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Abstract

In this 7th contribution to the Fungal Systematics and Evolution series published by Sydowia, the authors formally describe 14 species: Cantharomyces paschalis, Cryptandromyces pinguis, C. tricornis, Laboulbenia amblystomi (Laboulbeniales); Cortinarius squamosus, Entoloma brunneicoeruleum, E. callipygmaeum, E. minutigranulosum, E. perasprellum, E. pulchripes, E. tigrinum, E. timidum, E. violaceoserrulatum (Agaricales); and Suillus quercinus (Boletales). The following new country records are reported: Crepidotus malachioides from Italy, Leucoagaricus mucrocystis from French Guiana, Pluteus multiformis from Turkey (Agaricales); Herpomyces periplanetae from Bénin, the D.R. Congo, and Togo (Herpomycetales); Melanustilospora ari from Pakistan (Urocystidales); Neopestalotiopsis clavispora causing fruit rot on Zizyphus mauritiana from India (Amphisphaeriales); and Phytopythium chamaehyphon and Pp. litorale from Brazil (Peronosporales). Finally, a new combination is proposed based on morphology, ecology, and phylogenetic analysis: Rhodocollybia asema (Agaricales).
Sydowia 73 (2021) 271
DOI 10.12905/0380.sydowia73-2021-0271 Published online 14 June 2021
Fungal Systematics and Evolution: FUSE 7
Bálint Dima1, Tor Erik Brandrud2, Gilles Corriol3, Gerrit Maarten Jansen4, John Bjarne Jordal5,
Abdul N. Khalid6, Ellen Larsson7, Jostein Lorås8, Olga V. Morozova9, Arooj Naseer6, Machiel Evert
Noordeloos4, Walter Rossi10, Sergi Santamaria11, Samina Sarwar12, Ertug˘rul Sesli13, Muhammad Usman6,
Najam ul Sehar Afshan6, Ishtiaq Ahmad14, Arghya Banerjee15,16, Koushik Banerjee17, Egil Bendiksen18,
Débora Rodrigues da Silva Colombo19, André De Kesel20, Francesco Dovana21, Giuliano Ferisin22,
Shah Hussain23, Saidul Islam16,24, Ana Lucia Jesus19, Og˘ uzhan Kaygusuz25, Irmgard Krisai-Greilhuber26,
Sultan Mahammad27, Dilip Kumar Mishra28, Partha Sarathi Nath16, Sarah Cristina de Oliveira da Paixão19,
Birendranath Panja16, Viktor Papp29, Carmen Lidia Amorim Pires-Zottarelli19, Ágnes Radnóti29,
Debashis Rana16, Raina Saha30, I
˙brahim Türkekul31 & Danny Haelewaters32,33,*
1 Department of Plant Anatomy, Institute of Biology, Eötvös Loránd University, 1117 Budapest, Hungary
2 Norwegian Institute for Nature Research, 0855 Oslo, Norway
3 Conservatoire botanique national des Pyrénées et de Midi-Pyrénées, 65203 Bagnères-de-Bigorre, France
4 Naturalis Biodiversity Center, 2300 RA Leiden, The Netherlands
5 Miljøfaglig Utredning, 6630 Tingvoll, Norway
6 Department of Botany, University of the Punjab, Quaid-e-Azam Campus, 54590 Lahore, Pakistan
7 Gothenburg Global Biodiversity Center, Department of Biological and Environmental Sciences, University of Gothenburg,
405 30 Göteborg, Sweden
8 Nord University Nesna, 8700 Nesna, Norway
9 Komarov Botanical Institute of the Russian Academy of Sciences, 197376 Saint Petersburg, Russia
10 Sect. Environmental Sciences, Department of Life, Health and Environmental Sciences (MeSVA), University of L’Aquila,
67100 Coppito, Italy
11 Unitat de Botànica, Departament de Biologia Animal, de Biologia Vegetal i d’Ecologia, Facultat de Biociències, Universitat
Autònoma de Barcelona, 08193 Cerdanyola del Vallès (Barcelona), Spain
12 Department of Botany, Lahore College for Women University, 54000 Lahore, Pakistan
13 Department of Biology Education, Fatih Faculty of Education, Trabzon University, 61335 Trabzon, Turkey
14 Department of Botany, Islamia College Peshawar, 25120 Peshawar, Pakistan
15 Plant Quarantine Station, Haldia, West Bengal 721604, India
16 Department of Plant Pathology, Bidhan Chandra Krishi Viswavidyalaya, Mohanpur, West Bengal 741252, India
17 Division of Agricultural Physics, Indian Agricultural Research Institute (ICAR), New Delhi 110012, India
18 Norwegian Institute for Nature Research, 0855 Oslo, Norway
19 Instituto de Botânica, Núcleo de Pesquisa em Micologia, São Paulo, SP CEP 04301-912, Brazil
20 Meise Botanic Garden, 1860 Meise, Belgium
21 Department of Life Sciences and Systems Biology, University of Torino, 10125 Torino, Italy
22 Associazione Micologica Bassa Friulana, 33052 Cervignano del Friuli, Italy
23 Center for Plant Sciences and Biodiversity, University of Swat, 19130 Swat, Pakistan
24 Krishi Vigyan Kendra, Gayeshpur, West Bengal 741234, India
25 Department of Plant and Animal Production, Atabey Vocational School, Isparta University of Applied Sciences, 32670
Isparta, Turkey
26 Department of Botany and Biodiversity Research, University of Vienna, 1030 Wien, Austria
27 Institute for Agriculture Sciences and Forestry, University of Swat, 19130 Swat, Pakistan
28 ICAR–AICRP on Fruits, Directorate of Research, Bidhan Chandra Krishi Viswavidyalaya, Kalyani, West Bengal 741235, India
29
Department of Botany, Institute of Agronomy, Hungarian University of Agriculture and Life Sciences, 1118 Budapest, Hungary
30 Department of Plant Pathology, Uttar Banga Krishi Viswavidyalaya, Cooch Behar, West Bengal 736165, India
31 Department of Biology, Faculty of Science and Arts, Tokat Gaziosmanpas¸a University, 60010 Tokat, Turkey
32 Research Group Mycology, Department of Biology, Faculty of Sciences, Ghent University, 9000 Ghent, Belgium
33 Faculty of Science, University of South Bohemia, 370 05 C
ˇeské Budeˇjovice, Czech Republic
* e-mail: danny.haelewaters@gmail.com
Dima B., Brandrud T.E., Corriol G., Jansen G.M., Jordal J.B., Khalid A.N., Larsson E., Lorås J., Morozova O.V., Naseer A.,
Noordeloos M.E., Rossi W., Santamaria S., Sarwar S., Sesli E., Usman M., Afshan N.S., Ahmad I., Banerjee A., Banerjee K.,
Bendik-
sen E., Colombo D.R.S., De Kesel A., Dovana F., Ferisin G., Hussain S., Islam S., Jesus A.L., Kaygusuz O., Krisai-Greilhuber I.,
Mahammad S., Mishra D.K., Nath P.S., da Paixão S.C.O., Panja B., Papp V., Pires-Zottarelli C.L.A., Radnóti Á., Rana D., Saha R.,
Türkekul I
˙. & Haelewaters D. (2021) Fungal Systematics and Evolution: FUSE 7. – Sydowia 73: 271–340.
272 Sydowia 73 (2021)
Dima et al.: FUSE 7
In this 7th contribution to the Fungal Systematics and Evolution series published by Sydowia, the authors formally describe
14 species: Cantharomyces paschalis, Cryptandromyces pinguis, C. tricornis, Laboulbenia amblystomi (Laboulbeniales); Corti-
narius squamosus, Entoloma brunneicoeruleum, E. callipygmaeum, E. minutigranulosum, E. perasprellum, E. pulchripes, E. tigri-
num, E. timidum, E. violaceoserrulatum (Agaricales); and Suillus quercinus (Boletales). The following new country records are
reported: Crepidotus malachioides from Italy, Leucoagaricus mucrocystis from French Guiana, Pluteus multiformis from Turkey
(Agaricales); Herpomyces periplanetae from Bénin, the D.R. Congo, and Togo (Herpomycetales); Melanustilospora ari from Paki-
stan (Urocystidales); Neopestalotiopsis clavispora causing fruit rot on Zizyphus mauritiana from India (Amphisphaeriales); and
Phytopythium chamaehyphon and Pp. litorale from Brazil (Peronosporales). Finally, a new combination is proposed based on
morphology, ecology, and phylogenetic analysis: Rhodocollybia asema (Agaricales).
Keywords: 14 new species, 8 new records, 1 new combination, Agaricomycetes, Entolomataceae, integrative taxonomy, La-
boulbeniomycetes, Oomycota, Sordariomycetes, Ustilaginomycetes.
Materials and methods
Sample collection, isolation, and specimen exami-
nation
Most of the host insects bearing thalli of Can-
tharomyces Thaxt., Cryptandromyces Thaxt., and
Laboulbenia Mont. & C.P. Robin were supplied to S.
