotype_383_08_TUR_FIN
Entoloma_cf_venosum_WU_19672
MN809556_Entoloma_abbreviatipes_MICH9005_holotype_USA
S519_Entoloma_transvenosum_holotype_DNK
LN850544_Entoloma_rivulare_holotype_703_12_TUR_FIN
Entoloma_tiliae_NOBAS211316_NOR
NOBAS512218_Entoloma_politum_OF61875_NOR
PA165_Entoloma_nubooccultatum_PAN
NOBAS520518_Entoloma_sp_OF304982_NOR
PA727_Entoloma_nubooccultatum_PAN
C_F5917_Entoloma_pusillulum_holotype_DNK_GL
KaiR583_Entoloma_brunneorugulosum_PAN
Entoloma_rhodopolium_NOBAS460017_NOR
Entoloma_cf_alpicola_NOBAS235616_NOR
LN850508_Entoloma_melenosmum_holotype_JHP_338_C6229_DNK_GL
LN850591_Entoloma_griseorugulosum_isotype_RFS_020921_01_ESP
A376_Entoloma_sp_KBEB83_17_NOR
Entoloma_nidorosum_s_MEN_FR2016708_FRA
PAN532_Entoloma_brunneorugulosum_PAN
PA696_Entoloma_nubooccultatum_PAN
HQ445607_Entoloma_cf_alpicola_23_N1F31_2
Entoloma_rubrobasis_NOBAS453317_NOR
100
69
63
100
92
100
90
70
100
81
100
96
25
100
100
69
83
83
97
94
97
30
99
47
49
61
80
100
100
83
61
92
86
100
Fig. 3 Maximum Likelihood phylogram of species of subg. Entoloma based on ITS, rooted to E. festivum, bootstrap values above or below
branches, bar = estimated changes/nucleotide
105Mycological Progress (2022) 21:93–145
1 3
0.05
LN850540_Entoloma_caccabus_Podoba_G262081_FRA
17266_Entoloma_majaloides_GG160812_SWE
Entoloma_svalbardense_JF304386_MENJG228_NOR_SV
4003828298_MEN2012125_Entoloma_sp_NLD
LN850614_Entoloma_fasciculatum_holotype_TENN29376_USA
MF686497_Entoloma_cf_spadiceum_TENN_070791_USA_TN
LN850536_Entoloma_bisporigerum_holotype_2047_K108895_UK
Entoloma_lividoalbum_GE12_025_FRA
LN850621_Entoloma_brunneipes_holotype_TENN9029_USA
58_Entoloma_myriadophyllum_366VN10
Entoloma_leucocarpum_S568_holotype_NLD
LN850596_Entoloma_subradiatum_lectotype_G262076_FRA
EM6_Entoloma_dulciodorans_LGD_7580_IBUG_MEX
S481_Entoloma_griseopruinatum_holotype_FRA
EM13_Entoloma_myrmecophilum_Quezada_1_IBUG_MEX
LN850570_Entoloma_lupinum_holotype_13_14_TUR_FIN
LC088067_Entoloma_sarcopum_KUB205_JAP
EU807287_uncultured_fungus_USA
4012522693_C158_4001_Entoloma_caccabus_NLD
LN850519_Entoloma_borgenii_KK307_05_FIN
17304_Entoloma_atrosericeum_GG160811_SWE
4012522691_C157_4226_Entoloma_caccabus_NLD
LN850537_Entoloma_alnobetulae_holotype_K53_59_G262083_FRA
LN850517_Entoloma_paludicola_holotype_K178125_UK
KC710116_Entoloma_sinuatum_JVauras8181F_FIN
MF955123_Entoloma_sp_UBC_F_32218_CAN_BC
LN850526_Entoloma_serpens_holotype_410_09_TUR_FIN
LN850556_Entoloma_griseopruinatum_isotype_JLC030924_8_FRA
EM7_Entoloma_dulciodorans_Rodriguez322_IBUG_paratype_MEX
LN850462_Rhodophyllus_myrmecophilus_K398_holotype_FRA
LN850597_Entoloma_sordidulum_Romagn09_1935_syntype_FRA
MH234486_Entoloma_aurorae_borealis_O_F_254651_holotype_NOR
14662_Entoloma_jubatumPAM13_26_SWE
NOBAS531318_Entoloma_alpicola_OF61507_NOR
LN850615_Entoloma_brunneocinereum_holotype_TENN12881_USA
LC088033_Entoloma_subrhodopolium_KUB1_holotype_JAP
Entoloma_gerriae_S550_holotype_NLD
WU39739_Entoloma_noordeloosii
N128_Entoloma_sp_IK6540_USA
LN850444_Entoloma_sericatum_358_08_FIN
Entoloma_saussetiense_LN850594_holotype_FRA
LC088042_Entoloma_pseudorhodopolium_KUB102_holotype_JAP
LN850617_Entoloma_carolinianum_holotype_TENN21855_USA
LN850585_Entoloma_radicipes_holotype_42_14_TUR_FIN
N20_Entoloma_eminens_TEB699_17_NOR
LN850592_Entoloma_palustre_holotype_101_14_TUR_FIN
EM3_E_carolinianum_cf_D_Montanez222_IBUG_MEX
LN850445_Entoloma_sericatum_237_11_FIN
KY574307_uncultured_Entoloma_MEX
EM12_E_myrmecophilum_Santos_Barba5_IBUG_MEX
Entoloma_noordeloosii_758_WU18780_holotype_AUT
KaiR533_Entoloma_pruinosum_holotype_PAN
LN850489_Entoloma_majaloides_holotype_1168_K69959_UK
LN850539_Entoloma_bryorum_117_07_FIN
NOBAS531018_Entoloma_alpicola_OF7195_NOR
LN850524_Entoloma_borgenii_TB79_120_C5933_DNK_GL
CME14_Entoloma_pruinosum_PAN
JR64_Entoloma_mediorobustum_holotype_PAN
LC088049_Entoloma_lacus_KUB110_holotype_JAP
LN850607_Entoloma_atrosericeum_holotype_K69_310_G262084_FRA
S565_Entoloma_roseoalbum_holotype_NLD
JN021019_Entoloma_aff_sinuatum_TRTC156546_CAN
LN850616_Entoloma_lividomurinum_holotype_TENN28208_USA
17298_Entoloma_sp_JO160811_SWE
LN850584_Entoloma_eminens_holotype_417_12_TUR_FIN
Entoloma_eminens_NOBAS97115_NOR
PA728_Entoloma_mediorobustum_PAN
EM15_Entoloma_rhodopolium_Montanez149_150_MEX
LN850610_Entoloma_svalbardense_holotype_GG310_86_O74756_NOR_SV
Entoloma_inusitatum_S594_holotype_DEU
LN850618_Entoloma_nigrobrunneum_holotype_TENN9100_USA
Entoloma_bisporigerum_ALV8408_NOR
N93_Entoloma_platyphylloides_MD2017_10_ITA
Entoloma_myrmecophilum_var_coalescens_S582_holotype_DEU
S580_Entoloma_baeosporum_holotype_FRA
Entoloma_subarcticum_S510_isotype_DNK_GL
A456_Entoloma_rhodopolium_cf_DB03_06_2018_3_HUN
CME12_Entoloma_pruinosum_PAN
Entoloma_bryorum_NOBAS320816_NOR
86
39
27
24
7
31
41
93
100
97
74
100
57
98
81
65
97
8
87
100
100
100
100
8
100
98
94
93
75
100
40
39
47
76
100
49
16
100
25
27
44
100
86
46
89
10
54
21
100
100
100
58
100
100
14
100
6
95
69
Fig. 3 (continued)
106 Mycological Progress (2022) 21:93–145
1 3
Fig. 4 Basidiocarps, and basidi-
ospores as seen by SEM of
Entoloma brunneorugulosum
a (KaiR583) b, c, d (KaiR691,
holotype) bars = 10 µm
Fig. 5 Microscopic structures
of Entoloma brunneorugulosum
a pileipellis with intracellular
pigment (KaiR691, holotype),
bar = 20 μm b basidiospores
(KaiR583), bar = 10 μm
a
b
Fig. 6 Basidiocarp, and
basidiospores as seen by SEM
of Entoloma mediorobustum
(JR64, holotype)
107Mycological Progress (2022) 21:93–145
1 3
Habitat: Basidiocarps solitary to scattered in a group, in
Quercus-dominated montane forest at around 2000m asl. in
Chiriquí, Panama. Probably ectomycorrhizal with Quercus
spp. or Comarostaphylis arbutoides.
Additional specimens examined: PANAMA.
CHIRIQUÍ: near Volcán, Paso Ancho, PNVB, approx.
2000m asl., montane forest with abundant Quercus spp. and
Comarostaphylis arbutoides, 10 July 2016, H. Lotz-Winter,
PAN532 (M); ibid., 17 June 2017, E. Heinemann, KaiR583
(M); ibid., 28 June 2017, K. Reschke and T.A. Hofmann,
KaiR692 (M); ibid., 28 June 2017, K. Reschke and T.A.
Hofmann, KaiR695 (UCH9247, M).
Notes: Entoloma brunneorugulosum is characterised
by small to medium basidiocarps with a rugulose, brown,
umbonate pileus and a somewhat paler brown stipe, as well
as intracellular, granular pigment, and possibly by an associa-
tion to Quercus spp. Phylogenetically, it is a sister species to
Entoloma rivulare Kokkonen with a p-distance of 1.7% in the
ITS. Entoloma rivulare was described from a single collec-
tion in a moist Alnus incana/Populus tremula forest in Fin-
land. Its basidiocarps are like those of E. brunneorugulosum
in having a rather small, brown pileus, with a translucently
striate margin and a farinaceous smell and taste. It differs, how-
ever, by the non-umbonate, darker pileus and smaller basidi-
ospores with lower Q-values. Furthermore, these two species
have different mycorrhizal hosts. Entoloma griseorugulosum
Noordel. & Fernández Sas. is similar by its rugulose, dark
grey-brown pilei, which are not translucently striate, and the
size and shape of the basidiospores (Noordeloos 2004). Our
molecular phylogenetic analysis (Fig.5) suggests that E. gri-
seorugulosum is a later synonym of E. transvenosum Noordel.
Entoloma transvenosum was described with a translucently
striate pileus and larger basidiospores (9.0–13.0 × 7.0–10µm).
Both descriptions note greyish, often transvenose lamellae, a
stipe paler than pileus, intracellular pigment, and occurrence in
rather open forest, often together with Quercus spp. in western
Europe (Noordeloos 1982, 2004). Considering both concepts,
E. brunneorugulosum differs by the warmer, lighter brown col-
our of the pileus, the often concolorous stipe, and the different
habitat. Basidiocarps of E. rugosostriatum Largent & T.W.
Henkel are like those of E. brunneorugulosum by rugulose,
brown pilei with a striate margin; however, the basidiospores
of E. rugosostriatum are isodiametrical, weakly angled and
smaller (6.1–7.4 × 6.1–7.3µm) (Largent etal. 2008b). None of
the species described by Hesler (1967), Horak (1978, 1982),
Largent (1994), and Montañez etal. (2016) combine a dark
rugulose, non-hygrophanous pileus, and relatively paler stipe,
and an association with Quercus spp.
Entoloma mediorobustum Reschke, J. Rodríguez-
Cedeño & Noordel., sp. nov. Figs.6 and 7
Mycobank number MB840629.
Typification: PANAMA. CHIRIQUÍ: near Volcán,
Paso Ancho, PNVB, trail towards summit of volcano, N
8°48ʹ57.1'ʺ W 82°24ʹ35.2'ʺ, 2,040m asl., montane forest
with abundant Quercus spp. and Comarostaphylis arbuto-
ides, 1 August 2015, J. Rodríguez, S. Cáceres, K. Barrera, R.
Valdéz, JR64, holotype (UCH11749), isotype (M).
Etymology: medius (Latin)
=
medium, robustus
(Latin) = robust; refers to the shape of the basidiocarps.
Description: Basidiocarps tricholomatoid. Pileus 4.2–4.8cm
diameter, convex, with initially incurved, later straight and undu-
lating margin, yellowish brown (5E4, 5E5, 5D5, 5D6), pileal
surface smooth, sometimes partly covered by a whitish pruina,
not translucently striate, hygrophanous. Lamellae emarginate,
medium-spaced to rather distant, greyish white when young,
becoming dirty pink, with somewhat uneven to eroded, con-
colorous edges. Stipe 10.0 × 0.8–1.0cm, cylindrical with radi-
cating base, stuffed, whitish, fibrillose. Basal mycelium white,
somewhat cottony. Odour and taste not tested.