Santamaria and W. Rossi by entomologists. The host
insects of Laboulbenia amblystomi sp. nov. from
Spain and Thailand were collected in the eld ei-
ther by hand or by light trap. Permanent slide
mounts were prepared following the method de-
scribed by Rossi & Santamaria (2015) and were de-
posited at FI or BCB. Photomicrographs were cap-
tured with a ProgRes 10 Plus digital camera (Jenop-
tik, Jena, Germany) on a Leica DMR microscope
(Leice Biosystems, Wetzlar, Germany) with differen-
tial interference contrast optics (DIC); images were
processed with Photoshop CS5 (San Jose, Califor-
nia) and Dpx View Pro (Deltapix, Måløv, Denmark).
Basidiomata of Cortinarius (Pers.) Gray were
collected in Çilekli Village, Trabzon, Turkey. Photo-
graphs were taken in the laboratory. One of the ba-
sidiomata was used for a spore print; other basidi-
omata were dried for further studies and vouchering
purposes. Free-hand sections of pileus and lamellae
were treated in 5 % NH4OH, some of them were
stained with 1 % Congo red and photographed. Mi-
croscopic structures were observed and photo-
graphed under an Axio Imager A2 trinocular micro-
scope with Axio Imager software (ZEISS, Jena,
Germany). The description follows previous litera-
ture, including Moser et al. (1995), Bidaud et al.
(1997), Høiland & Holst-Jensen (2000), Moser (2002),
Bidaud et al. (2006), Niskanen & Kytövuori (2008),
and Niskanen et al. (2012). Specimens are deposited
in the herbarium of the Fatih Education Faculty at
Trabzon University (KATO).
Collections of Entoloma spp. were photographed
in the eld. Macroscopic characters were noted
straight after collecting. Color codes in the descrip-
tions follow Kornerup & Wanscher (1978), except
for Entoloma perasprellum sp. nov. that follows
Munsell Color Company (1954). Microscopic char-
acters were studied with a Leica DMLS microscope,
using a drawing tube and a Touptek Phototronics
camera (Zhejiang, China); a ZEISS Axioscope A1
microscope with AxioCam 1Cc 3; and a ZEISS Axi-
ophot microscope with DC controlled Cree XP-G3
R3 CRI 90+ LED illumination, Plan Neouar objec-
tives 40×/1.30 Oil, 100×/1.30 Oil (ZEISS), differen-
tial interference contrast (DIC) optics, a 12MP
Touptek video camera with SONY Exmor IMX226
CMOS sensor (Tokyo, Japan), and Toupview video &
image processing software (Touptek Photonics). Ba-
sidiospores, basidia, and cystidia were observed in
squash preparations of small parts of the lamellae
in 5 % KOH or 1 % Congo Red in concentrated
NH4OH. Pileipellis was examined on a radial sec-
tion of the pileus in 5 % KOH. Stipitipellis was ex-
amined in 10 % Ammonia solution. Basidiospore
dimensions are based on measuring 20 spores, di-
mensions of cystidia and basidia on observing at
least 10 structures per collection. Basidia were
measured without sterigmata, and the basidio-
spores without apiculus. Spore length to width ra-
tios are reported as Q. Collections are deposited at
the following herbaria: BBF, GB, L, LE, O, WU (sen-
su Thiers continuously updated).
For the Suillus Gray study, specimens were col-
lected from moist temperate forests in Khyber Pa-
khtunkhwa Province, Pakistan. Basidiocarps were
photographed in the eld. Fresh morphological
characters were recorded in the eld; colors follow
Munsell Color Company (1954). Field data on ba-
sidiocarps (site, habitat, association, etc.) was not-
ed. Specimens were studied microscopically follow-
ing Bessette et al. (2000). For basidiospore dimen-
sions, the rst and last values represent the range of
lengths and widths, whereas the values in parenthe-
ses represent mean lengths and widths ± standard
deviations; Q, length/width ratio of an individual
basidiospore; Qav, the mean of Q. Other measure-
ments are given as a range with outlier values in
Sydowia 73 (2021) 273
Dima et al.: FUSE 7
parentheses. Voucher specimens are deposited at
LAH (sensu Thiers continuously updated).
The macroscopic description of Crepidotus mal-
achioides is based on observations of fresh material.
Photographs were taken with a Canon EOS 80D
camera (Tokyo, Japan). The micro-morphological
characters are based on the study of both fresh and
dried material. Dry specimens were rehydrated in
distilled water or 10 % KOH before observation and
mounted in aqueous Congo Red. In the notation of
basidiospore dimensions, (a–)b–c(–d), ‘b–c’ is the
range including 90 % of the measured values, with
‘a’ and ‘d’ being extreme values. Q represents the
range of the length/width ratio for all measured
spores. For all microscopic structures, 20 elements
were measured. Voucher specimens are deposited at
GDOR (sensu Thiers continuously updated).
For the Herpomyces Thaxt. study, Periplaneta
americana cockroaches (Blattodea, Blattidae) were
collected by hand and screened for the presence of
Herpomyces thalli (Ascomycota, Laboulbeniomy-
cetes) under 40–50× magnication. Antennae were
dislodged from the host and embedded in cotton
blue, resulting in clearly colored thalli. Next, thalli
were removed from the antenna using a stainless
steel No. 000 insect pin (Ento Sphinx, Pardubice,
Czech Republic). Thalli were embedded in Amann
solution (Benjamin 1971) with the help of a droplet
of Hoyer’s medium as described by De Kesel et al.
(2020). Permanent slides are deposited at BR.
Mounted thalli were viewed at 200–1,000× magni-
cation using an Olympus BX51 light microscope
(Tokyo, Japan) with drawing tube, digital camera,
and AnalySIS software (Soft Imaging System
GmbH, Münster, Germany).
For the Leucoagaricus Locq. ex Singer study,
macromorphological features were studied on fresh
collections as well as by analyzing photos taken in
the eld. Micromorphological data were obtained
from dried material and observed under a Zeiss
Axio Imager.A2 light microscope, equipped with
AxioVision Release 4.8.2. software. Measurements
were done with a 100× oil immersion objective
(1,000× magnication). Dried basidiomata were
sectioned with a razor blade, and the sections were
mounted in water, stained with ammoniacal Congo
red and Melzer’s reagent, separately. The specimens
are deposited at the Department of Plant Anatomy,
Eötvös Loránd University, Budapest, Hungary
(ELTE).
For the Melanustilospora Denchev study, infect-
ed plants of Arum jacquemontii (Alismatales, Are-
caceae) were collected in Kalam, a sub-valley of
Swat (Pakistan), situated at 2,085 m a.s.l. Swat val-
ley is an important phytogeographic region in
northwestern Pakistan with a very rich ora (Ali &
Qaiser 1986, Ahmad et al. 2015). The genus Arum is
native to Europe, Northern Africa, and Central and
Western Asia, with the highest species diversity in
the Mediterranean region (Govaerts et al. 2002). In
Pakistan, only one species is currently known, Arum
jacquemontii, found in northern areas of the coun-
try, including Swat (Stewart 1972). Spores were
studied by light and scanning electron microscopy
(SEM). Spores were observed under an Olympus
BX40 compound microscope in lactic acid. For
SEM, spores were attached to specimen holders us-
ing double-sided adhesive tape and coated with a
50-nm lm of gold in a Polaron E5300 freeze dryer
(Quorum Technologies, Laughton, UK). The gold-
coated stubs were photographed in a 3-30BM scan-
ning electron microscope (CamScan Inc., Cranberry
Township, PA). Measurements of teliospores are
presented as (a–)b–c(–d), with ‘b–c’ representing
90 % of the measured values, and ‘a’ and ‘d’ being
extreme values. Studied materials are deposited at
LAH.
The Neopestalotiopsis Maharachch., K.D. Hyde
& Crous study was conducted in November–Decem-
ber 2017, with the detection of symptoms of fruit
rot on ripened skins of Ber fruit, Zizyphus mauriti-
ana (Rosales, Rhamnaceae). Observations were
made in ve different orchards in Cooch Behar Dis-
trict, West Bengal, India. Diseased tissue was cut
into small pieces and surface-sterilized with 0.1 %
HgCl2 for 30 sec, then plated on potato-dextrose
agar (PDA) and incubated for 10 days at 25 °C and
photoperiod of 12:12 (L:D) h. To conrm patho-
genicity, both mycelial plug inoculation (5 mm
diam.) and conidial suspension injection (1×106 co-
nidia/ml) were performed under laboratory condi-
tion (20 °C, relative humidity 90 %) on detached Ber
fruit; control fruits were treated with PDA plug in-
oculation (5 mm diam.) and sterile water injection.
Diseased plant material and cultures are preserved
at the National Herbarium of Cultivated Plants, In-
dian Agricultural Research Institute, and at the In-
dian Type Culture Collection (New Delhi, India).
For the study of Phytopythium Abad, De Cock,
Bala, Robideau, A.M. Lodhi & Lévesque in Brazil,
samples of water, soil, submerged leaves, and sur-
face sediments were collected in several areas of the
Atlantic Rainforest. Soil, submerged leaves, and
surface sediments were diluted in 30 ml of sterile
ddH2O and, along with water samples collected
from freshwater bodies and rainbow trout farming
tanks, plated and baited using the multiple baiting
method (Milanez 1989). Plates were incubated for
274 Sydowia 73 (2021)
Dima et al.: FUSE 7
4−7 days in an acclimatized room at ~22 ºC. After
this period, the baits (Sorghum spp. seeds, onion-
skin, corn leaves, and snakeskin) were observed un-
der a Leica DMLB2 compound microscope. Phyto-
pythium isolates were puried onto cornmeal agar
(CMA) medium (Fuller & Jaworski 1987), with
0.5g/l streptomycin sulphate and 0.5g/l penicillin
G. Species were identied morphologically based
on the original descriptions using a Leica MC170
HD camera with Leica Qwin 3.1 software. Speci-
mens were incorporated into the CCIBt culture col-
lection (Coleção de Culturas de Algas, Cianobacté-
rias e Fungos do Instituto de Botânica).
Collections of Pluteus Fr. were collected in the
province of Mug˘la, Turkey in 2015. Morphological
features were described from young to mature ba-
sidiomata. The macro-morphological descriptions
and images of basidiomata were based on fresh ma-
terial. For micro-morphological structures, dried
basidiomata were rehydrated in 3 % KOH, and sub-
sequently stained with Congo Red. The following
abbreviations are used in the descriptions: Lm for
the average length of all measured basidiospores,
Wm for the average width, Q for length/width ratio,
and Qm for the average of all calculated Q values. At
least thirty basidiospores from spore prints were
measured. Microstructures were drawn with the
help of Adobe Illustrator CS4 (San Jose, CA). Speci-
mens are deposited in the personal fungarium of O.
Kaygusuz at Isparta University of Applied Sciences
(Turkey).
Collections of Rhodocollybia Singer were made
in the framework of the Norwegian Barcode of Life
project (NorBOL). Macromorphological observa-
tions are based on eld notes, fresh or dried basidi-
omata were photographed by NorBOL staff in the
laboratory as part of the standard NorBOL docu-
mentation method under a Creative Common Attri-
bution NonCommercial ShareAlike (CC BY-SA) li-
cense. Specimens were dried in a dehydrator at
45 °C and deposited at O (sensu Thiers continuously
updated).
DNA extraction, PCR amplication, and sequenc-
ing
For the Cortinarius study, total DNA was ex-
tracted from dried specimens following Murray &
Thompson (1980). Primers ITS1F and ITS4 (White
et al. 1990, Gardes & Bruns 1993) were used to am-
plify the internal transcribed spacer (ITS) barcode
region, and LR0R and LR5 (Vilgalys & Hester 1990,
Hopple 1994) were used for the large subunit (LSU)
ribosomal RNA gene. Cycling conditions were as
follows: initial denaturation at 95 ºC for 5 min; fol-
lowed by 35 cycles of denaturation at 94 ºC for 45 s,
annealing at 54 ºC for 30 s, and extension at 72 ºC
for 45 s; followed by a nal extension at 72 ºC for
10 min (Mullis & Faloona 1987). PCR products were
checked in 1% agarose gels, and positive reactions
were sequenced with one or both PCR primers.
Chromatograms were checked to search for puta-
tive reading errors, which were corrected manually.
For the Entoloma study, DNA was extracted
from dried herbarium material using the Nucle-
oSpin Plant II kit (Macherey-Nagel, Germany). For-
ward primers ITS1F and reverse primers ITS4 and
ITS4B (White et al. 1990, Gardes & Bruns 1993)
were used in both PCR and sequencing reactions for
the ITS barcode region. PCR products were puried
with the Fermentas Genomic DNA Purication Kit
(Thermo Scientic, Pittsburg, PA). Puried PCR
products were sequenced on an ABI model 3130 Ge-
netic Analyzer (Applied Biosystems, Foster City,
CA) or commercially at LGC Genomics (Berlin,
Germany). Alternatively, DNA extraction, PCR, and
sequencing procedures were performed through the
Norwegian Barcode of Life project (NorBOL) and
followed Larsson & Jacobsson (2004) and Larsson
et al. (2018). Chromatograms were checked and ed-
ited with the CodonCode Aligner package (Codon-
Code Corporation, Centerville, MA) and MEGA X
(Kumar et al. 2018). Sequence comparison with
public and personal databases followed Noordeloos
et al. (2017). Newly generated sequences were sub-
mitted to GenBank (Tab. 1).
Total DNA of Suillus quercinus sp. nov. was ex-
tracted from dried material following a modied
CTAB extraction method (Lee et al. 1988). The ex-
tracted genomic DNA was evaluated using gel elec-
trophoresis with 1 % agarose gels and a UVIpro
Platinum gel documentation system (Uvitec, Cam-
bridge, UK) with default settings. Genomic DNA
was suspended in nuclease-free H20 and stored at
20 ºC. The ITS region was amplied using primers
ITS1F and ITS4 (White et al. 1990, Gardes & Bruns
1993). Cycling conditions included an initial dena-
turation step at 94 °C for 1 min; then 35 cycles of
denaturation at 94 °C for 1 min, annealing at 53 °C
for 1 min, and extension at 72 °C for 1 min; and nal
extension at 72 °C for 8 min (Saba et al. 2020). Puri-
cation and sequencing were outsourced to Macro-
gen (Seoul, South Korea). Consensus sequences
were generated from the obtained sequence reads in
BioEdit version 7.0.9 (Hall 1999). Newly generated
sequences were submitted to NCBI GenBank with
accession numbers MT361744, MT361745, and
MT361746.
Sydowia 73 (2021) 275
Dima et al.: FUSE 7
Tab. 1. Details of sequences and isolates included in the BLAST searches and molecular analyses for the new species and interesting reports.
Species name ID (isolate, strain1, status2,
voucher) Country, isolation source SSU ITS LSU Reference(s)
Agrocybe pediades CIGYA 003 Mexico MG817376 A.M. Montiel Gonzalez & R. Ortega
Avila, unpubl.
Collybia butyracea olrim421 Sweden AY781251 Mata et al. (2004)
Collybia butyracea olrim426 Sweden AY805607 Mata et al. (2004)
Cortinarius aavae VMS8 Canada FJ717539 Harrower et al. (2011)
Cortinarius aciculisporus G: 257 France MT934844 Liimatainen et al. (2020)
Cortinarius albolens PC:A. Bidaud 97-10-368 France MT934855 Liimatainen et al. (2020)
Cortinarius aureifer G:351 France MT934893 Liimatainen et al. (2020)
Cortinarius badioavidus WTU:J.F. Ammirati 13668, T USA NR_153055 Li et al. (2016)
Cortinarius badioavidus WTU:M. Beug 11MWB111913 USA KU041733 Li et al. (2016)
Cortinarius buxiolens G:3300 France MT934935 Liimatainen et al. (2020)
Cortinarius carcharias G:4276 France MT934948 Liimatainen et al. 2020
Cortinarius conicoumbonatus KATO 3455 Turkey MF696141 Sesli & Liimatainen (2018)
Cortinarius ferrugineifolius IB:M. Moser 1964-0110 Sweden MT935277 Liimatainen et al. (2020)
Cortinarius ferrugineifolius IB:M. Moser 1991-0305 USA MT935044 Liimatainen et al. (2020)
Cortinarius ferrugineovelatus UBC:F17141 Canada GQ159884 Harrower et al. (2011)
Cortinarius fulvoisabellinus JB-8342/14 Spain KU953939 Ballarà et al. (2016)
Cortinarius fulvoisabellinus PC:RH1891 France KJ206485 Dima et al. (2014)
Cortinarius griseascens G:4263 France MT935100 Liimatainen et al. (2020)
Cortinarius helvolus TUB 011905 France AY669667 Garnica et al. (2005)
Cortinarius herculinus G:4195 France MT935120 Liimatainen et al. (2020)
Cortinarius hinnuleoarmillatus G: 00052098, T France NR_131790 Niskanen et al. (2006)
Cortinarius hinnuleoarmillatus G:GK16160 France DQ499464 Niskanen et al. (2006)
Cortinarius hinnuleoarmillatus H:IK01-021 Finland DQ499462 Niskanen et al. (2006)
Cortinarius hinnuleoarmillatus H:TN03-093 Finland DQ499460 Niskanen et al. (2006)
Cortinarius hinnuleoarmillatus S:F39953 Sweden DQ499461 Niskanen et al. (2006)
Cortinarius hinnuleocervinus Ammirati 13502 USA MT935133 Liimatainen et al. (2020)
Cortinarius hinnuleocervinus Niskanen 12-175, T USA MG136827 Liimatainen (2017)
Cortinarius hinnuleovelatus G:4203 France MT935136 Liimatainen et al. (2020)
Cortinarius hinnuleus CFP332 Sweden DQ117926 Kytövuori et al. (2005)
Cortinarius hinnuleus IB19960139 India AY083183 Peintner et al. (2003)
Cortinarius cf. hinnuleus var. minutalis 2071876 Canada MG786239 K. Liimatainen & J. Ammirati, unpubl.
Cortinarius hinnuloides G:4285 France MT935139 Liimatainen et al. (2020)
Cortinarius lepidus G:765 France MT935194 Liimatainen et al. (2020)
Cortinarius nauseosmus G:4214 France MT935240 Liimatainen et al. (2020)
Cortinarius ochraceoplicatus G:378 France MT935256 Liimatainen et al. (2020)
Cortinarius paraphaeochrous IB:M. Moser 1991-0323 USA MT935282 Liimatainen et al. (2020)
Cortinarius pseudohinnuleus G:4224 France MT935343 Liimatainen et al. (2020)
Cortinarius radicosissimus G:142 France MT935369 Liimatainen et al. (2020)
Cortinarius roseonudipes G:37 France MT935391 Liimatainen et al. (2020)
Cortinarius salicum G:4479 France MT935414 Liimatainen et al. (2020)
Cortinarius scabridipileus H:7000550, T Sweden MT112168 Niskanen (2020)
Cortinarius semiodoratus PC:R. Henry 91.6 France MT935427 Liimatainen et al. (2020)
Cortinarius solidus G:4253 France MT935438 Liimatainen et al. (2020)
Cortinarius sp. 1745193 Spain MG739379 B. McAdoo, unpubl.