Basidiospores 7.5–8.7–10.0 × 7.0–7.9–8.5 µm,
Q = 1.00–1.11–1.20 (n = 50 spores of 2 specimens), spatial
shape not determined, at least with 12 facets, base often built
by 3 facets, isodiametrical to subisodiametriacal, with 5–7
rather blunt angles in outline, weakly pigmented yellowish
pink, somewhat thick-walled. Basidia 30–43 × 9.0–11.0µm,
clavate to subclavate, hyaline, 4-spored, sterigmata up to
4.0µm long. Lamellar edge fertile, hymenial cystidia
Fig. 7 Microscopic structures
of Entoloma mediorobustum
(JR64, holotype): a pileipel-
lis with incrusting pigment
indicated on the right hand side,
bar = 20μm; b basidiospores,
bar = 10μm
a
b
108 Mycological Progress (2022) 21:93–145
1 3
absent. Hymenophoral trama regular, formed by sub-
cylindrical to fusiform cells, (40) 50–200 × 6.0–30µm.
Pileipellis a cutis composed of narrow, cylindrical cells,
70–250 × 3.5–9.0µm, pigment incrusting, brownish in KOH.
Stipitipellis a cutis composed of rather hyaline, cylindrical
cells, 55–150 × 3.5–8.0 (9.5) µm. Caulocystidia absent.
Clamp connections present in all parts of the basidiocarps.
Habitat: Basidiocarps solitary on soil in a montane forest
with abundant Quercus spp. and Comarostaphylis arbutoides,
between 1950 and 2050m asl. in Chiriquí, Panama. Prob-
ably ectomycorrhizal with Quercus spp. or Comarostaphylis
arbutoides.
Additional specimen examined: PANAMA. CHIRIQUÍ:
near Volcán, Paso Ancho, PNVB, N 8°48ʹ55.4'ʺ W 82°34ʹ47.1'ʺ,
1,970m asl., montane forest with abundant Quercus spp. and
Comarostaphylis arbutoides, 9 November 2015, T.A. Hofmann,
J. Rodríguez, S. Cáceres, R. Villarreal, PA728 (M).
Notes: Entoloma mediorobustum is distinctive by the non-
hygrophanous pileus with a distinct pruina, which is rather
unusual in this subgenus. It clusters with two collections from
Mexico, labelled as E. myrmecophilum and E. cf. carolinianum,
respectively (Fig.5), that differ by glabrous, relatively dark brown
pilei (Montañez etal. 2016). These species have in common
their possible association with Quercus spp. and the presence
Fig. 8 Basidiocarps and basidi-
ospores as seen by SEM of
Entoloma nubooccultatum:
a, e (KaiR687, holotype) bar
= 10 µm; b (PA165) picture
by S. Cáceres c (KaiR588); d
(PA727)
Fig. 9 Microscopic structures
of Entoloma nubooccultatum
(KaiR687, holotype): a pileipel-
lis, intracellular pigment not
indicated, bar = 20μm; b basidi-
ospores, bar = 10μm
a
b
109Mycological Progress (2022) 21:93–145
1 3
of incrusting pigments in the pileipellis. Entoloma pruinosum is
somewhat similar, but its basidiocarps are slightly darker, more
robust, and have a more differentiated stipe surface. The two spe-
cies share the relatively small, isodiametrical to subisodiametrical
spores. Phylogenetically, however, they are quite distant (Fig.5).
Entoloma nubooccultatum Reschke & Noordel., sp. nov.,
Figs.8 and 9
Mycobank number: MB840632.
Typification: PANAMA. CHIRIQUÍ: near Volcán, Paso
Ancho, PNVB, approximately 2000m asl., montane forest
with abundant Quercus spp. and Comarostaphylis arbuto-
ides, 28 June 2017, K. Reschke and T.A. Hofmann KaiR687,
holotype (UCH9243), isotype (M).
Etymology: nubes (Latin)
=
cloud, occultatum
(Latin)
=
hidden; refers to the hidden life of this species in
cloud forests.
Description: Basidiocarps collybioid. Pileus 1.5–6.5cm
diameter, convex umbilicate to depressed, with deflexed and
uneven margin which is slightly incurved in young basidiocarps
and straight in older basidiocarps, greyish yellow (4B4, 4B5,
4C5, 4C6), brownish orange (5C4, 5C5, 5C6), yellowish brown
(5D5, 5D6, 5D7, 5E4, 5E5) to olive brown (4D5, 4D6), pileal
surface glabrous, with a waxy appearance when wet, translu-
cently striate at the margin about 1/3 to 2/3 the centre, hygropha-
nous, fading to yellowish grey (3B2). Lamellae broadly adnate
to somewhat decurrent, segmentiform, close, whitish in young
specimens, pinkish at maturity, with somewhat undulating and
concolorous edges. Stipe 4.5–10 × 0.3–0.8cm, cylindrical, often
bent, stuffed to hollow, white, somewhat grooved, finely innately
fibrillose, base often covered by adhering soil and litter. Basal
mycelium white, with many small rhizomorphs. Odour indis-
tinct or slightly sweetish, fruity, taste not tested.
Basidiospores 7.0–7.8–8.5 × 6.0–6.9–7.5 µm,
Q = 1.05–1.13–1.25 (n =
146 spores of 6 specimens), spatial
shape not determined, at least with 11–13 facets, mainly with a
dièdre basal, sometimes incomplete, sometimes with a simple
base, isodiametrical to broadly heterodiametrical, predominantly
subisodiametrical, with 6–7 weak to pronounced, somewhat
rounded angles in outline, weakly pigmented yellowish pink,
somewhat thick-walled. Basidia (30) 35–43 (48) × 8.5–12.5µm,
subclavate, often almosturniform (formed like an urn), hyaline,
4-spored, with up to 6.0µm long sterigmata, few 2-spored.
Lamellar edge fertile, hymenial cystidia absent. Hymenophoral
trama regular, formed by cylindrical, somewhat fusiform to inflated
cells, 45–150 × 6–20 (28) µm. Pileipellis a cutis of cylindrical cells,
35–185 × (4.0) 6.0–12.5µm, with intracellular pigment. Stipitipellis
a cutis composed of short, cylindrical cells, 30–70 × 3.0–6.0µm, not
pigmented. Caulocystidia absent. Oleiferous hyphae present in the
trama. Clamp connections abundant in all parts of the basidiocarp.
Habitat: Basidiocarps solitary or in small groups
in a montane Quercus-dominated forest with abundant
Comarostaphylis arbutoides between 1900 and 2200m
asl. in Chiriquí, Panama. Probably ectomycorrhizal with
Quercus spp. or Comarostaphylis arbutoides.
Additional specimens examined: PANAMA. CHIRIQUÍ:
near Volcán, Paso Ancho, PNVB, approximately 2000m asl.,
montane forest with abundant Quercus spp. and Comarostaphy-
lis arbutoides, 17 June 2017, K. Reschke, KaiR588 (M); ibid.,
N 8°48ʹ55.3'ʺ W82°34ʹ43.5, 2000m asl., montane forest
with abundant Quercus spp. and Comarostaphylis arbutoides,
28 September 2014, S. Cáceres, K. Barrera, T.A. Hofmann,
R. Valdés, PA165 (M); ibid., N 8°48ʹ53.4ʺ W82°34ʹ48.0ʺ,
1,960m asl., montane forest with abundant Quercus spp. and
Comarostaphylis arbutoides, 9 November 2015, S. Cáceres,
R. Valdés, J. Rodríguez, T.A. Hofmann, PA696 (M); ibid.,
N 8°48ʹ55.4'ʺ W 82°34ʹ47.1ʺ, 1970m asl., montane forest
with abundant Quercus spp. and Comarostaphylis arbutoides,
9 November 2015, T.A. Hofmann, J. Rodríguez, S. Cáceres,
R. Valdés, PA727 (M); ibid., N 8°48ʹ52.9ʺ W 82°34ʹ47.7ʺ,
1920m asl., montane forest with abundant Quercus spp. and
Comarostaphylis arbutoides, 11 September 2016, T.A. Hof-
mann and M. Cuevas, PA864 (M).
Notes: Entoloma nubooccultatum is characterised by
basidiocarps with collybioid habit, a brown, translucently stri-
ate, hygrophanous pileus, a white stipe, intracellular pigment
in the pileipellis and rather small basidiospores. Phylogeneti-
cally, it forms part of a clade including E. rhodopolium (Fr.) P.
Kumm. and several similar, closely related species, which have
not yet been completely resolved (Kokkonen 2015; Brandrud
etal. 2018). Its p-distance to these species ranges from 2.8 to
3.2% in the ITS. Habitually, it is not unlike E. melenosmum
Noordel. However, E. nuboocculatatum has basidiospores that
are on average smaller than those of most species in the E.
rhodopolium clade, and its distribution is probably restricted
to tropical montane Quercus forests, while E. rhodopolium and
related species are mainly distributed in northern temperate and
boreal deciduous forests, predominantly associated with species
of Betulaceae, sometimes Fagus sylvatica L., but rarely with
Quercus spp. (Noordeloos 2004).
Entoloma pruinosum Reschke, Manz & Noordel., sp.
nov., Figs.10 and 11
Mycobank number: MB840622.
Typification: PANAMA. CHIRIQUÍ: near Bajo Boquete,
Jaramillo Arriba, El Musgo, forest trail towards rock look-
out, N 08°47ʹ32.3ʺ W 82°24ʹ50.3ʺ, 1650m asl., mon-
tane forest dominated by Quercus spp., 14 June 2017, K.
Reschke, KaiR533, holotype (UCH009184), isotype (M).
Etymology: pruinosus (Latin) = pruinose; refers to the
pruinose pileal surface.
Description: Basidiocarps tricholomatoid. Pileus
3.0–7.0cm diameter, convex when young, becoming broadly
convex to almost plane in old basidiocarps, with or without
110 Mycological Progress (2022) 21:93–145
1 3
an obtuse umbo, margin initially deflexed and still slightly
deflexed in old basidiocarps, greyish yellow, pale olive brown,
greyish brown, brownish grey to grey (4B4, 4D5, 4D6, 5B1,
5C2, 5D2, 5D3), uniformly coloured, pileal surface glabrous,
with a waxy appearance, covered by a whitish, slightly fibril-
lose pruina, not translucently striate, not hygrophanous.
Lamellae sinuate, segmentiform to ventricose, crowded, ini-
tially almost white to pale greyish, later pink, with slightly
blunt serrate, concolorous edges and moderately to strongly
transvenose sides. Stipe 8.0–12.0 × 0.8–1.0cm, cylindrical to
somewhat tapering towards base, solid, greyish yellow (4B4,
4B5, 4C3, 4C4, 4C5, 4C6, 4C7) under a whitish fibrillose to
Fig. 10 Basidiocarps and
basidiospores as seen by SEM
of Entoloma pruinosum: a
(CME14); b (PA167); c, d
(KaiR533, holotype)bar = 10
µm
Fig. 11 Microscopic struc-
tures of Entoloma pruinosum
(KaiR533, holotype): a pileipel-
lis with incrusting pigment
indicated at the right hand side,
bar = 20µm; b stipitipellis with
caulocystidia, bar = 20µm; c
basidiospores, bar = 10µm
a
c
b
111Mycological Progress (2022) 21:93–145
1 3
fibrillose-flocculose layer, staining brownish yellow (5C7,
5C8) after damage. Basal mycelium cottony, white. Odour
not distinctive to weakly farinaceous when undamaged, fari-
naceous when cut, taste farinaceous.