Cortinarius speciosior G:4280 France MT935443 Liimatainen et al. (2020)
276 Sydowia 73 (2021)
Dima et al.: FUSE 7
Species name ID (isolate, strain1, status2,
voucher) Country, isolation source SSU ITS LSU Reference(s)
Cortinarius squamosus KATO Fungi 3386, T Turkey MW314263 MW314262 This study
Cortinarius squamulifer G:4260 France MT935451 Liimatainen et al. (2020)
Cortinarius sublamentosus G:1196 France MT935484 Liimatainen et al. (2020)
Cortinarius subhelvolus G:470 France MT935491 Liimatainen et al. (2020)
Cortinarius subrigidipes IB:M. Moser 1991-0309 USA MT935523 Liimatainen et al. (2020)
Cortinarius subulatus G:4229 France MT935536 Liimatainen et al. (2020)
Cortinarius tigris G:4269 France MT935553 Liimatainen et al. (2020)
Cortinarius umbrinolens TUB 011918 Germany AY669658 Garnica et al. (2005)
Crepidotus albolanatus PDD:72865 holotype New Zealand KY827292 Horak (2018)
Crepidotus luteolus 16834 Italy JF907963 Osmundson et al. (2013)
Crepidotus malachioides 303022 USA MK607561 S.D. Russell & D. Grootmyers, unpubl.
Crepidotus malachioides BD46 China JQ666669 Zhiguang et al. (2016)
Crepidotus malachioides FLAS-F-61634 USA MH212015 B.S. Kaminsky, M.E. Smith, R. Healy &
B. Spakes Richter, unpubl.
Crepidotus malachioides GDOR5069 Italy MW504470 This study
Crepidotus malachioides SLO 1250, T Slovakia NR_132047 Jancˇovicˇová et al. (2014)
Crepidotus malachioides WU 31421 Austria KF879616 Jancˇovicˇová et al. (2014)
Crepidotus malachioides WU 32709 Austria KF879615 Jancˇovicˇová et al. (2014)
Crepidotus cf. malachioides TENN 055381 Argentina KY559326 P.B. Matheny, H.B. Korotkin &
M.E. Smith, unpubl.
Crepidotus nyssicola S.D. Russell MycoMap # 7399 USA MN906236 S.D. Russell, unpubl.
Crepidotus nyssicola S.D. Russell MycoMap # 7426 USA MN906237 S.D. Russell, unpubl.
Crepidotus sp. MushroomObserver.org/307011 USA MH087459 A. Rockefeller, unpubl.
Crepidotus sp. SDR NAMA 2017-156 USA MK575449 A. Rockefeller, unpubl.
Crepidotus sphaerosporus 11253 Italy JF907960 Osmundson et al. (2013)
Crepidotus sphaerosporus HMAS 255466 MK966515 T.Z. Wei, unpubl.
Crepidotus sphaerosporus HMAS 290002 MK966514 T.Z. Wei, unpubl.
Crepidotus subverrucisporus 15720 Italy JF907961 Osmundson et al. (2013)
Entoloma anatinum [as Entoloma
longistriatum]
TUF106928 Estonia UDB015648* I. Saar, unpubl.
Entoloma asprellum [as Entoloma
lividocyanulum]
TUF106064 Estonia UDB011486* I. Saar, unpubl.
Entoloma atrocoeruleum [as Entoloma
poliopus va r. parvisporigerum]
TUF120520 Estonia UDB031517* UNITE
Entoloma azureosquamulosum HKAS53408 China JQ410334 He et al. (2012)
Entoloma brunneicoeruleum AKS-E1-15 Norway MZ145169 This study
Entoloma brunneicoeruleum JL124-16 Norway MZ145168 This study
Entoloma brunneicoeruleum L0608198 The Netherlands MZ145166 This study
Entoloma brunneicoeruleum LE 302098, T Russia MZ145170 This study
Entoloma brunneicoeruleum O-F-291139 Norway MZ145167 This study
Entoloma aff. caesiellum SAAS1410 China KP329587 X.-L. He, unpubl.
Entoloma callipygmaeum LE 253784, T Russia MZ145207 This study
Entoloma callipygmaeum LE 312487 Russia MZ145206 This study
Entoloma callipygmaeum LE 312488 Russia MZ145205 This study
Entoloma cf. catalaunicum TU106338 Estonia UDB011680* I. Saar, unpubl.
Entoloma chalybaeum TUF105760 Denmark UDB034191* I. Saar, unpubl.
Entoloma ekaterinae LE312053, T Russia MK693215 Crous et al. (2019)
Sydowia 73 (2021) 277
Dima et al.: FUSE 7
Species name ID (isolate, strain1, status2,
voucher) Country, isolation source SSU ITS LSU Reference(s)
Entoloma erhardii LE312051, T Russia MK693218 Crous et al. (2019)
Entoloma exile K(M)157760 Great Britain MF977951 J.J. Elsey, unpubl.
Entoloma cf. foliocontusum
[as Leptonia cf. foliocontusa]
4954SL USA KX574457 M. Gordon, unpubl.
Entoloma fuscosquamosum TENN064384 USA KY777405 P.B. Matheny, S.A. Trudell &
M.G. Wood, unpubl.
Entoloma holmvassdalenense O-F-75311, T Norway KM610321 Weholt et al. (2014)
Entoloma incanum K(M)190322 Great Britain MF977955 J.J. Elsey, unpubl.
Entoloma kauffmanii KA13-1202 South Korea KR673675 Kim et al. (2015)
Entoloma cf. largentii [as Leptonia
cf. convexa]
OSC144006 USA KX574458 M. Gordon, unpubl.
Entoloma longistriatum TENN070451 USA KY744164 P.B. Matheny, S.A. Trudell &
M.G. Wood, unpubl.
Entoloma minutigranulosum GB-0204540 Sweden MZ145209 This study
Entoloma minutigranulosum JL54-14 Norway MZ145221 This study
Entoloma minutigranulosum JL69-17 Norway MZ145220 This study
Entoloma minutigranulosum JL99-11 Norway MZ145216 This study
Entoloma minutigranulosum JL99-16 Norway MZ145217 This study
Entoloma minutigranulosum L0607681 The Netherlands MZ145215 This study
Entoloma minutigranulosum L0607941 / MD-2014-7 Germany MZ145211 This study
Entoloma minutigranulosum LE 302096, T Russia MZ145214 This study
Entoloma minutigranulosum LE 312483 Russia MZ145212 This study
Entoloma minutigranulosum LE 312484 Russia MZ145210 This study
Entoloma minutigranulosum LE 312675 Russia MZ145219 This study
Entoloma minutigranulosum O-F-175913 / GG 3568 Norway MZ145218 This study
Entoloma minutigranulosum O-F-304886 / TEB 121-16 Norway MZ145208 This study
Entoloma minutigranulosum WU25057 Austria MZ145213 This study
Entoloma montanum GB-0191635, T Sweden MW340896 Noordeloos et al. (2021)
Entoloma mougeotii LE254352 Russia KC898446 Morozova et al. (2014)
Entoloma nigrovelutinum LE295077, T Vietnam MF898426 Crous et al. (2017)
Entoloma nipponicum TNS F70747, T Japan MK693223 Crous et al. (2019)
Entoloma nordlandicum O-F-76176, T Norway MW340899 Noordeloos et al. (2021)
Entoloma ochromicaceum TU120040 Estonia UDB023715* I. Saar, unpubl.
Entoloma perasprellum GB-0204546 / EL28-19 Sweden MZ145182 This study
Entoloma perasprellum GB-0204547 / JBJ 19-107 Sweden MZ145179 This study
Entoloma perasprellum GB-0204548 / JBJ 19-122 Sweden MZ145180 This study
Entoloma perasprellum GB-0204549 / JBJ 19-180 Sweden MZ145183 This study
Entoloma perasprellum GB-0204550 / JBJ 19-119 Sweden MZ145181 This study
Entoloma perasprellum GC01100310, T France MZ145177 This study
Entoloma perasprellum LE 312499 Russia MZ145186 This study
Entoloma perasprellum LE 312500 Russia MZ145185 This study
Entoloma perasprellum LE 312501 Russia MZ145178 This study
Entoloma perasprellum O-F-256732 / JBJ 18-004 Sweden MZ145184 This study
Entoloma poliopus G4742 Estonia
UDB0332859*
R. Pau, unpubl.
Entoloma pulchripes LE 311808, T Russia MZ145188 This study
Entoloma pulchripes LE 311809 Russia MZ145189 This study
Entoloma pulchripes LE 312485 Russia MZ145187 This study
278 Sydowia 73 (2021)
Dima et al.: FUSE 7
Species name ID (isolate, strain1, status2,
voucher) Country, isolation source SSU ITS LSU Reference(s)
Entoloma querquedula 18.XI.2011 TUR Finland LN850627 Kokkonen (2015)
Entoloma aff. rectangulum [as Leptonia
rectangula]
SCL8524 USA KU574744 M. Gordon, unpubl.
Entoloma roseotinctum WU13070 Austria LN850611 Kokkonen (2015)
Entoloma sarcitulum TUR-31-VII-1967 FIN LN850562 Kokkonen (2015)
Entoloma septentrionale O-F-254295, T Norway MW340904 This study
Entoloma serrulatum LE254361 Russia KC898447 Morozova et al. (2014)
Entoloma sp. CM13-233 New Caledonia KY774214 Carriconde et al. (2019)
Entoloma sp. MEL2382758 Australia KP012941 G. Bonito, M. Barrett, F. Udovicic & T.
Lebel, unpubl.
Entoloma sp. MES-2627 Chile MH930251 A.B. Mujic & M.E. Smith, unpubl.
Entoloma sp. MES-534 Chile KY462681 Truong et al. (2017)
Entoloma sp. MushroomObserver.org/240552 USA MG966312 A. Rockefeller, unpubl.
Entoloma sp. soil sample USA EF619690 Parrent & Vilgalys (2007)
Entoloma sp. soil sample USA KP889939
S.H.A. Guichon & S.W. Simard, unpubl.
Entoloma sp. TRTC156923 Canada JN021015 Dentinger et al. (2011)
Entoloma sp. TUF120259 Estonia UDB024650* I. Saar, unpubl.
Entoloma sp. [as Entoloma cf. sericeum] UBC-F14053 Canada AF335439 M.L. Berbee, P. Inderbitzin &
G. Zhang, unpubl.
Entoloma subcaesiellum LE253776, T Russia MK693224 Noordeloos & Morozova (2010)
Entoloma subcaesiocinctum SAAS2238 China KY711234 He et al. (2017)
Entoloma subcorvinum TENN070435 USA KY744169 P.B. Matheny, S.A. Trudell &
M.G. Wood, unpubl.
Entoloma subfarinaceum TENN070395 USA KY777374 P.B. Matheny, S.A. Trudell &
M.G. Wood, unpubl.
Entoloma tigrinum GB-0204535 / EL156-18 Sweden MZ145172 This study
Entoloma tigrinum GB-0204536 / JBJ 19-128 Sweden MZ145171 This study
Entoloma tigrinum GB-0204537 / JBJ 19-109 Sweden MZ145173 This study
Entoloma tigrinum GB-0204538 Sweden MZ145175 This study
Entoloma tigrinum GB-0204539 Sweden MZ145174 This study
Entoloma tigrinum O-F-304580, T Norway MZ145176 This study
Entoloma timidum JL13-16 Norway MZ145195 This study
Entoloma timidum JL15-14 Norway MZ145194 This study
Entoloma timidum LE 311800 Russia MZ145191 This study
Entoloma timidum LE 312382 Russia MZ145198 This study
Entoloma timidum LE 312480, T Russia MZ145197 This study
Entoloma timidum LE 312481 Russia MZ145199 This study
Entoloma timidum O-F-252355 Norway MZ145196 This study
Entoloma timidum O-F-75148 Norway MZ145190 This study
Entoloma timidum TEB73-20 Norway MZ145193 This study
Entoloma timidum Weholt E16-10 Norway MZ145192 This study
Entoloma turci MCVE3882 Italy JF907993 Osmundson et al. (2013)
Entoloma cf. unicolor TENN070383 USA KY777373 P.B. Matheny, S.A. Trudell &
M.G. Wood, unpubl.
Entoloma violaceoserrulatum JV 8329F (TUR), T Finland MF476913 Morozova et al. (2018)
Entoloma violaceoserrulatum L0607704 The Netherlands MZ145200 This study
Sydowia 73 (2021) 279
Dima et al.: FUSE 7
Species name ID (isolate, strain1, status2,
voucher) Country, isolation source SSU ITS LSU Reference(s)
Entoloma violaceoserrulatum LE 312676 Sweden MZ145204 This study
Entoloma violaceoserrulatum O-F-260353 / TEB 339-15 Norway MZ145201 This study
Entoloma violaceoserrulatum O-F-75151 Norway MZ145203 This study
Entoloma violaceoserrulatum TEB128-19 Norway MZ145202 This study
Entoloma yanacolor QCAM 6312, T Ecuador MG947210 Crous et al. (2018)
Herpomyces chaetophilus D. Haelew. 1097b Panama, Periplaneta
americana
MG438321 MG438294 MG438352 Haelewaters et al. (2019)
Herpomyces chaetophilus D. Haelew. 1097c
Panama, Periplaneta americana
MG438322 MG438295 MG438353 Haelewaters et al. (2019)
Herpomyces chaetophilus D. Haelew. 435b USA Massachusetts,
Periplaneta americana
MG438318 MG438292 Haelewaters et al. (2019)
Herpomyces chaetophilus D. Haelew. 483b USA Massachusetts,
Periplaneta americana
MG438319 MG438293 MG438350 Haelewaters et al. (2019)
Herpomyces chaetophilus D. Haelew. 483e USA Massachusetts,
Periplaneta americana
MG438320 MG438351 Haelewaters et al. (2019)
Herpomyces chaetophilus D. Haelew. 602b USA Massachusetts, Periplan-
eta americana
KT800023 KT800039 KT800009 Haelewaters et al. (2015)
Herpomyces ectobiae MG001 Poland, Blattella germanica KT800024 KT800040 Haelewaters et al. (2015)
Herpomyces ectobiae TW793a USA California, Blattella
germanica
MG438296 Haelewaters et al. (2019)
Herpomyces leurolestis 2017/0199 Hungary, Phoetalia pallida MG438299 Haelewaters et al. (2019)
Herpomyces leurolestis D. Haelew. 1417b Hungary, Phoetalia pallida MG438297 Haelewaters et al. (2019)
Herpomyces leurolestis Debr_Ppal Hungary, Phoetalia pallida MG438323 MG438298 MG438354 Haelewaters et al. (2019)
Herpomyces paranensis D. Haelew. 1365a
Panama, Blaberus giganteus
MG438300 Haelewaters et al. (2019)
Herpomyces paranensis D. Haelew. 1365b
Panama, Blaberus giganteus
MG438301 Haelewaters et al. (2019)
Herpomyces periplanetae ADK6485 Bénin, Periplaneta americana MZ145233 MZ145234 MZ144000 This study
Herpomyces periplanetae D. Haelew. 1187d USA Massachusetts,
Periplaneta americana
MG438331 MG438309 MG438359 Haelewaters et al. (2019)
Herpomyces periplanetae D. Haelew. 602a USA Massachusetts,
Periplaneta americana
MG438326 MG438304 Haelewaters et al. (2019)
Herpomyces periplanetae D. Haelew. 602c USA Massachusetts,
Periplaneta americana
KT800025 KT800041 KT800010 Haelewaters et al. (2015)
Herpomyces periplanetae D. Haelew. 602d USA Massachusetts,
Periplaneta americana
MG438327 MG438305 MG438357 Haelewaters et al. (2019)
Herpomyces periplanetae D. Haelew. 620a USA New York, Periplaneta
americana
MG438328 MG438306 MG438358 Haelewaters et al. (2019)
Herpomyces periplanetae D. Haelew. 654b Panama, Periplaneta
americana
MG438329 MG438307 Haelewaters et al. (2019)
Herpomyces periplanetae D. Haelew. 654c Panama, Periplaneta
americana
MG438330 MG438308 MG438308 Haelewaters et al. (2019)
Herpomyces periplanetae TW437c USA Massachusetts,
Periplaneta americana
MG438324 MG438302 MG438355 Haelewaters et al. (2019)
Herpomyces periplanetae TW448b USA Massachusetts,
Periplaneta americana
MG438325 MG438303 MG438356 Haelewaters et al. (2019)
Herpomyces shelfordellae Bud_Slat Hungary, Shelfordella lateralis MG438333 MG438312 MG438361 Haelewaters et al. (2019)
Herpomyces shelfordellae D. Haelew. 1415a Hungary, Shelfordella lateralis MG438313 Haelewaters et al. (2019)
Herpomyces shelfordellae D. Haelew. 1427a USA Massachusetts,
Shelfordella lateralis
MK299847 Haelewaters et al. (2019)
280 Sydowia 73 (2021)
Dima et al.: FUSE 7
Species name ID (isolate, strain1, status2,
voucher) Country, isolation source SSU ITS LSU Reference(s)
Herpomyces shelfordellae DE_HerpBL1 Hungary, Shelfordella lateralis KT800026 KT800042 KT800011 Haelewaters et al. (2015)
Herpomyces shelfordellae MH202b, T Poland, Shelfordella lateralis MK299848 Haelewaters et al. (2019)
Herpomyces sp. H77-1 USA North Carolina,
Parcoblatta cf. lata
KY523239 KY293260 KY350529 Sundberg et al. (2018)
Herpomyces spegazzinii CEPHe27-1 Argentina, Periplaneta
fuliginosa
MN597427 Gutierrez et al. (2020)
Herpomyces spegazzinii LPS:49123-2, T Argentina, Periplaneta
fuliginosa
NR_169702 Gutierrez et al. (2020)
Herpomyces stylopygae Bud_Bori Hungary, Blatta orientalis MG438332 MG438310 MG438360 Haelewaters et al. (2019)
Herpomyces stylopygae Bud_Bori_2 Hungary, Blatta orientalis MG438311 Haelewaters et al. (2019)
Leucoagaricus americanus BHI-F360a USA Massachusetts MF161229 Haelewaters et al. (2018)
Leucoagaricus americanus S.D. Russell HRL0779 Canada MH979431 S. Russell & R. Lebeuf, unpubl.