Basidiospores 7.0–7.9–9.0 × 6.0–7.0–8.0 µm,
Q = 1.05–1.13–1.25 (n = 94 spores of 4 specimens), decahe-
droid, composed of an adaxial facet (D), a pair of apico-adaxial
facets (AD), an apical facet (A), an abaxial facet (B), two pairs
of lateral facets (L1–L2), and a basal facet (Ba) forming a sim-
ple base, isodiametrical to subisodiametrical, with 5–6 pro-
nounced angles in outline, weakly pigmented yellowish pink,
somewhat thick-walled. Basidia 44–60 × 7.5–11.0µm, clavate,
hyaline, 4-spored, sterigmata up to 4.0µm long. Lamellar edge
fertile, hymenial cystidia absent. Hymenophoral trama regular,
composed of somewhat fusiform to inflated cells in the centre,
50–120 × 6.0–14.0µm, longer, up to 240µm and narrower,
narrowest 4.0µm, towards the subhymenium. Pileipellis a cutis
formed by cylindrical cells, 65–130 × 3.0–6.0µm, with finely
incrusting pigment, without distinct subpellis. Pileitrama com-
posed of rather short and wide, subcylindrical to inflated cells,
40–65 × 9.0–11.5µm. Stipitipellis a cutis composed of cylin-
drical cells, 40–200 × 3.5–7.0µm, with incrusting pigment
0.05
KC257436_Entoloma_pulchellum_HMLD1300_CN
KF679348_Nolanea_albertinae_FK0935_holotype_BR
JX454803_Entoloma_olivaceohebes_WU20798_IT
CME6_Entoloma_paraconfere ndum_holotype_PAN
KF698730_Entoloma_luteofuscum_K_M_188309_holotype_IN
KC898450_Entoloma_cetratum_LE235480_RU
CME9_Entoloma_transitionisporum_PAN
KaiR628_Entoloma_flavoconicum_holotype_PAN
PDD87270_Entoloma_readiae_NZL
KaiR508_Entoloma_cremeostriatum_PAN
KF679350_Nolanea_albertinae_FK1731_BR
KF738923_Nolanea_pallidosalmonea_FK0891_BR
KaiR630_Entoloma_belouvense_var_albertinae_PAN
KR673495_Entoloma_pulchellum_KA12_1242_KR
KaiR609_Entoloma_flavoconicum_PAN
KaiR693_Entoloma_sp_PAN
KM262035_Entoloma_piceinum_LE254131_holotype_RU
KF679354_Nolanea_atropapillata_FK0898_holotype_BR
CME5_Entoloma_belouvense_var_albertinae_PAN
KaiR1342_Entoloma_conferendum_AUT
WU32902_Entoloma_subelegans_holotype_MUS
KaiR503_Entoloma_transitionisporum_holotype_PAN
CME7_Entoloma_paraconferendum_PAN
KaiR626_Entoloma_sp_PAN
KaiR988_Entoloma_infula_DEU
PDD71294_Entoloma_readiae_NZL
KF679349_Nolanea_albertinae_FK0912_BR
CME10_Entoloma_cremeostriatum_PAN
KaiR978_Entoloma_conferendum_DEU
KF679351_Nolanea_albertinae_FK1732_BR
KaiR1358_Entoloma_conferendum_DEU
JX454804_Entoloma_olivaceohebes_WU17841_holotypus_IT
WU27132_Entoloma_belouvense_holotype_FRA_RE
MH211959_Nolanea_sp_FLAS_F_61537_US
MF476911_Entoloma_vernum_LE_312418_RU
KF679353_Nolanea_parvispora_FK1140_BR
KF679352_Nolanea_tricholomatoidea_FK1049_holotype_BR
KaiR1282_Entoloma_clandestinum_SWE
KaiR989_Entoloma_infula_DEU
KaiR613_Entoloma_cremeostriatum_holotype_PAN
KaiR1267_Entoloma_sericeum_SWE
KC898421_Entoloma_euchroum_KR_M_0032474_neotype_DE
29
61
61
68
98
100
100
47
87
84
87
91
100
90
98
100
97
100
61
99
82
100
100
92
44
85
97
74
47
68
100
86
78
99
99
61
50
Fig. 12 Maximum Likelihood phylogram of species of subg. Nolanea based on ITS, rooted to E. subelegans and E. euchroum, bootstrap values
above or below branches, bar = estimated changes/nucleotide
112 Mycological Progress (2022) 21:93–145
1 3
at some hyphae. Caulocystidia clavate to subcylindrical,
35–65 × 6.5–10.0µm. Oleiferous hyphae present in the trama.
Clamp connections abundant in all parts of the basidiocarp.
Habitat: Basidiocarps solitary or gregarious, in Quercus-
dominated montane forests between 1650 and 1900m asl. in
Chiriquí, Panama. Probably ectomycorrhizal with Quercus
spp.
Additional specimens examined: PANAMA.
CHIRIQUÍ: near Volcán, Paso Ancho, PNVB, N 08°48ʹ55ʺ
W82°34ʹ46.8ʺ, 1,900m asl., montane forest with abundant
Quercus spp. and Comarostaphylis arbutoides, 28 Septem-
ber 2014, S. Cáceres, K. Barrera, T.A. Hofmann, R. Villar-
real, PA167 (M); near Bajo Boquete, Jaramillo Arriba, El
Musgo, N 08°47ʹ26.9ʺ W 82°24ʹ34.4ʺ, 1,670m asl., mon-
tane forest dominated by Quercus spp., 24 June 2018, T.A.
Hofmann, CME12 (M); Bajo Boquete, Finca Lérida, forest
trail, N 08°49ʹ10.3ʺ W 82°29ʹ09.2ʺ, 1820m asl., montane
forest with Quercus seemannii and other Quercus spp., 28
June 2018, C. Manz and F. Hampe, CME14 (M).
Notes: Basidiocarps of E. pruinosum are characterised by
a tricholomatoid habit, a brown surface of the pileus with
a distinct pruina and relatively small basidiospores. This
combination of characters is rather unique. No other known
species in subg. Entoloma has such a strongly developed
pruina on the pileus. The small, isodiametrical spores are
more or less like those found in the /prunuloides clade, and
the European E. prunuloides may have a slight, but not so
strong pruina on the pileus. Entoloma prunuloides, however,
like some other similar species, is phylogenetically very dis-
tant from subg. Entoloma (Morgado etal. 2013). Entoloma
pruinosum belongs to a clade of species known from Central
and North America. In the ITS phylogeny (Fig.5), it is close
to E. brunneocinereum Hesler, with a p-distance of 2.7%
in the ITS1. This species has, however, somewhat smaller
basidiospores (7.0–8.0
×
6.0–7.0µm) and intracellular pig-
ment (Noordeloos 1988). Based on type studies, Kokkonen
(2015) demonstrated that E. lividomurinum Hesler is con-
specific with E. brunneocinereum. Entoloma griseopruina-
tum Cheype & Noordel., a European species with a pruinose
pileus, has larger basidiospores (9.0–11.0 × 7.0–8.0µm),
more greyish colours in stipe and pileus, intracellular
pigment in the pileipellis, and is phylogenetically distant
(Fig.5).
Subgenus Nolanea (Fr.) Noordel (Fig.12)
Traditionally, species of the subg. Nolanea are character-
ised by basidiocarps with a mycenoid habit, a rather smooth
pileus surface, and relatively long cells in the hymenopho-
ral trama (Noordeloos 1980). A preliminary study on this
subgenus in Europe was published by Vila etal. (2013).
Karstedt etal. (2020) described five new species from Bra-
zil, using Nolanea at generic rank. Their molecular phylog-
eny based on three loci supports Nolanea as a monophyletic
clade. The results of a large, worldwide study on this sub-
genus are currently being prepared for publication (Reschke
etal., in prep.).
Entoloma belouvense Noordel. & Hauskn., Fungal
Divers. 27: 112 (2007) var. belouvense.
Holotype WU27132: Several basidiocarps in good
condition. Basidiospores heterodiametrical, 10.5–11
.3–12.0 × 6.5–7.1–7.5µm, Q = 1.45–1.60–1.85 (n = 26).
Basidia 4-spored, rather broadly clavate. Hymenopho-
ral trama with abundant oily content. Stipe surface
composed of long and narrow cells. Clamp connections
absent.
Specimen examined: LA RÉUNION, Saint-Benoît, Forêt
de Belouve, on decaying, mossy wood of broadleaf tree in
tropical highland rainforest, 28 March 2005, A. Hausknecht,
holotype (WU 27,132).
Notes: Entoloma belouvense is a member of the clade
around E. conferendum (Britzelm.) Noordel. In contrast to
the original description (Noordeloos and Hausknecht 2007),
no clamp connections were observed at the base of basidia.
Fig. 13 Basidiocarp of Entoloma belouvense var. albertinae
(KaiR630)
Fig. 14 Basidiospores of Ento-
loma belouvense var. albertinae
(KaiR630), bar = 10 µm
113Mycological Progress (2022) 21:93–145
1 3
Entoloma belouvense Noordel. & Hauskn. var. alberti-
nae (Karstedt & Capelari) Reschke & Noordel. comb. & stat.
nov., Figs.13 and 14
Basionym: Nolanea albertinae Karstedt & Capelari in
Karstedt etal., Mycotaxon 135: 596 (2020).
Mycobank: MB840620.
Basidiocarps mycenoid. Pileus 1.5–2.0cm diameter,
broadly conical to expanded with pronounced papilla and
straight to deflexed, somewhat uneven margin, greyish
yellow–golden, brown to dark brown (4C6, 6E5, 6E6, 6F5,
6F6) with dark brown (6F6, 7F5) centre, pileal surface
glabrous, somewhat innately fibrillose, translucently striate
almost to the centre, hygrophanous and then with sericeous
appearance. Lamellae adnexed to free, ventricose, medium-
spaced to rather distant, greyish white when young, becoming
brownish pink, with smooth and concolorous edges. Stipe
Fig. 15 Basidiocarps and
basidiospores as seen by SEM
of Entoloma cremeostriatum:
a (KaiR508); b (CME10)
bar = 2cm; c, d (KaiR613, holo-
type) bar = 10µm
Fig. 16 Microscopic structures
of Entoloma cremeostriatum
(KaiR613, holotype): a pileipel-
lis with incrusting pigment
indicated on the right hand side,
bar = 20μm; b basidiospores,
bar = 10μm
a
b
2.0–3.5 × 0.20–0.25cm, cylindrical, getting slightly broader
towards base, hollow, greyish yellow, greyish yellow–golden
to yellowish brown (4C5, 4C6, 5D5), polished to somewhat
innately fibrillose towards base, with silvery white fibrils
when contorted. Basal mycelium white, with many short and
fine rhizomorphs. Odour indistinct, taste not tested.
Basidiospores 9.0–9.9–11.0 × 6.0–7.1–7.5 µm,
Q = 1.25–1.40–1.60 (n = 53 spores of 2 specimens), het-
erodiametrical, with 5–6 pronounced angles in outline,
weakly pigmented yellowish pink, somewhat thick-
walled. Basidia 26–34 × 9.5–11.0µm, subclavate, hyaline,
4-spored, sterigmata up to 5.0µm long. Lamellar edge het-
erogeneous, cheilocystidia scattered, solitary or in small
groups between basidia, rather infrequent, approximately
25–50 × 5.0–8.5µm, septate tibiiform to cylindrical capitate.
Pileipellis a cutis of relatively long and narrow, 4.0 to 8.0
114 Mycological Progress (2022) 21:93–145
1 3
µm wide elements, pigment intracellular, brownish in KOH.
Clamp connections absent from all parts of the basidiocarp.
Habitat: Basidiocarps found solitary or scattered in a
small group in montane forests between 1600 and 2100m
asl. in Chiriquí, Panama.
Specimens examined: PANAMA. CHIRIQUÍ: near
Bajo Boquete, PNVB, Sendero Culebra, N 08°50ʹ42.6ʺ W
82°28ʹ52.5ʺ, 1,670m asl., montane forest dominated by
Quercus spp., 22 June 2017, K. Reschke, J. Rodríguez, B.
Wergen, KaiR630 (UCH11745, M); near Cerro Punta, Los
Quetzales Lodge, N 08°52ʹ30.4ʺ W 82°33ʹ14.0ʺ, 2030m
asl., montane tropical forest, 18 June 2018, C. Manz and F.
Hampe, CME5 (M).
Notes: Nolanea albertinae was described from Brazil by
Karstedt etal. (2020). Before, it had been treated as E. bel-
ouvense by Karstedt etal. (2019). In the phylogeny (Fig.14),
the ITS sequence of the holotype of E. belouvense var. bel-
ouvense is nested among the sequences of specimens of E.
belouvense var. albertinae. However, the differences in spore
size, resulting in higher Q-values in E. belouvense var. bel-
ouvense, are confirmed here. Apart from this, no differing
characters were observed. Nolanea albertinae is treated here
as a variety of E. belouvense because only one specimen of
the type variety is available and ITS sequences may not be
suitable to resolve such close relationships. More specimens
from the type region, La Réunion, are needed to evaluate if
this treatment is justified. Similar to the type variety, no clamp
connections were observed in the specimens from Panama.
Entoloma cremeostriatum Reschke, Manz & Noordel.,
sp. nov., Figs.15 and 16
Mycobank number: MB840936.
Typification: PANAMA. CHIRIQUÍ: near Cerro Punta,
Entre Ríos, Montaña Azul, PILA, approx. 2300m asl., mon-
tane forest dominated by Quercus spp., 21 June 2017, J.
Rodríguez, KaiR613, holotype (M).
Etymology: crème (French) = cream (here: colour), stria-
tus (Latin) = striate; refers to the morphology of the pileus.
Description: Basidiocarps collybioid to omphalinoid.
Pileus 2.5–3.9cm diameter, convex umbilicate, margin
deflexed to straight, slightly exceeding the lamellae, uni-
formly yellowish brown (5E7) in immature basidiocarps,
soon paler, light yellow (4A4), greyish yellow (4B4), grey-
ish orange (5B3, 5B4) to brownish orange (5C6), often
darker, greyish yellow (4B6) to yellowish brown (5F7) in
the centre, whitish at the very margin, pileal surface gla-
brous, with a waxy appearance when wet, innately fibril-
lose at the centre, with some fine, protruding fibrils all over,
translucently striate almost to the centre, hygrophanous
reaction not observed. Lamellae emarginate, segmenti-
form, medium-spaced to rather distant, greyish to cream
when young, becoming pinkish, with slightly undulating and
concolorous edges. Stipe 3.0–3.5 × 0.30–0.35cm, cylindri-
cal, hollow, similarly coloured as the pileus, greyish yellow
(3B4, 4B5, 4C5), brownish orange (5C5, 5C6) to yellowish
brown (5D6), innately fibrillose, partly overlaid by whitish
fibrils, but not silvery fibrillose, slightly whitish fibrillose-
flocculose at the tip. Basal mycelium white, with some short
and fine rhizomorphs. Odour slightly nitrous or somewhat
sweetish, taste not tested.
Basidiospores 6.5–7.6–8.5 × 5.5–6.2–6.5 µm,
Q = 1.10–1.23–1.40 (n = 83 spores of 3 specimens), hendec-
ahedroid, composed of an adaxial facet (D), a pair of apico-
adaxial facets (AD), an apical facet (A), an abaxial facet (B),
and three pairs of lateral facets (L1–L3), of which the basal
pair (L3) forms a dièdre basal, subisodiametrical to broadly
heterodiametrical, with 5–6 pronounced angles in outline,
weakly pigmented yellowish pink, thick-walled. Basidia
21–29 × 8.5–10.0µm, clavate, hyaline, 4-spored, sterig-
mata up to 4.0µm long, with clamp connections. Lamellar
edge fertile, hymenial cystidia absent. Hymenophoral trama
regular, formed by wide, somewhat fusiform cells in the
mediostratum, and narrower, cylindrical cells towards the
subhymenium, 100–360 × 3.5–25µm, with slightly incrust-
ing pigment at some hyphae. Pileipellis a cutis of narrow,
cylindrical cells, 60–200 × 3.0–6.5 (10.0) µm, pigment finely
incrusting, without a subpellis, gradually passing into pileit-
rama, composed of wider cells. Stipitipellis a cutis formed
by long, cylindrical cells, 55–220 × 3.0–8.0µm, with finely
incrusting pigment. Caulocystidia absent. Oleiferous hyphae
present in the trama. Clamp connections abundant in the
hymenium, rarely present elsewhere.
Habitat: Basidiocarps solitarily to gregarious on soil,
between mosses and litter, in Alnus acuminata-dominated
montane forest and Quercus-dominated montane forest
between 2300 and 2400m asl. in Chiriquí, Panama.
Additional specimens examined: PANAMA.
CHIRIQUÍ: Cerro Punta, Las Nubes, PILA Sendero Cas-
cada, N 08°53ʹ51.6ʺ W 82°37ʹ10.4ʺ, 2380m asl., mon-
tane forest dominated by Alnus acuminata, 11 June 2017,
K. Reschke and J. Rodríguez, KaiR508 (M); ibid., N
08°53ʹ37.5ʺ W 82°36ʹ54.4ʺ, 2,030m asl., montane forest
dominated by Alnus acuminata, 22 June 2018, C. Manz and
T.A. Hofmann, CME10 (M).
Notes: Entoloma cremeostriatum is characterised by
a convex-umbilicate, pale, striate pileus, small, hetero-
diametrical basidiospores, and clamp connections at the
base of basidia. With a p-distance of 5.5% between the
ITS sequences, it is relatively close to E. readiae G. Stev.,
described from New Zealand (Stevenson 1962). This species
has basidiocarps which are like those of E. cremeostriatum
in shape. Moreover, the two species share finely incrusting
pigments in the hyphae of the pileipellis and clamp con-
nections at the base of the basidia. The basidiospores of E.
115Mycological Progress (2022) 21:93–145
1 3
readiae are larger than those of E. cremeostriatum according
to the original description (9–10 × 7–8µm). Our measure-
ments of two authentic specimens (Table1) of E. readiae,
however, resulted in somewhat smaller basidiospore sizes,
7.5–8.4–9.0 × 6.5–7.4–8.0µm, Q = 1.00–1.14–1.25 (n = 46
of 2 specimens). However, these are still somewhat larger
and have smaller Q-values than those of E. cremeostria-
tum. In addition, basidiocarps of E. readiae are brown and
have a strongly farinaceous odour, while those of E. cre-
meostriatum are generally paler and have a slightly nitrous
odour. Two sequences annotated as E. pulchellum (Hongo)
Hongo are relatively close to those of E. cremeostriatum
with p-distances of 5.2% between the ITS sequences. Ento-
loma pulchellum is described as having a convex-umbilicate,
yellow to flesh-coloured pileus with brown tones, often a
minutely squamulose pileus surface, especially in the centre,
a striate pileus margin, heterodiametrical basidiospores of
10.0–12.5 × 7.0–9.0µm, clavate to capitate cheilocystidia
of 25–37 × 13.0–14.5µm, and no clamp connections at the
base of basidia (Hongo 1957). These characters, especially
the squamulose pileus surface together with the absence of
clamp connections and presence of distinct cheilocystidia,
indicate a species of subg. Cyanula, probably close to E.
formosum (Fr.) Noordel. Thus, the specimens correspond-
ing to the ITS sequences (KR673495, KC257436) annotated
as E. pulchellum in GenBank apparently are misidentified.
Entoloma avoconicum Reschke & Noordel., sp. nov.,
Figs.17 and 18
Mycobank number: MB840631.
Fig. 17 Basidiocarps and
basidiospores as seen by SEM
of Entoloma flavoconicum:
a (KaiR609); b (PA891);
c, d (KaiR628, holotype),
bar = 10μm
Fig. 18 Microscopic structures
of Entoloma flavoconicum
(KaiR628, holotype): a pileipel-
lis with incrusting pigment
indicated on the right hand side,
bar = 20μm; b basidiospores,
bar = 10μm
a
b
116 Mycological Progress (2022) 21:93–145
1 3
Typification: PANAMA. CHIRIQUÍ: near Cerro
Punta, Entre Ríos, Montaña Azul, PILA, N 08°53ʹ41ʺ W
82°35ʹ01.5ʺ, 2,340m asl., montane forest dominated by
Quercus spp., 21 June 2017, K. Reschke, KaiR628, holotype
(UCH9226), isotype (M).
Etymology: flavus (Latin) = yellow, konikos
(Greek) = conical; refers to the colour and shape of the
pileus.
Description: Basidiocarps mycenoid. Pileus 0.7–2.2cm
diameter, conical when young, remaining conical or becom-
ing convex with a pronounced, acute umbo, either dark
brown (6F7) to yellowish brown (5F6) or light yellow (4A5)
to orange yellow (4A6) in the centre, towards the margin at
first light brown (6D6, 6D7) to yellowish brown (5D7, 5D8,
5E8), then greyish yellow (4B6, 4C6, 4C7) to orange yel-
low (4B7) and eventually yellow (3A7) at the very margin,
pileal surface glabrous, but overlaid by an ephemeral, fine,
white pruina in young basidiocarps, translucently striate
from the margin almost to the centre, hygrophanous reac-
tion not observed. Pileal margin initially slightly incurved,
later straight, slightly overlapping the lamellae. Lamellae
emarginate to adnexed, ventricose, distant, relatively thick
and broad, initially greyish, sometimes with yellow tinges,
later greyish pink, with somewhat transvenose sides and
smooth, concolorous edges. Stipe 1.0–3.5 × 0.14–0.25cm,
cylindrical to slightly clavate, hollow, brownish yellow
(5C7, 5C8) sometimes darker, yellowish brown (5D7, 5D8)
towards base, glabrous to innately fibrillose, finely whitish
pruinose all over, especially towards the apex of the stipe.
Basal mycelium white, often cottony. Odour indistinct, taste
not tested.
Basidiospores 8.5–9.4–10.5 × 6.0–6.4–7.0 µm,
Q = 1.30–1.46–1.65 (n = 70 spores of 3 specimens), hendec-
ahedroid, composed of an adaxial facet (D), a pair of apico-
adaxial facets (AD), an apical facet (A), an abaxial facet (B),
and three pairs of lateral facets (L1–L3), of which the basal
pair (L3) forms a dièdre basal, heterodiametrical, with 5–6,
predominantly 6, pronounced to nodulose angles in outline,
weakly pigmented yellowish pink, somewhat thick-walled.
Basidia 27–34 × 11.0–13.0µm, clavate, hyaline, predomi-
nantly 4-spored, sterigmata up to 6.0µm long. Lamellar edge
fertile, hymenial cystidia absent. Hymenophoral trama regu-
lar, formed by narrow, cylindrical and centrically broad fusi-
form cells, 125–520 × 3.5–27µm, some hyphae with incrust-
ing pigment, yellowish in KOH. Pileipellis a cutis formed by
narrow, cylindrical cells, 40–260 × 3.5–9.5µm, with incrust-
ing and in addition scattered intracellular pigment, cells
wider, somewhat inflated, towards pileitrama. Stipitipellis
a cutis composed of cylindrical cells, 85–400 × 3.0–7.5µm,
with some incrusting pigment. Caulocystidia absent. Oleifer-
ous hyphae present in the trama. Clamp connections absent
from all parts of the basidiocarp.
Fig. 19 Basidiocarps and
basidiospores as seen by SEM
of Entoloma paraconferendum
(CME6, holotpye), bar = 10µm
Fig. 20 Microscopic structures
of Entoloma paraconferendum
(CME6, holotype): a pileipel-
lis with subpellis composed of
inflated cells, intracellular pig-
ment not indicated, bar = 20μm;
b basidiospores, bar = 10μm
ab
117Mycological Progress (2022) 21:93–145
1 3
Habitat: Basidiocarps scattered in a group, in Quercus-
dominated montane forests from 2000 to 2340m asl. in
Chiriquí, Panama.
Additional specimens examined: PANAMA.
CHIRIQUÍ: near Cerro Punta, Entre Ríos, Montaña Azul,
PILA, N 08°53ʹ35.1ʺ W 82°34ʹ53.2ʺ, 2260m asl., mon-
tane forest dominated by Quercus spp., 21 June 2017, K.
Reschke, T.A. Hofmann, J. Rodríguez, KaiR609 (M); near
Volcán, Paso Ancho, PNVB, N 08°48ʹ55.1ʺ, W 82°34ʹ42.6ʺ,
2000m asl., montane forest with abundant Quercus spp. and
Comarostaphylis arbutoides, 26 October 2016, J. Rodríguez,
T.A. Hofmann, K. Barrera, PA891 (M).