Leucoagaricus americanus S.D. Russell MO #107715 USA Indiana MW567919 S.D. Russell, unpubl.
Leucoagaricus americanus S.J.W. Verduin (L) USA North Carolina AF295928 Vellinga (2000)
Leucoagaricus americanus
[as Leucoagaricus bresadolae]
CCBAS802 Czech Republic LN714565 Veˇtrovský et al. (2016)
Leucoagaricus americanus
[as Leucoagaricus bresadolae]
MCVE:756 Italy GQ329047 Osmundson et al. (2013)
Leucoagaricus americanus
[as Leucoagaricus sp.]
Royan 3 Iran MT573394 Alimadadi et al. (2019)
Leucoagaricus cf. majusculus MFLU 09-0164 Thailand HM488764 Vellinga et al. (2011)
Leucoagaricus gongylophorus MyrE002 Panama KF572016 Kooij et al. (2015)
Leucoagaricus meleagris AJ 542 USA Massachusetts MN483027 Justo et al. (2021)
Leucoagaricus meleagris E.C. Vellinga 1990 The Netherlands AY176419 Vellinga (2004)
Leucoagaricus meleagris E.C. Vellinga 2095 The Netherlands AF482867 Vellinga et al. (2003)
Leucoagaricus meleagris HAW:JKS90 USA Hawai’i MK412590 Stallman (2019)
Leucoagaricus meleagris iNAT:58855005 USA New York MW031133 S. Jakob, unpubl.
Leucoagaricus meleagris WA0000072219 Laos MT252565 Łuczaj et al. (2021)
Leucoagaricus mucrocystis AJ 476 USA US Virgin Islands MN483025 Justo et al. (2021)
Leucoagaricus mucrocystis CA 16 (JBSD) Dominican Republic MN483026 Justo et al. (2021)
Leucoagaricus mucrocystis ELTE:DB-FG-167-19 French Guiana MZ047586 MZ047585 This study
Leucoagaricus mucrocystis ELTE:DB-FG-168-19 French Guiana MZ047587 This study
Leucoagaricus mucrocystis
[as Lepiota besseyi]
D.E. Hemmes deh1867 USA Hawai’i HM488763 Vellinga et al. (2011)
Leucoagaricus mucrocystis
[as Lepiota besseyi]
HAW:JKS109 USA Hawai’i MK412602 Stallman (2019)
Leucocoprinus sp. RB29 Brazil MN473907 R. Bizarria Jr. & A. Rodrigues, unpubl.
Neopestalotiopsis clavispora ncb01 India, Ziziphus mauritiana MW012901 This study
Phytopythium boreale CBS:551.88 China AB725879 HQ665261 Robideau et al. (2011), Baten et al.
(2014)
Phytopythium carbonicum CBS:112544 France AB725876 AB996605 Baten et al. (2014)
Phytopythium carbonicum CBS:292.37 The Netherlands AB690620 M.A. Baten, K. Kageyama &
H. Suga, unpubl.
Phytopythium chamaehyphon CBS:259.30 USA Hawai’i AB690609 HQ665177 Robideau et al. (2011), Baten et al.
(2014)
Phytopythium chamaehyphon CCIBt 4338 Brazil MT620968 MT612336 This study
Phytopythium chamaehyphon CCIBt 4407 Brazil MT620969 MT612337 This study
Sydowia 73 (2021) 281
Dima et al.: FUSE 7
Species name ID (isolate, strain1, status2,
voucher) Country, isolation source SSU ITS LSU Reference(s)
Phytopythium chamaehyphon CPZ68 Brazil MT620967 MT612335 This study
Phytopythium chamaehyphon PPRI8625 South Africa FJ415975 McLeod et al. (2009)
Phytopythium citrinum CBS:119171 France AY197328 AB948195 Baten et al. (2014)
Phytopythium curcubitacearum CBS:748.96 Japan AB725877 AB690598 Baten et al. (2014)
Phytopythium delawarense 382B USA Ohio AB725875 AB690591 Baten et al. (2014)
Phytopythium dogmae USTCMS 4101 Philippines MF353170 MF373431 Bennett et al. (2017)
Phytopythium fagopyri CBS:293.35 Japan AB690617 AB690590 Baten et al. (2014)
Phytopythium fagopyri FP1 Japan AB690621 AB690599 Baten et al. (2014)
Phytopythium fagopyri HonMa Japan AB690615 AB690588 Baten et al. (2014)
Phytopythium helicoides CBS:286.31 USA AB725878 MF375637 Baten et al. (2014), Bennett et al. (2017)
Phytopythium helicoides CCIBt 4103 Brazil KR092138 This study
Phytopythium helicoides CCIBt 4104 Brazil KR092137 This study
Phytopythium iriomotense GUCC0025 AB690622 AB690600 M.A. Baten, K. Kageyama & H. Suga,
unpubl.
Phytopythium iriomotense GUCC7020 AB690629 AB690607 M.A. Baten, K. Kageyama & H. Suga,
unpubl.
Phytopythium kandeliae CBS:111.91 Taiwan HQ643134 HQ665065 Robideau et al. (2011)
Phytopythium kandeliae CCIBt 4023 Brazil KJ399962 KJ399965 Marano et al. (2014)
Phytopythium leanoi CBS:113.91 Taiwan MF355474 KJ399963 Marano et al. (2014), Bennett et al.
(2017)
Phytopythium leanoi USTCMS 4102 Philippines MF353169 MF373430 Bennett et al. (2017)
Phytopythium litorale CBS:118360 Germany HQ643386 HQ665082 Robideau et al. (2011)
Phytopythium litorale CCIBt 4659 Brazil MT620970 MT612338 This study
Phytopythium litorale Dyrbkr01 Turkey MN203107 MN197634 Dervis¸ et al. (2020)
Phytopythium litorale Dyrbkr02 Turkey MN203108 MN197635 Dervis¸ et al. (2020)
Phytopythium litorale GUCC7167 AB690612 AB690583 Baten et al. (2014)
Phytopythium litorale SCP82 Brazil MT620972 MT612340 This study
Phytopythium megacarpum CBS:112351 France AB725881 HQ665067 Robideau et al. (2011), Baten et al.
(2014)
Phytopythium mercuriale CBS:112443 South Africa AB725882 KF853236 de Cock et al. (2015); M.A. Baten,
K. Kageyama & H. Suga, unpubl.
Phytopythium mirpurense CBS:124524 Pakistan KJ831614 de Cock et al. (2015)
Phytopythium montanum ADC9762 Germany HQ643391 Robideau et al. (2011)
Phytopythium montanum CBS:111349 Germany AB725883 HQ665064 Robideau et al. (2011), Baten et al.
(2014)
Phytopythium nanginjense Chen216 China MF459636 Chen et al. (2019)
Phytopythium nanginjense Chen218 China MF459635 Chen et al. (2019)
Phytopythium oedochilum CBS:292.37 USA AB690619 HQ665191 Robideau et al. (2011), Baten et al.
(2014)
Phytopythium oedochilum CBS:738.94 South Africa HQ643394 Robideau et al. (2011)
Phytopythium ostracodes CBS:768.73 Spain AB108022 HQ665295 Robideau et al. (2011); M.A. Baten, K.
Kageyama & H. Suga, unpubl.
Phytopythium palingenes CCIBt 3981 Brazil KR092139 KR092143 This study
Phytopythium palingenes CCIBt 4397 Brazil MT620974 MT612342 This study
Phytopythium palingenes CCIBt 4428 Brazil MT620973 MT612341 This study
Phytopythium sindhum CBS:124518 Pakistan HM244825 Bala et al. (2010)
Phytopythium sindhum JKI KJ755089 König et al. (2015)
282 Sydowia 73 (2021)
Dima et al.: FUSE 7
Species name ID (isolate, strain1, status2,
voucher) Country, isolation source SSU ITS LSU Reference(s)
Phytopythium sp. ‘amazonianum’ WPC:8239B1845 Ecuador GU258624 M.D. Coffey, A.K. Brar, E. Xu, E.A.
Sarhan & I.M. Cunningham, unpubl.
Phytopythium sp. ‘amazonianum’ WPC:8243B519 Ecuador GU258937 M.D. Coffey, A.K. Brar, E. Xu, E.A.
Sarhan & I.M. Cunningham, unpubl.
Phytopythium vexans CBS:119.80 Iran AY598713 HQ665090 Lévesque & de Cock (2004), Robideau
et al. (2011)
Phytopythium vexans CCIBt 4101 Brazil KR092141 KR092144 This study
Phytopythium vexans CCIBt 4383 Brazil MT620976 MT612344 This study
Phytopythium vexans CCIBt 4398 Brazil MT620975 MT612343 This study
Pluteus beniensis RSPF 0299 Brazil JQ065029 A. Justo, N. Menolli Jr. & A.M. Minnis,
unpubl.