Notes: Entoloma flavoconicum is characterised by
mycenoid, strongly yellowish basidiocarps, conical to
acutely umbonate pilei, heterodiametrical basidiospores,
incrusting pigment on the cells of the pileipellis, and
absence of clamp connections. It forms a clade together
with E. olivaceohebes Noordel. & Hauskn. and E. vernum
S. Lundell (Fig.14). However, these two species have
basidiocarps with rather dark brown colours. In addition
Fig. 21 Basidiocarp and basidi-
ospores as seen by SEM of
Entoloma transitionisporum
(KaiR503, holotype): a stipe b
pileus from above; c basidi-
ospores, including reduced
forms, bar = 10µm
Fig. 22 Microscopic structures
of Entoloma transitionisporum
(CME6, holotype): a pileipel-
lis with subpellis composed of
inflated cells, intracellular pig-
ment not indicated, bar = 20μm;
b basidiospores, bar = 10μm
a
b
118 Mycological Progress (2022) 21:93–145
1 3
to this, they can be distinguished from E. flavoconicum
by micromorphological characters, like the presence of
clamp connections in the hymenium (Noordeloos 1992,
2004; Horak 2008). Entoloma subelegans Noordel. &
Hauskn., described from Mauritius (Noordeloos and
Hausknecht 2016), is somewhat similar in colour and
striation of the pileus. It differs, however, by depressed
pilei, shape and size of the basidiospores, presence of
cheilocystidia, and clamp connections in the hymenium.
Phylogenetically, E. subelegans does not belong to E.
subg. Nolanea (Fig.14). Entoloma aromaticum E. Horak
from Australia and New Zealand is another yellowish and
clampless species, but the colour of the basidiocarps is
somewhat more reddish yellow, and they have a strongly
aromatic smell and taste (Horak 2008; Noordeloos and
Gates 2012). No similar species is described in the mono-
graphs of Hesler (1967), Largent (1994), and Noordeloos
(2004).
Entoloma paraconferendum Reschke, Manz, F. Hampe
& Noordel., sp. nov., Figs.19 and 20
Mycobank number: MB840628.
Typification: PANAMA. CHIRIQUÍ: near Cerro
Punta, Entre Ríos, Montaña Azul, PILA, N 08°53ʹ39.7ʺ
W 82°34ʹ55.0ʺ, 2340m asl., montane forest dominated by
Quercus spp., 19 June 2018, C. Manz and F. Hampe CME6,
holotype (UCH11747), isotype (M).
Etymology: para (Latin) = beside; refers to Entoloma
conferendum, which is closely related and has morphologi-
cally similar basidiocarps.
Description: Basidiocarps mycenoid. Pileus 2.5–4.5cm
diameter, broadly conical to campanulate, slightly papil-
late, with straight, slightly uneven margin, greyish yellow
(4B4, 4B5), with darker greyish yellow (4C5), delimited
centre, whitish at the very margin, pileal surface glabrous,
but finely scaly-pruinose if seen through a lens, translucently
striate at about 1/3 the radius, hygrophanous. Lamellae
adnexed to almost free, ventricose, close, initially whitish,
later pink, with rather even and concolorous edges. Stipe
4.5–7.0
×
0.25–0.40cm, cylindrical, hollow, greyish yel-
low (4B4, 4B5), fibrillose, overlaid by whitish, silky fibrils.
Basal mycelium white, somewhat cottony. Odour somewhat
musty, taste not tested.
Basidiospores 8.5–10.5–12.0 × 6.5–7.3–8.0 µm,
Q = 1.25–1.44–1.70 (n =
54 spores of 2 specimens),
heptahedroid, composed of an adaxial facet (D), a pair
of apico-adaxial facets (AD), an apical facet (A), a pair
of lateral facets (L), and a basal facet (Ba) forming a
simple base, heterodiametrical, rarely irregularly cru-
ciform, with 4–6, predominantly 5, pronounced, blunt
angles in outline, weakly pigmented yellowish pink,
somewhat thick-walled. Basidia 30–42 ×
10.5–12.5µm,
clavate to subclavate, hyaline, 4-spored, sterigmata up
to 4.0µm long. Lamellar edge fertile, hymenial cystidia
absent. Hymenophoral trama regular, formed by long,
cylindrical to somewhat inflated cells, rather wide cen-
trally, narrower towards hymenium, 80–550 × 3.5–25µm.
Pileipellis a cutis composed of cylindrical cells,
45–145 × 5.0–12.0µm, often disrupted by trichodermal
sections formed by a few erect hyphal ends, composed
Fig. 23 Maximum likelihood
phylogram of species of subg.
Alboleptonia based on ITS,
rooted to E. serrulatum, boot-
strap values above branches,
bar = estimated changes/nucleo-
tide
0.05
KaiR672_Entoloma_amistadosericellum_ PAN
KaiR1214_Entoloma_sericellum_SWE
KaiR501_Entoloma_amistadosericellum_ PAN
KaiR670_Entoloma_confusissimum_holotype_PA
N
KaiR489_Entoloma_confusissimum_PAN
KaiR675_Entoloma_amistadosericellum_ PAN
KaiR502_Entoloma_amistadosericellum_ PAN
KC898453_Entoloma_sericellum_LE254362_RUS
KaiR429_Entoloma _nubilosilvae_PAN
KP965765_Alboleptonia_sericella_Lueck1_DEU
KaiR474_Entoloma _nubilosilvae_PAN
KaiR1203_Entoloma_sericellum_SWE
KC898447_Entoloma_serrulatum_LE254361_RUS
KaiR465_Entoloma _nubilosilvae_holotype_PAN
KaiR612_Entoloma_amistadosericellum_ holotype_PAN
KaiR490_Entoloma_amistadosericellum_ PAN
KaiR492_Entoloma_amistadosericellum_ PAN
KaiR495_Entoloma _amistadosericellum_ PAN
PDD104142_Entoloma_niveum_NZL
KaiR430_Entoloma_nubilosilva e_PAN
88
100
62
98
66
100
56
98
98
119Mycological Progress (2022) 21:93–145
1 3
of cells of the same size or of relatively short cells,
15.0–40 × 6.5–12.0 µm, pigment intracellular, with a
subpellis composed of inflated cells, 40–90 × 20–50µm.
Stipitipellis a cutis composed of long, cylindrical cells,
60–400 × 3.0–5.5µm. Caulocystidia absent. Oleiferous
hyphae frequently present in the trama. Clamp connec-
tions absent from all parts of the basidiocarp.
Habitat: Basidiocarps in small groups in Quercus-dom-
inated montane forest at 2340m asl. in Chiriquí, Panama.
Additional specimen examined: PANAMA. CHIRIQUÍ:
near Cerro Punta, Entre Ríos, Montaña Azul, PILA, N
08°53ʹ39.7ʺ W 82°34ʹ55.0ʺ, 2,340m asl., montane forest
dominated by Quercus spp., 19 June 2018, C. Manz and F.
Hampe, CME7 (M).
Notes: Entoloma paraconferendum is characterised
by mycenoid basidiocarps with a translucently striate
pileus, a silky, fibrillose stipe, and heterodiametrical
basidiospores. Its basidiocarps are macromorphologically
like those of E. conferendum. These two species are also
closely related based on ITS sequences (p-distance = 1.8%).
However, they can easily be distinguished by the almost
exclusively cruciform basidiospores of E. conferendum and
predominantly heterodiametrical, only rarely subcruciform
basidiospores of E. paraconferendum.
Entoloma transitionisporum Reschke, Manz & Noordel.,
sp. nov., Figs.21 and 22
Mycobank number: MB840626.
Typification: PANAMA. CHIRIQUÍ: Cerro Punta,
Las Nubes, PILA, Sendero Cascada, N 08°53ʹ48.0ʺ W
82°37ʹ05.7ʺ, 2320m asl., montane forest dominated by
Alnus acuminata, 11 June 2017, K. Reschke and J. Rod-
ríguez KaiR503, holotype (UCH11743), isotype (M).
Etymology: transitio (Latin)
=
transition; refers to the
transitional shape of the basidiospores, between heterodia-
metrical and cruciform.
Description: Basidiocarps mycenoid. Pileus 3.5–4.0cm
diameter, broadly conical to convex umbilicate, margin
slightly incurved and uneven, brown to light brown (5E7,
5D6, 5D7) at centre, paler, brownish orange to greyish orange
(5C4, 5C5 to 5B3), towards margin, pileal surface glabrous,
translucently striate up to 2/3 of the centre, hygrophanous.
Lamellae adnexed, almost free, medium-spaced to rather
close, white to slightly greyish in immature basidiocarps, pink
upon maturity, with slightly undulating and concolorous edges
and transvenose sides. Stipe 7.0 ×
0.4–0.5cm, cylindrical to
slightly broadening towards base, hollow, brownish orange
(5C5, 5C6), overlaid by silvery whitish fibrils, innately fibril-
lose, sometimes contorted. Basal mycelium not recorded.
Odour indistinct, taste not tested.
Basidiospores 7.5–9.2–10.5 × 6.0–6.6–7.5 µm,
Q = 1.15–1.40–1.65 (n =
65 spores of 2 specimens), pre-
dominantly heptahedroid, composed of an adaxial facet
(D), a pair of apico-adaxial facets (AD), an apical facet (A),
a pair of lateral facets (L), and a basal facet (Ba) forming
a simple base, reduced forms hexahedroid or more rarely
pentahedroid (prismatic), with single AD-facet and/or
reduced to absent A-facet, heterodiametrical to irregularly
cruciform, with 4–5 pronounced angles in outline, weakly
pigmented yellowish pink, somewhat thick-walled. Basidia
Fig. 24 Basidiocarps and
basidiospores as seen by SEM
of Entoloma amistadosericel-
lum: a (KaiR612, holotype);
b (KaiR502); c (KaiR672);
d (KaiR612, holotype)
bar = 10µm
120 Mycological Progress (2022) 21:93–145
1 3
34–41 × 11.0–13.0µm, subclavate, hyaline, 4-spored, ster-
igmata up to 5.0µm long. Lamellar edge fertile, hyme-
nial cystidia absent. Hymenophoral trama regular, formed
by long, cylindrical to somewhat inflated cells, centrally
140–430 × 6.5–20µm, narrower, 170–450 × 2.5–7.0 µm,
towards hymenium. Pileipellis a cutis of relatively narrow
cells, 70–110 × 3.0–9.0µm, with yellowish to brownish,
intracellular pigment and a distinct subpellis of inflated cells,
40–65 × 28–40µm. Stipitipellis a cutis composed of long,
cylindrical cells, 290–950 × 4.0–14.0µm, with some intra-
cellular, pale yellowish brown pigment, difficult to locate.
Caulocystidia absent. Oleiferous hyphae present, abundant
in hymenophoral trama. Clamp connections absent from all
parts of the basidiocarp.
Habitat: Basidiocarps found solitary among mosses in
Alnus acuminata-dominated montane forests between 2000
and 2400m asl. in Chiriquí, Panama.
Additional specimen examined: PANAMA. CHIRIQUÍ:
Cerro Punta, Las Nubes, PILA, Sendero Cascada, N
08°53ʹ37.5'ʺ W 82°36ʹ54.4ʺ, 2,030m asl., montane forest
dominated by Alnus acuminata, 22 June 2018, C. Manz and
T.A. Hofmann, CME9 (M).
Notes: Entoloma transitionisporum belongs to the clade
of E. conferendum and its allies. It is characterised by a coni-
cal to convex umbilicate, brown, translucently striate pileus,
a silvery fibrillose stipe, absence of clamps, and the shape of
the basidiospores, which can be interpreted as a transitional
state between heterodiametrical and cruciform. Entoloma
belouvense is closely related based on ITS sequence data
(Fig.14). Its basidiocarps are on average smaller and paler
than those of E. transitionisporum and their pilei have a
contrasting darker centre, which was not observed in pilei
of the latter species. In addition, the basidiospores of E.
belouvense are somewhat longer and without transitional
states to a cruciform shape. Basidiocarps of E. conferen-
dum can have a similar habit, but E. conferendum is eas-
ily distinguished from E. transitionisporum by exclusively
cruciform basidiospores. Entoloma luteifuscum K.N.A. Raj
& Manim., described from India, differs by yellowish tones
of the basidiocarps and slightly larger basidiospores, which
are exclusively heterodiametrical (Raj etal. 2014). Entoloma
dissimile (Singer) Horak, described from Nothofagus-forest
in Argentinia, may be similar to E. transitionisporum. How-
ever, the pilei of E. dissimile are darker, basidiospores are
slightly broader, and pigment is incrusting in the pileipel-
lis (Horak 1978). No similar species were found in Largent
(1994), Horak (1982), and Hesler (1967).
Subgenus
Alboleptonia
(Largent & R.G. Benedict)
Noordel
The subgenus Alboleptonia in the traditional sense (Lar-
gent and Benedict 1970; Noordeloos 1987, 2004) is an
assemblage of white or whitish species, often with a differ-
entiated pileipellis. It appears to be polyphyletic, considering
recent phylogenies (He etal. 2015c; Largent etal. 2016).
Here, we focus on the core group of Alboleptonia, i.e., spe-
cies around the type species E. sericellum (Fr.) P. Kumm.
This species is widespread in Europe and appears to be a
complex of several species, both in and outside of Europe.
Entoloma cuboidoalbum Noordel. & Hauskn. was described
from Austria (Noordeloos and Hausknecht 2009), but there
are more species waiting to be described in this group, as
evident by the results of the phylogenetic analysis (Fig.23).
In the present study, three new species from Panama are
Fig. 25 Microscopic structures
of Entoloma amistadosericel-
lum (KaiR612, holotype):
a pileipellis, bar = 20μm; b
basidiospores, bar = 10μm; c
cheilocystidia, bar = 20µm
a
c
b
121Mycological Progress (2022) 21:93–145
1 3
described in the clade of E. sericellum that can be well dif-
ferentiated by their basidiospore characters.
Entoloma amistadosericellum Reschke & Noordel., sp.
nov., Figs.24 and 25
Mycobank number: MB840937.
Typification: PANAMA. CHIRIQUÍ: near Cerro Punta,
Entre Ríos, Montaña Azul, PILA, around 2300m asl.,
montane forest dominated by Quercus spp., 21 June 2017,
J. Rodríguez, KaiR612, holotype (M).
Etymology: named after its similarity to Entoloma seri-
cellum and its occurrence in the Parque Internacional La
Amistad, honouring the principle of friendship in nature
conservation.
Description: Basidiocarps collybioid. Pileus 1.0–5.5cm
diameter, convex and often with an obtuse umbo when
Fig. 26 Basidiocarps and
basidiospores as seen by SEM
of Entoloma confusissimum: a
(KaiR489); b, c, d (KaiR670,
holotype) bars = 10µm
Fig. 27 Microscopic struc-
tures of Entoloma confusis-
simum (KaiR670, holotype): a
stipitipellis with caulocystidia,
bar = 20µm; b pileipellis
bar = 20µm; c basidiospores,
bar = 10µm; d cheilocystidia,
bar = 20µm
a
b
c
d
122 Mycological Progress (2022) 21:93–145
1 3
young, becoming depressed, often remaining a small papilla,
margin initially incurved, later straight, white in young and
fresh basidiocarps, older basidiocarps with yellowish to
pale brownish patches, pileal surface radially fibrillose, not
translucently striate. Lamellae sinuate, adnate to decurrent,
medium-spaced to rather distant, initially white, later pink,
with smooth to slightly uneven, concolorous edges. Stipe
1.0–4.7 × 0.20–0.60 cm, cylindrical to slightly clavate
towards base, rather fragile and hollow, translucently
whitish, inconspicuously fibrillose and slightly pruinose all
over. Basal mycelium white. Odour not distinctive, taste not
tested.
Basidiospores 8.5–9.4–10.5 × 6.5–7.6–8.5 µm,
Q = 1.10–1.25–1.40 (n = 111 spores of 5 specimens),
enneahedroid, composed of an adaxial facet (D), a pair of
apico-adaxial facets (AD), an apical facet (A), an abaxial
facet (B), and two pairs of lateral facets (L1–L2), of which
the basal pair (L2) forms a dièdre basal, reduced forms
octa- or heptahedroid, often with single AD-facet and/
or reduced to absent A-facet, mainly broadly heterodia-
metrical, some subisodiametrical, rhomboid, cruciform or
almost quadratic, with 4–6, predominantly 5, pronounced
angles in outline, weakly pigmented yellowish pink, some-
what thick-walled. Basidia 36–55 × 10.0–12.5µm, clavate,
hyaline, 4-spored, sterigmata up to 5.5µm long. Lamellar
edge almost sterile, cheilocystidia cylindrical, clavate to
broadly lageniform, sometimes capitate, 37–73 × 10–19µm.
Hymenophoral trama composed of rather broadly cylindri-
cal cells, 30–175
×
5–22µm. Pileipellis a cutis formed by
cylindrical cells, 70–175 × 3.5–11.5µm, pigment absent;
subpellis absent, gradually passing into pileitrama, com-
posed of somewhat wider cells. Stipitipellis a cutis com-
posed of cylindrical cells, 50–200
×
4.0–8.5µm. Caulo-
cystidia absent. Oleiferous hyphae present in the trama but
rather infrequent. Clamp connections abundant in all parts
of the basidiocarp.
Habitat: Basidiocarps solitary to scattered, in Alnus
acuminata-dominated montane forest and Quercus-dominated
montane forest at around 2300m asl. in Chiriquí, Panama.
Additional specimens examined: PANAMA.
CHIRIQUÍ: Cerro Punta, Las Nubes, PILA, Sendero Cas-
cada, N 08°53ʹ46.0ʺ W 82°37ʹ01.ʺ 2270m asl., montane
forest dominated by Alnus acuminata, 11 June 2017, K.
Reschke and J. Rodríguez, KaiR490 (UCH11736, M);
ibid., N 08°53ʹ46.8ʺ W 82°37ʹ01.4ʺ, 2280m asl., mon-
tane forest dominated by Alnus acuminata, 11 June 2017,
K. Reschke and J. Rodríguez, KaiR492 (UCH11737, M);
ibid., N 08°53ʹ46.8ʺ W 82°37ʹ01.4ʺ, 2280m asl., montane
forest dominated by Alnus acuminata, 11 June 2017, K.
Reschke and J. Rodríguez, KaiR495 (UCH11738, M); ibid.,
N 08°53ʹ48.0ʺ W 82°37ʹ05.7ʺ, 2320m asl., montane forest
dominated by Alnus acuminata, 11 June 2017, K. Reschke
and J. Rodríguez, KaiR501 (M); ibid., N 08°53ʹ48.0ʺ W
82°37ʹ05.7ʺ, 2320m asl., montane forest dominated by
Alnus acuminata, 11 June 2017, K. Reschke and J. Rod-
ríguez, KaiR502 (M); ibid., N 08°53ʹ46.2ʺ W 82°37ʹ01.4ʺ,
2280m asl., montane forest dominated by Alnus acuminata,
27 June 2017, K. Reschke, KaiR672 (M).
Notes: Entoloma amistadosericellum is characterised
by whitish basidiocarps with convex to depressed pilei, a
radially fibrillose pileal surface, abundant clamp connec-
tions, relatively broad cheilocystidia, and the presence of
rhomboid to cruciform basidiospores. It is closely related
to E. nubilosilvae, described below. Most species of subg.
Alboleptonia s.l. are macromorphologically similar, how-
ever, they differ in micromorphological characters. Ento-
loma cylindrocapitatum (T.J. Baroni & Ovrebo) Noordel.
& Co-David, also described from Panama, has rather small
basidiospores as well as long, narrow, and capitate cheilo-
cystidia (Ovrebo and Baroni 2007). Entoloma cuboidoal-
bum has larger basidiospores (9.0–11.5 × 7.5–10.5 µm)
and lower Q-values (1.0–1.3) (Noordeloos and Hausknecht
2009) than E. amistadosericellum (8.5–10.5 × 6.5–8.5µm,
Q = 1.10–1.40). Entoloma crocotillum Xiao L. He has
large, exclusively heterodiametrical basidiospores (He
etal. 2015c). Entoloma davidii Noordel. & Co-David (syn.
Alboleptonia largentii T.J. Baroni & Lodge), E. laccari-
oides T.H. Li etal. and E. subroseum (T.J. Baroni & Lodge)
Noordel. & Co-David have cuboid basidiospores (Baroni
and Lodge 1998; He etal. 2015a). Entoloma sericellum s.
auct. has a temperate and boreal distribution, and hetero-
diametrical, 5–8 angled basidiospores. Entoloma shwethum
Manim etal. has rather large basidiocarps (pileus 3–5cm
diameter), sterile lamellar edges, and large basidiospores
(9–12 × 6–9µm) (Manimohan etal. 1995). Entoloma subo-
chraceum Blanco-Dios (syn. Alboleptonia ochracea Largent
& R.G. Benedict) has basidiospores with an average length
of 10.5µm and only scattered clamp connections (Largent
and Benedict 1970). Basidiocarps of Entoloma theekshna-
gandhum Manim. etal. can be differentiated from those of
E. amistadosericellum by their odour of fermented rice soup
and cuboid to isodiametrical basidiospores (Manimohan and
Leelavathy 1988; Manimohan etal. 1995).
Entoloma confusissimum Reschke & Noordel., sp. nov.,
Figs.26 and 27
Mycobank number: MB840938.
Typification: PANAMA. CHIRIQUÍ: Cerro Punta,
Las Nubes, PILA, Sendero Cascada, N 08°53ʹ46.2ʺ W
82°37ʹ01.4ʺ, 2280m asl., montane forest dominated by
Alnus acuminata, 27 June 2017, K. Reschke, KaiR670,
holotype (M).
Etymology: confusus (Latin)
=
confused; refers to
the fact, that basidiocarps of this species can easily be
123Mycological Progress (2022) 21:93–145
1 3
confused with those of several other species of E. subg.
Alboleptonia.
Description: Basidiocarps mycenoid to collybioid. Pileus
0.5–5.0cm diameter, broadly conical to convex, becoming
depressed in old basidiocarps, margin initially somewhat
incurved, later straight, white in young and fresh basidi-
ocarps, older basidiocarps with slightly yellowish patches,
pileal surface finely fibrillose squamulose, not translucently
striate, not hygrophanous. Lamellae emarginate with decur-
rent tooth to decurrent, segmentiform, medium-spaced to
distant, sometimes distinctly transvenose, initially white,
later pink, with even to slightly undulating, concolorous
edges. Stipe 2.0–5.0 × 0.15–0.25cm, cylindrical, translu-
cently whitish, polished to slightly fibrillose and pruinose,
especially towards the apex, staining slightly yellowish after
damage. Odour indistinct, taste not tested.