Pluteus cinereofuscus AJ229 Portugal HM562108 Justo et al. (2011b)
Pluteus aff. cinereofuscus LE 303665 Russia KX216324 Malysheva et al. (2016)
Pluteus eludens MA50497 Portugal HM562118 HM562240 Justo et al. (2011b)
Pluteus eludens SF15 USA HM562185 Justo et al. (2011b)
Pluteus exiguus O-F-21721 Norway UDB036730* A. Molia, unpubl.
Pluteus extremiorientalis LE 303463 Russia KM658282 Crous et al. (2014)
Pluteus extremiorientalis, T LE<RUS>:262872 Russia NR_153249 Crous et al. (2014)
Pluteus occipes BRNM Czech Republic LN794642 Ševcˇíková & Borovicˇka (2015)
Pluteus uminensis SP393710 Brazil FJ816655 Menolli et al. (2010)
Pluteus uminensis SP393711 Brazil FJ816664 FJ816650 Menolli et al. (2010)
Pluteus fuligineovenosus SP393705 Brazil FJ816662 Menolli et al. (2010)
Pluteus hubeiensis HMJAU45199 China MH167350 W. Fengjian & L. Yu, unpubl.
Pluteus hubeiensis HMJAU45200 China MH167353 W. Fengjian & L. Yu, unpubl.
Pluteus jamaicensis SP393706 Brazil FJ816657 Menolli et al. (2010)
Pluteus cf. jamaicensis SP416738 Brazil KM983709 Menolli et al. (2015)
Pluteus cf. jamaicensis SP417454 Brazil KM983711 Menolli et al. (2015)
Pluteus keselakii BRNM 817402 Slovakia MN250223 MN250223 Ševcˇíková et al. (2020)
Pluteus keselakii LIP Mycologie 0401385 France MN250224 Ševcˇíková et al. (2020)
Pluteus keselakii, T BRNM:817402 Slovakia NR_169977 Ševcˇíková et al. (2020)
Pluteus ludwigii MCVE30136 Slovenia MK834525 MK834527 Crous et al. (2019)
Pluteus ludwigii, T MCVE:30136 Slovenia NR_164496 Crous et al. (2019)
Pluteus multiformis AC4249 Spain HM562201 MK278503 Justo et al. (2011b), Varga et al. (2019)
Pluteus multiformis AH 40107 Spain MN250225 MN250225 Ševcˇíková et al. (2020)
Pluteus multiformis OKA-TR1750 Turkey MT982425 MT982429 This study
Pluteus multiformis OKA-TR1751 Turkey MT982426 MT982430 This study
Pluteus multiformis, T AH:40107 Spain NR_119877 Justo et al. (2011a)
Pluteus nanus UC1859980 USA KF306030 N.H. Nguyen, E.C. Vellinga,
G.M. Cobian, A.J. Fernandez &
T.D. Bruns, unpubl.
Pluteus nanus f. griseopus NL-2546 Hungary MK278504 Varga et al. (2019)
Pluteus pallescens AJ214 Spain HM562056 HM562231 Justo et al. (2011b)
Pluteus rimosoafnis SP394379 Brazil HM562145 Menolli et al. (2015)
Pluteus rimosoafnis SP416740 Brazil KM983706 Menolli et al. (2015)
Pluteus sapiicola SP394382 Brazil HM562148 Justo et al. (2011b)
Pluteus sapiicola SP394387 Brazil HM562146 Justo et al. (2011b)
Pluteus sp. JLF1767 USA MK634597 J.L. Frank, unpubl.
Sydowia 73 (2021) 283
Dima et al.: FUSE 7
Species name ID (isolate, strain1, status2,
voucher) Country, isolation source SSU ITS LSU Reference(s)
Pluteus sp. PDD:106511 New Zealand MN738653 J.A. Cooper, unpubl.
Pluteus sp. PDD:110518 New Zealand MN738674 MN738605 J.A. Cooper, unpubl.
Pluteus tenebromarginatus GC17102401 France MT079860 Corriol et al. (2020)
Pluteus terricola PDD:107339 New Zealand MN738665 MN738585 J.A. Cooper, unpubl.
Pythium takayamanum CBS:122.492 Japan HQ643853 HQ665094 Robideau et al. (2011)
Pythium takayamanum CCIBt 4040 Brazil KM058758 KM058755 This study
Pyxidiophora arvernensis CBS 657.82 The Netherlands FJ176839 FJ176894 Schoch et al. (2009)
Pyxidiophora cf. microspora MG200 Poland MG438334 MG438314 MG438362 Haelewaters et al. (2019)
Rhodocollybia asema O-F-248185 Norway MZ156765 This study
Rhodocollybia asema O-F-248288 Norway MZ156764 This study
Rhodocollybia asema O-F-74975 Norway MZ156766 This study
Rhodocollybia asema O-F-75562 Norway MZ156767 This study
Rhodocollybia butyracea FB10726 (TENN) Russia AF505750 Mata et al. (2004)
Rhodocollybia butyracea FB11456 (TENN) Austria AF505751 Mata et al. (2004)
Rhodocollybia butyracea O-F-74979 Norway MZ156768 This study
Rhodocollybia butyracea O-F-75421 Norway MZ156769 This study
Rhodocollybia butyracea TENN53580 Sweden AY313293 Mata et al. (2004)
Rhodocollybia butyracea TU106219, UNITE RefSeq Estonia UDB011434* V. Liiv, unpubl.
Rhodocollybia butyracea TU118269 Estonia UDB015439* V. Liiv, unpubl.
Rhodocollybia butyracea f. asema TU106218 Estonia UDB015078* V. Liiv, unpubl.
Rhodocollybia butyracea f. asema TU106229 Estonia UDB019799* V. Liiv, unpubl.
Rhodocollybia butyracea f. asema TU118574 Estonia UDB017989* V. Liiv, unpubl.
Rhodocollybia fodiens TU106942 Estonia UDB015266* V. Liiv, unpubl.
Rhodocollybia maculata BRNM699408 Italy GU947370 Antonin & Noordeloos (2010)
Rhodocollybia maculata BRNM714632 Czech Republic GU947369 Antonin & Noordeloos (2010)
Rhodocollybia maculata TU106940 Estonia UDB015655* V. Liiv, unpubl.
Rhodocollybia cf. maculata O-F-75733 Norway MZ156770 This study
Rhodocollybia prolixa TU118816 Estonia UDB019494* V. Liiv, unpubl.
Rhodocollybia sp. h43 Czech Republic LN714597 Veˇtrovský et al. (2016)
Suillus americanus 1008-NC USA North Carolina AF166500 Wu et al. (2000)
Suillus americanus F1185163 USA West Virginia KU663182 R. Zhang, X. Shi, P. Liu &
G.M. Mueller, unpubl.
Suillus americanus F1185445 USA Indiana KU663183 R. Zhang, X. Shi, P. Liu &
G.M. Mueller, unpubl.
Suillus americanus MAV-5625 USA Ohio AF166503 Wu et al. (2000)
Suillus americanus TDB-581 USA Michigan L54103 Wu et al. (2000)
Suillus americanus TJB-7683 USA New York AF166502 Wu et al. (2000)
Suillus cf. americanus YNP:2355 USA California KU663196 R. Zhang, X. Shi, P. Liu &
G.M. Mueller, unpubl.
Suillus cothurnatus MA-Fungi 47683 Spain AJ419218 Martin & Raidl (2002)
Suillus cothurnatus MA-Fungi 49403 Spain AJ419217 Martin & Raidl (2002)
Suillus cothurnatus NSW-4662 USA Louisiana L54092 Kretzer et al. (1996)
Suillus decipiens DG-1451 USA Texas AF166508 Wu et al. (2000)
Suillus decipiens DG-1451 USA Texas L54079 Kretzer et al. (1996)
Suillus decipiens DPL-5724 USA Texas AF166510 Wu et al. (2000)
Suillus avidus DG66 UK JQ888208 Jones et al. (2012)
Suillus avidus FFP909 Canada JQ711962 Jones et al. (2012)
284 Sydowia 73 (2021)
Dima et al.: FUSE 7
Species name ID (isolate, strain1, status2,
voucher) Country, isolation source SSU ITS LSU Reference(s)
Suillus avidus FFP962 Canada JQ711908 Jones et al. (2012)