Fig. 28 Basidiocarps and
basidiospores as seen by SEM
of Entoloma nubilosilvae: a
(KaiR429); b (KaiR465, holo-
type) bar = 1cm; c, d (KaiR465,
holotype) bars = 10µm
Fig. 29 Microscopic struc-
tures of Entoloma nubilosilvae
(KaiR465, holotype): a pileipel-
lis, bar = 20μm; b cheilocys-
tidia, bar = 20µm; c basidi-
ospores, bar = 10μm
bc
a
124 Mycological Progress (2022) 21:93–145
1 3
Basidiospores 9.0–10.3–12.0 × 6.5–7.2–8.0 µm,
Q = 1.35–1.44–1.60 (n = 44 spores of 2 specimens), tetra-
decahedroid, composed of an adaxial facet (D), a pair of
apico-adaxial facets (AD), an apical facet (A), two abaxial
facets (B1, B2), and four pairs of lateral facets (L1–L4), of
which the basal pair (L4) forms a sometimes incomplete
dièdre basal, heterodiametrical, with 6–9 nodulose angles
in outline, weakly pigmented yellowish pink, somewhat
thick-walled. Basidia 30–40 × 10.0–13.0µm, clavate to
subclavate, hyaline, predominantly 4-spored, with up to
4.0µm long sterigmata, few 2-spored. Lamellar edge het-
erogeneous, at some parts almost sterile, cheilocystidia
35–60 × 7.0–11.0µm, subcylindrical to lageniform, often
in clusters. Hymenophoral trama regular, mainly composed
of rather short, broadly cylindrical to somewhat inflated
cells, 70–200 × 7–25µm, longer and narrower cells towards
0.05
KaiR638_Entoloma_melleosquamulosum_PAN
KaiR534_Entoloma_griseocaeruleum_holotype_PAN
LN850561_Entoloma_sarcitulum_K378_holotype_GB
Entoloma_indutoides_G1397_WU19603_MK278000
KY774214_Entoloma_sp_NewCaledonia
UDB015211_MK278008_Entoloma_viiduense_G1602_Estonia
MW340899_Entoloma_norlandicum_O_F_76176_holotype_NO
OTU039_Entoloma_montanum_NOBAS418317_N41_03_09_2010
Entoloma_indutoides_cf_TU106924_UDB015261
KC898449_MH259311_Entoloma_inocephalum_LE262922_Vietnam
KaiR413_Entoloma_microserrulatum_ holotpye_PAN
CME2_Entoloma_cf_violaceobrunneum_PAN
MW934593_MW934255_Entoloma_subcoracis_holotype_RU
Entoloma_nigrovelutinum_LE_295077
KY462681_Entoloma_sp_Chile
KaiR536_Entoloma_griseocaeruleum_PAN
MW340896_Entoloma_montanum_GB_0191635_holotype_SW
Entoloma_ochromicaceum_UDB023715_AT250_DMS9201008
CME13_Entoloma_griseocaeruleum_PAN
MG947210_Entoloma_yanacolor_holotype_EC
E194_Entoloma_longistriatum_KY744164_USA
Entoloma_serrulatum_NOBAS219616_AT244_DMS730296
KY744177_Entoloma_subserrulatum_USA
KaiR535_Entoloma_caeruleomarginatum_holotype_PAN
Entoloma_aff_griseocyaneum_G0074_NL5097_USA
Entoloma_odoratum_AT323_DMS166826_Denmark
KP191935_Entoloma_sp_NewZealand
Entoloma_holmvassdalenense_DB6073_TEB271_16_NOBAS432617
OW135_Entoloma_roseotinctum_NOR
Cro16_Entoloma_cf_pseudoturci_HRV
KY777374_Entoloma_subfarinaceum_USA
KaiR614_Entoloma_arcanum_holotype_PAN
KaiR646_Entoloma_aff_necopinatum_PAN
KaiR1004_Entoloma_isborscianum_DEU
CME8_Entoloma_griseocaeruleum_PAN
KaiR488_Entoloma_arcanum_PAN
KaiR664_Entoloma_microserrulatum_ PAN
CME16_Entoloma_melleosquamulosum_holotype_PAN
CME3_Entoloma_caeruleomarginatum_PAN
KC898446_Entoloma_mougeotii_LE254352_RU
KY706167_Entoloma_subserrulatum_Canada
LN850627_Entoloma_querquedula_TUR_18_XI_2011_FI
WPR004_Entoloma_turci_DE
KY972699_Entoloma_subcaesiocinctum_China
MK693223_MK696392_Entoloma_nipponicum_TNS_F70747_JP
KM610321_Entoloma_holmvassdalenense_O_F75311_holotype_NO
KaiR990_Entoloma_incanum_DEU
KaiR997_Entoloma_griseocyaneum_DEU
KY706188_Entoloma_aff_griseocyaneum_Canada
MF476913_MF487803_Entoloma_violaceoserrulatum_TUR_JV_8329F_FI
MK277991_Entoloma_glaucobasis_S569_Roth16_6_95_G0209
KaiR632_Entoloma_cf_violaceobrunneum_PAN
UDB011680_Entoloma_cf_catalaunicum_Estonia
KY706185_Entoloma_sp_Canada
Cro26_Entoloma_mediterraneense_HRV
KX574458_Entoloma_cf_largentii_USA
KY744158_Entoloma_fuscosquamosum_US_TN
MW934571_MW934251_Entoloma_coracis_O_F_256850_holotype_NOR
KY774215_Entoloma_sp_NewCaledonia
CME11_Entoloma_caeruleomarginatum_PAN
KaiR511_Entoloma_sp_PAN
KY777373_Entoloma_cf_unicolor_PBM3995_USA
100
64
54
100
100
81
75
78
91
83
100
36
100
59 100
100
100
99
91
100
61
100
50
47
41
84
57
63
99
70
80
74
84
95
60
89
87
98
67
81
31
84
100
47
98
84
100
97
100
86
73
28
97
98
89
100
Fig. 30 Maximum likelihood phylogram of species of subg. Cyanula based on concatenated ITS and LSU sequences, rooted to E. inocephalum,
E. indutoides, and E. cf. indutoides, bootstrap values above or below branches, bar = estimated changes/nucleotide
125Mycological Progress (2022) 21:93–145
1 3
the subhymenium, approximately 150–300 × 4.0–5.0µm.
Pileipellis a cutis formed by cylindrical to somewhat fusi-
form or locally inflated cells, 70–250 × 3.0–16.0µm, with-
out pigment, subpellis not developed, pileipellis gradually
passing into pileitrama, composed of similar to somewhat
wider cells. Stipitipellis a cutis composed of cylindrical
cells, 60–180 × 4.5–8.0µm. Caulocystidia present in dense
bundles, cylindrical, often subcapitate, 30–65 (85) × (5.0)
6.5–10.0 (12.5) µm. Oleiferous hyphae present in the
trama. Clamp connections abundant in all parts of the
basidiocarp.
Habitat: Basidiocarps scattered in small groups in Alnus
acuminata-dominated montane forest between 2200 and
2300m asl. in Chiriquí, Panama.
Additional specimen examined: PANAMA.
CHIRIQUÍ: Cerro Punta, Las Nubes, PILA, Sendero Cas-
cada, N 08°53ʹ46.0ʺ W 82°37ʹ01.5ʺ, 2270m asl., montane
forest dominated by Alnus acuminata, 11 June 2017, K.
Reschke and J. Rodríguez, KaiR489 (UCH11735, M).
Notes: Entoloma confussisimum is characterised by
its E. sericellum-like habit and rather large and complex
basidiospores with 6–9 angles in outline. Its ITS sequences
have a p-distance of 5% to that of a specimen identified
as E. sericellum from Caucasus, Russia (Fig.23). Ento-
loma sericellum differs by smaller basidiospores with
lower Q-values (1.1–1.5) and 5–8, predominantly 6
angles (Noordeloos 1992). Entoloma subsericellum Mur-
rill, described from a coniferous forest of North America,
has basidiospores of similar size, however, with only
5–6 angles (Murrill 1917; Largent and Benedict 1970).
The absence of cheilocystidia, rare clamp connections,
and a farinaceous odour indicates that E. subsericellum
does not belong to the core clade of subg. Alboleptonia.
Entoloma aripoanum Dennis, described from Trinidad, is
similar in macromorphology and spore size, but differs
by its very narrow lamellae, large, cylindrical cheilocyst-
idia (70–160 × 10–16µm), and occurrence on rotten wood
(Dennis 1953).
Entoloma nubilosilvae Reschke & Noordel., sp. nov.,
Figs.28 and 29
Mycobank number: MB840623.
Typification: PANAMA. CHIRIQUÍ: near Boquete,
PNVB, Sendero Culebra, N 08°51ʹ12.5ʺ W 82°29ʹ01ʺ,
1,830m asl., montane forest with abundant Quercus spp.,
9 June 2017, K. Reschke, KaiR465, holotype (UCH11733),
isotype (M).
Etymology: nubilus (Latin) = cloudy, silva (Latin) = for-
est; refers to the occurrence in cloud forest.
Description: Basidiocarps collybioid. Pileus 0.8–2.7cm
diameter, convex, becoming depressed in old basidiocarps,
margin initially incurved, later straight, white in young and
fresh basidiocarps, older basidiocarps with yellowish to
pale brownish patches, pileal surface radially fibrillose, not
translucently striate. Lamellae slightly to distinctly decur-
rent, rather distant, initially white, later pink, with slightly
uneven and finely pruinose, concolorous edges. Stipe
1.8–3.5
×
0.15–0.50cm, cylindrical, rather fragile and hol-
low, translucently whitish, inconspicuously fibrillose and
slightly pruinose all over, staining yellowish after damage.
Odour sometimes perfume-like, taste not tested.
Basidiospores 7.0–8.8–10.0 × 6.0–7.1–8.0 µm,
Q = 1.05–1.24–1.40 (n =
137 spores of 4 specimens), octa-
hedroid, composed of a adaxial facet (D), predominantly
a single apico-adaxial facet (sAD), an apical facet (A), an
abaxial facet (B), and two pairs of lateral facets (L1–L2),
of which the basal pair (L2) forms a dièdre basal, subiso-
diametrical to broadly heterodiametrical, often rhomboid to
almost quadratic or cruciform, with 4–5 pronounced angles
in outline, weakly pigmented yellowish pink, somewhat
thick-walled. Basidia 36–45 × 9.0–10.5µm, clavate, hya-
line, mainly 4-spored with up to 5.0µm long sterigmata,
some 2-spored. Lamellar edge heterogeneous, cheilocystidia
40–60 × 5.0–13.0µm, subcylindrical, lageniform to capitate,
often in clusters, with variable abundance. Hymenophoral
trama regular, composed of rather short, broadly cylindrical
cells, 40–100 × 5.0–18.0µm. Pileipellis a cutis formed by
Fig. 31 Basidiocarps of Ento-
loma arcanum: a (KaiR488); b
(KaiR614)
126 Mycological Progress (2022) 21:93–145
1 3
cylindrical to subcylindrical cells, 65–250 × 4.5–12.0µm,
pigment absent, without subpellis, gradually passing into
pileitrama, composed of somewhat wider cells. Stip-
itipellis a cutis composed of narrow cylindrical cells,
60–210
×
3.0–6.0µm. Caulocystidia absent. Oleiferous
hyphae rare, but present at least in the pileitrama. Clamp
connections abundant in all parts of the basidiocarp.
Habitat: Basidiocarps found solitary to scattered in
Quercus-dominated montane forest between 1750 and
1900m asl. in Chiriquí, Panama.
Additional specimens examined: PANAMA.
CHIRIQUÍ: near Boquete, PNVB, Sendero Culebra, N
08°51ʹ00.7ʺ W 82°28ʹ55.2ʺ, 1,760m asl., montane forest
with abundant Quercus spp., 6 June 2017, K. Reschke,
KaiR429 (M); ibid., N 08°51ʹ00.7ʺ W 82°28ʹ55.2ʺ,
1,760m asl., montane forest with abundant Quercus
spp., 6 June 2017, K. Reschke, KaiR430 (M); ibid., N
08°51ʹ21.1ʺ W 82°29ʹ02.5ʺ, 1880m asl., montane forest
with abundant Quercus spp., 9 June 2017, K. Reschke,
KaiR474 (M).
Notes: Entoloma nubilosilvae is characterised by whitish
basidiocarps with a convex to depressed pileus, a radially
fibrillose pileal surface, abundant clamp connections,
rather narrow cheilocystidia, and the presence of rhomboid
to cruciform basidiospores. It is closely related to E.
amistadosericellum, described above, which has broader
cheilocystidia and slightly larger basidiospores. As
basidiocarps of E. nubilosilvae are quite similar to those of
E. amistadosericellum micro- and macromorphologically,
it differs from all the species mentioned in the notes of the
latter in the same way. Entoloma adnatifolium (Murrill)
Blanco-Dios has basidiospores and cheilocystidia of similar
size as E. nubilosilvae; however, rhomboid basidiospores
were not mentioned in the description of E. adnatifolium and
clamp connections are rare to scattered on the hyphae of the
pileipellis and the stipe surface (Largent and Benedict 1970).
Fig. 32 Microscopic struc-
tures of Entoloma arcanum:
a–c (KaiR488); a pileipellis,
bar = 20µm; b stipitipellis with
caulocystidia, bar = 20µm; c
basidiospores, bar = 10µm;
d (KaiR614) cheilocystidia,
bar = 20µm
a
b
c
d
127Mycological Progress (2022) 21:93–145
1 3
Subgenus Cyanula (Romagn.) Noordel
Species of the subgenus Cyanula form a well-supported
clade in the ITS/LSU phylogeny (Fig.30). They develop
basidiocarps with often bright colours, a differentiated
pileipellis with a trichodermal structure, intracellular pig-
ment, and clampless hyphae. The lamellar edge of many
species is formed by tramal hyphae that grow through the
hymenium forming a sterile band along the edge, with dense
clusters of more or less clavate or fusiform terminal cells.
These terminal cells often contain blue or brown intracel-
lular pigment, causing a coloured border along the edge, the
“serrulatum-type” lamellar edge (Noordeloos 2004). Spe-
cies of this clade occur worldwide, from the arctic tundra to
the tropical rainforest. In Europe, they are characteristic of
oligotrophic grasslands, habitats which are generally threat-
ened in Europe (Janssen etal. 2016). The subgenus Cyanula
is the most species-rich subgenus within Entoloma, with
approximately 500–600 species already described (Hesler
1967; Horak 1980, 2008; Largent 1994; Noordeloos 2004;
Noordeloos and Gates 2012; Crous etal. 2021; Noordeloos
etal. 2021; Dima etal. 2021). Species delimitation is often
rather difficult. Morphological look-alikes occurring on sev-
eral continents are often phylogenetically distant and must
therefore be considered as semi-cryptic.
Entoloma arcanum Reschke & Noordel., sp. nov.,
Figs.31 and 32
Mycobank number: MB840624.
Typification: PANAMA. CHIRIQUÍ: Cerro Punta,
Las Nubes, PILA, Sendero Cascada, N 08°53ʹ46.0ʺ W
82°37ʹ01.5ʺ, 2,270m asl., montane forest dominated by
Alnus acuminata, 11 June 2017, K. Reschke and J. Rod-
ríguez, KaiR488, holotype (UCH11734), isotype (M).
Etymology: arcanus (Latin) = arcane; refers to the dark
appearance of the basidiocarps and its isolated position in
the phylogenetic tree.
Description: Basidiocarps collybioid. Pileus 3.0–4.5cm
diameter, hemispherical to convex with umbilicate centre,
margin initially incurved, remaining somewhat incurved in
old basidiocarps, blackish blue (19F5, 19F6, 19F7), blue tones
becoming brownish while ageing, pileal surface squamulose-
velutinous in the centre, squamulose to squamulose-fibrillose
towards margin, not translucently striate, not hygrophanous.
Lamellae broadly adnate to slightly decurrent, segmentiform,
close, white with creamish tinge, becoming pink at maturity, with
somewhat crenate and concolorous edges. Stipe 2.0–4.0 × 0.5cm,
cylindrical, sometimes compressed, hollow, initially concolorous
to the pileus, blue tones soon becoming brownish, finely
squamulose. Basal mycelium white, cottony, sometimes with
obvious, thick rhizomorphs. Odour unpleasant, similar to old
basdidiocarps of Tricholoma spp. or burnt potatoes, taste not
tested.
Basidiospores 9.5–10.3–11.0 × 6.5–7.0–7.5 µm,
Q = 1.30–1.48–1.65 (n =
53 spores of 2 specimens), hetero-
diametrical, with 5–6 pronounced angles in outline, weakly
pigmented yellowish pink, somewhat thick-walled. Basidia
30–38 (40) × 8.5–11.0µm, clavate to subclavate, hyaline,
Fig. 33 Basidiocarp and cellular
structures as seen by SEM of
Entoloma caeruleomarginatum
(KaiR535, holotype): a view
from below, depicting lamellae
with serrulatum structure; b
view from above c basidiospore,
bar = 10µm; d cheilocystidia,
bar = 10µm
128 Mycological Progress (2022) 21:93–145
1 3
4-spored, with up to 5.5µm long sterigmata. Lamellar edge
sterile, cheilocystidia predominantly broadly fusiform,
sometimes lageniform, more rarely irregularly cylindrical
to moniliform, 25–55 × 8.0–14.0µm. Hymenophoral trama
regular, formed by subcylindrical cells, somewhat constricted
at septae, 55–275 × 3–20µm. Pileipellis a cutis with transi-
tions to a trichoderm, composed of broadly cylindrical to
inflated end cells, 40–140 × 10.5–19.0µm, upper pileipellis
cells 43–110 × 5.0–15.0µm, and a subpellis of narrower cells,
4.0–7.0µm wide, pigment intracellular as clumps, violet in
water, soluble and then brownish in KOH. Stipitipellis simi-
lar to pileipellis, composed of subcylindrical to somewhat
inflated cells, 50–180 × 7.5–15.0µm, with emerging end cells
in bundles and below a layer of narrower hyphae, 4.0–7.5µm
wide. Caulocystidia absent. Oleiferous hyphae present in the
trama. Brilliant granules abundant. Clamp connections absent
from any part of the basidiocarp.
Habitat: Basidiocarps solitary to scattered in a small
group in Quercus- and Alnus-dominated montane forests
between 2250 and 2300m asl. in Chiriquí, Panama.
Additional specimen examined: PANAMA.
CHIRIQUÍ: near Cerro Punta, Entre Ríos, Montaña Azul,
PILA, approximately 2300m asl., montane forest domi-
nated by Quercus spp., 21 June 2017, E. Heinemann,
KaiR614 (M).
Notes: Entoloma arcanum is characterised by dark,
blackish blue basidiocarps and a sterile lamellar edge with
mainly fusiform cheilocystidia. It is phylogenetically rela-
tively close to a specimen identified as E. fuscosquamosum
Hesler (Fig.30). This species has basidiocarps with more
greyish colours, larger basidiospores (9–13 × 6–7.5µm), and
cylindric-capitate cheilocystidia (Hesler 1967) or no cheilo-
cystidia at all (Noordeloos 1988). The RPB2 sequence of
E. arcanum is close to a sequence (KR233903) labelled E.
trichomatum (Largent) Noordel. & Co-David with a p-dis-
tance of 3%. This species is morphologically similar to E.
arcanum but differs by clavate to cylindrical cheilocystidia.
Entoloma trichomatum is known from coniferous forests in
western North America (Largent 1994), while E. arcanum
was found in tropical montane forests with broadleaved
trees.
Entoloma caeruleomarginatum Reschke, Manz, &
Noordel., sp. nov., Figs.33 and 34
Mycobank number: MB840627.
Typification: PANAMA. CHIRIQUÍ: near Bajo Boquete,
Jaramillo Arriba, El Musgo, N 08°47ʹ27.3ʺ W 82°24ʹ34.3ʺ,
1660m asl., montane forest dominated by Quercus spp., 14
June 2017, K. Reschke and M. Cuevas, KaiR535, holotype
(UCH11744), isotype (M).
Etymology: caeruleus (Latin)
=
dark blue, margo
(Latin) = edge; refers to the dark blue lamellar edge.
Description: Basidiocarps collybioid to some-
what omphalinoid. Pileus 1.5–2.0cm diameter, convex
Fig. 34 Microscopic structures
of Entoloma caeruleomargi-
natum (KaiR535, holotype):
a pileipellis, bar = 20µm; b
basidiospores, bar = 10µm;
c stipitipellis, bar = 20µm; d
cheilocystidia, bar = 20µm
a
b
d
c
129Mycological Progress (2022) 21:93–145
1 3
umbilicate with incurved margin in young basidiocarps,
expanded umbilicate with straight margin in older basidio-
carps, bluish grey to greyish blue (20C2, 20D6), centre and
squamules dark blue (20E4) to blackish blue (20F8), pileal
surface squamulose, not translucently striate in young and
fresh basidiocarps, but somewhat translucently striate
at the margin of old and soaked basidiocarps. Lamellae
broadly adnate, slightly emarginate to decurrent, segmenti-
form to arcuate, medium-spaced to rather distant, pale blue
in young basidiocarps, later pinkish, edges blackish blue
(20F6) serrulate. Stipe 4.0–5.0 × 0.30–0.35cm, cylindrical,
hollow, bluish grey to greyish blue (20D3, 21C6, 21C7,
21D7) with dark blue (20E5) fine, fibrillose squamules.
Bluish tones discolouring to brown in old basidiocarps.
Basal mycelium white, cottony. Odour indistinct, taste not
tested.
Basidiospores 9.0–9.8–11.0 × 6.0–7.0–8.0 µm,
Q = 1.20–1.41–1.60 (n = 69 spores of 3 specimens),
Fig. 35 Basidiocarps and cel-
lular structures as seen by SEM
of Entoloma griseocaeruleum:
a, b (KaiR534, holotype)
bar = 10µm; c (KaiR536); d
(KaiR534, holotype) cheilocyst-
idia, bar = 10µm
Fig. 36 Microscopic structures
of Entoloma griseocaeruleum
(KaiR534, holotype): a pileipel-
lis, bar = 20µm; b cheilocys-
tidia, bar = 20µm; c basidi-
ospores, bar = 10µm
a
b
c
130 Mycological Progress (2022) 21:93–145
1 3
hendecahedroid, composed of an adaxial facet (D), a pair
of apico-adaxial facets (AD), an apical facet (A), an abax-
ial facet (B), and three pairs of lateral facets (L1–L3), of
which the basal pair (L3) forms a dièdre basal, heterodia-
metrical, with 5–6 (7) pronounced, sometimes rather nodu-
lose angles in outline, weakly pigmented yellowish pink,
somewhat thick-walled. Basidia 34–48 × 8.0–11.0µm, sub-
clavate, hyaline, 4-spored, with up to 4.0µm long sterig-
mata. Lamellar edge sterile, of the serrulatum-type, cheilo-
cystidia 40–85 (135) × (5.5) 7.5–12.0 (17.0) µm, lageniform,
subfusiform to cylindrical, sometimes subcapitate, with
dark violet, intracellular pigment in clumps, soluble, and
then becoming brown in KOH. Hymenophoral trama regu-
lar, formed by long, cylindrical to slightly fusiform cells,
55–175 × 6.0–16.0µm. Pileipellis a cutis with transitions to a
trichoderm, especially towards centre, composed of clavate,
sometimes fusiform end cells, 70–210 × 13–26µm, with vio-
let, intracellular pigment, cells narrower, 6.0–12.0µm wide,
towards pileitrama, cylindrical to slightly fusiform. Stipiti-
pellis similar to pileipellis, composed of cylindrical cells,
60–210 × 4.0–7.5 (9.5) µm, with subcylindrical to somewhat
fusiform terminal cells, 75–150 × 10.5–16.0µm. Oleiferous
hyphae absent or not distinctive. Clamp connections absent
from any part of the basidiocarp.
Habitat: Basidiocarps found solitary in a Quercus-
dominated montane forest between 1650 and 1700m asl.
in Chiriquí, Panama.
Additional specimens examined: PANAMA.
CHIRIQUÍ: near Bajo Boquete, Jaramillo Arriba, El Musgo,
N 08°47ʹ26.9ʺ W 82°24ʹ34.4ʺ, 1670m asl., montane forest
dominated by Quercus spp., 15 June 2018, C. Manz and F.
Hampe, CME3 (M); ibid., 24 June 2018, C. Manz and F.
Hampe, CME11 (M).
Notes: Entoloma caeruleomarginatum belongs to the
clade of E. serrulatum and closely related species. It is
characterised by basidiocarps with a dark blue pileus,
blue serrulate lamellar edges, a fibrillose-squamulose
stipe surface, and heterodiametrical basidiospores with
an average length of approximately 10µm. The European
E. serrulatum is morphologically very similar and hard to
distinguish. The latter, however, is widely distributed and
common in northern hemisphere temperate to boreal grass-
lands. It is phylogenetically rather distant with a p-distance
of 10.5% in the ITS sequences. For this reason, we con-
sider them semi-cryptic species, separated by ecological
and geographical characters as well as molecular sequence
data. Both, E. subserrulatum (Peck) Hesler and E. subcae-
siocinctum Xiao-Lan He & W.H. Peng are phylogeneti-
cally closer to E. caeruleomarginatum based on ITS/LSU-
data than E. serrulatum (Fig.30). Entoloma subserrulatum
differs by basidiocarps with a pale and translucently striate
pileus (Hesler 1967). Entoloma subcaesiocinctum differs
from E. caeruleomarginatum by basidiocarps with a pale
pileus and by smaller basidiospores (He etal. 2017). Ento-
loma proximum Horak, described from Argentina, differs
by basidiocarps with an appressed fibrillose to subsqua-
mulose pileal surface, slightly larger basidiospores, and
relatively short, cylindrical to subclavate cheilocystidia
(Horak 1978).
Entoloma griseocaeruleum Reschke, Manz & Noordel.,
sp. nov., Figs.35 and 36
Mycobank number: MB840630.
Typification: PANAMA. CHIRIQUÍ: near Bajo
Boquete, Jaramillo Arriba, El Musgo, N 08°47ʹ26.1ʺ W
82°24ʹ29.9ʺ, 1,650m asl., montane forest dominated by
Quercus spp., 14 June 2017, K. Reschke, KaiR534, holo-
type (UCH9185), isotype (M).
Etymology: griseus (Latin)
=
grey, caeruleus
(Latin) = dark blue; refers to the colour of the basidiocarps.
Description: Basidiocarps collybioid to somewhat
omphalinoid. Pileus 1.5–3.5cm diameter, hemispherical to
convex with depressed to umbilicate centre, margin initially
incurved, later straight, initially greyish blu