Suillus granulatus K-15-1 Russia MK414509 O.B. Vaishlya, K.S. Karbysheva,
I. Bakhtinskaya & E. Murina, unpubl.
Suillus granulatus KA17-0554 South Korea MN294845 Cho et al. (2020)
Suillus himalayensis SNW-03 India KJ472765 Verma & Reddy (2014a)
Suillus intermedius ACAD-15271 Canada L54074 Kretzer et al. (1996)
Suillus placidus TDB-725 USA Michigan L54108 Kretzer et al. (1996)
Suillus placidus VC-1022 Nepal L54118 Kretzer et al. (1996)
Suillus quercinus LAH240711 Pakistan MT361746 This study
Suillus quercinus LAH36421, T Pakistan MT361745 This study
Suillus quercinus LAH36422 Pakistan MT361744 This study
Suillus sibiricus EA24040 Pakistan JN119750 Sarwar et al. (2011)
Suillus sibiricus EA24104 Pakistan JN119751 Sarwar et al. (2011)
Suillus sibiricus HMAS:66061 China AF166512 Kretzer et al. (1996)
Suillus sibiricus QXW-2092 Austria AF166513 Kretzer et al. (1996)
Suillus sibiricus UC2023481 Switzerland KX213817 Kretzer et al. (1996)
Suillus sibiricus VC-1040 Nepal L54117 Kretzer et al. (1996)
Suillus spraguei GMM-5703 China AF166518 Wu et al. (2000)
Suillus spraguei QXW-2409 China AF166520 Wu et al. (2000)
Suillus spraguei QXW-2435 China AF166522 Wu et al. (2000)
Suillus spraguei Tissue_lib TDB-638 USA Michigan M91617 Wu et al. (2000)
Suillus spraguei TJB-6228 USA New York AF166525 Wu et al. (2000)
Suillus subalutaceus ACAD-15288 Canada L54075 Kretzer et al. (1996)
Suillus subaureus TDB-780 USA Massachusetts L54109 Kretzer et al. (1996)
Suillus subluteus IB-13-8/19/72 USA Michigan L54088 Kretzer et al. (1996)
Suillus triacicularis MSM 0027 Pakistan KM677929 Sarwar et al. (2015)
Suillus triacicularis PUN 5534 India KF977188 Verma & Reddy (2014b)
Suillus triacicularis PUN 5534, T India NR_153233 Verma & Reddy (2014b)
Uncultured Basidiomycota 4S1_A12 USA EU489965 Hollister et al. (2010)
Uncultured Basidiomycota A2 Australia DQ672275 Midgley et al. (2007)
Uncultured fungus FDBC50 Mexico JQ247381 Romero-Olivares et al. (2013)
Uncultured fungus 4248_284 Lithuania MT236490 Mariulynas et al. 2020
Uncultured fungus TUN11 UK KM374523 Johnson et al. (2014)
1 Herbarium abbreviations follow Index Herbariorum (Thiers continuously updated).
2 T: ex-type strain. * from UNITE database (https://unite.ut.ee).
Sydowia 73 (2021) 285
Dima et al.: FUSE 7
DNA was extracted from dried Crepidotus mal-
achioides basidiomata with the CTAB protocol
from Doyle & Doyle (1987). The ITS region was am-
plied with primers ITS1F (Gardes & Bruns 1993)
and ITS4 (White et al. 1990). Cycling conditions
were as in Saba et al. (2020). The generated sequence
reads were assembled and edited in Geneious ver-
sion 11.1.5 (Kearse et al. 2012) and then submitted
to NCBI GenBank. Accession numbers are reported
in Tab. 1.
Genomic DNA was isolated from 1 female thal-
lus of Herpomyces periplanetae using the REPLI-g
Single Cell Kit (Qiagen, Valencia, CA) following the
modications by Haelewaters et al. (2019). Ampli-
cation of the internal transcribed spacer (ITS) bar-
code region and the nuclear ribosomal RNA small
and large subunits (SSU and LSU) was done using
the following primer sets: NSL1/NSL2 for SSU
(Haelewaters et al. 2015), ITS1f/ITS4 for ITS (White
et al. 1990, Gardes & Bruns 1993), and LIC24R/LR3
for LSU (Vilgalys & Hester 1990, Miadlikowska &
Lutzoni 2000). Amplications were performed on a
pro S Mastercycler (Eppendorf, Hauppauge, NY) in
25-μl reactions containing 12.5 μl of 2× MyTaq Mix
(Bioline, Swedesboro, NJ), 9.5 μl of ddH2O, 1.0 μl of
each 10 mM primer, and 1.0 μl of DNA extract. Cy-
cling conditions followed Haelewaters et al. (2019)
for ITS and Liu et al. (2020) for SSU and LSU. For
SSU: initial denaturation at 95 °C for 5 min; then
40 cycles of denaturation at 95 °C for 30 s, anneal-
ing at 55 °C for 45 s, and extension at 72 °C for 45 s;
and nal extension at 72 °C for 1 min. For ITS: ini-
tial denaturation at 94 °C for 3 min; then 35 cycles
of denaturation at 94 °C for 1 min, annealing at
50 °C for 45 s, and extension at 72 °C for 90 s; and
nal extension at 72 °C for 10 min. For LSU: initial
denaturation at 94 °C for 5 min; then 35 cycles of
denaturation at 94 °C for 30 s, annealing at 50 °C for
45 s, and extension at 72 °C for 1 min; and nal ex-
tension at 72 °C for 7 min. Purication and Sanger
sequencing were outsourced to Genewiz (Plaineld,
NJ). Raw sequence reads were assembled, trimmed,
and edited in Sequencher version 5.2.3 (Gene Codes
Corporation, Ann Arbor, MI).
For the Leucoagaricus study, the ITS and LSU
regions were amplied directly from samples with
the Phire Plant Direct PCR Master Mix Kit (Thermo
Scientic). Amplication of the regions ITS and
LSU was done using primer sets ITS1F/ITS4 (White
et al. 1990, Gardes & Bruns 1993) and LR0R/LR5
(Vilgalys & Hester 1990, Hopple 1994), respectively.
Sequencing of the amplicons was carried out with
the primers used for amplication by LGC Genom-
ics (Berlin, Germany). Chromatograms were checked
and edited with CodonCode Aligner version 8.0.1
(CodonCode Corporation). Newly generated se-
quences were submitted to GenBank (Tab. 1).
Neopestalotiopsis DNA was extracted from co-
nidia, conidiophores, and mycelium using the CTAB
method from Doyle & Doyle (1990). DNA was resus-
pended in 100 ml of MilliQ H2O and quantied us-
ing a Nanodrop 2000c spectrophotometer (Thermo
Scientic) at A260/280 and A230/260. DNA was then di-
luted to 20 ng/ml and stored at –20±2 °C. Amplica-
tion of the ITS region was performed with primers
ITS1 and ITS4 (White et al. 1990). PCR was done in
15-μl reactions consisting of 0.18 μl of each 10 mM
primer, 0.18 μl of dNTPs, 0.90 U of GoTaqVR DNA
Polymerase (Promega, Madison, WI), and 3 μl of the
diluted DNA suspension. Amplied PCR products
were checked by electrophoresis on 1.5 % agarose
gel and cleaned with ExoSAP-IT PCR Product
Cleanup Reagent (Thermo Scientic). Sequencing
was done by Macrogen. The generated sequence was
submitted to GenBank (accession no. MW012901).
Mycelium biomass from each Phytopythium
specimen was grown for 3−5 days in falcon tubes
containing 20 ml of MP5 (4 g of maltose, 1 g of pep-
tone, and 1 l of ddH2O) liquid medium at room tem-
perature (~22 ºC). Biomass was transferred to Ep-
pendorf tubes, washed with sterile water, and cen-
trifuged at 13,000 rpm for 15 min to obtain pellets.
The DNA extraction was performed according to
the protocol described in the PureLink Genomic
DNA kit (Invitrogen, Carlsbard, CA). The ITS and
LSU regions were amplied using the primers UN–
up18S42 and UN–1o28S22 (Robideau et al. 2011)
and LR0R/LR6-O, respectively (Hopple 1994, Rieth-
müller et al. 2002). DNA was amplied with Jump-
Start TM Taq DNA Polymerase (Sigma-Aldrich, St.
Louis, MO) for a nal volume of 25 μl using the PCR
conditions described by Marano et al. (2014). Am-
plicons were puried manually according to the
protocol of Schmitz & Riesner (2006). Sanger se-
quencing was performed by Macrogen. Assembly of
contigs and correction of ambiguous bases were
manually edited using Sequencher version 4.1.4. All
sequences were submitted to GenBank (Tab. 1).
Genomic DNA of Pluteus basidiomata was ex-
tracted using the ZR Fungal/Bacterial DNA Mini-
Prep kit (Zymo research, Irvine, CA). PCR ampli-
cation was performed with the following primer
pairs: ITS1F and ITS4 for ITS (White et al. 1990,
Gardes & Bruns 1993), and LR0R and LR5 for LSU
(Vilgalys & Hester 1990, Hopple 1994). PCR proce-
dures were after Kaygusuz et al. (2020). Purication
of PCR products was done using the UltraClean
PCR Clean-Up kit (MoBio Laboratories, Carlsbad,
286 Sydowia 73 (2021)
Dima et al.: FUSE 7
CA) following the manufacturer’s instructions.
Sanger sequencing of puried PCR products using
the PCR primers was outsourced to Source Biosci-
ence (Berlin, Germany). Newly generated sequences
were submitted to GenBank (accession numbers in
Tab. 1).
Seven Rhodocollybia collections from Norway
were studied by molecular methods (Tab. 1). Six of
them belong to the R. butyracea (Bull.) Lennox
complex. DNA extraction and PCR amplication of
the ITS region as well as Sanger sequencing were
done in collaboration with the Norwegian Barcode
of Life Network (NorBOL) as part of BOLD (Bar-
code of Life Data System). For BOLD methods, see
Ratnasingham & Hebert (2007, 2013).
Phylogenetic analyses
The newly generated Cortinarius sequence was
subjected to a BLAST search against NCBI Gen-
Bank’s standard nr/nt nucleotide database (https://
blast.ncbi.nlm.nih.gov/Blast.cgi) to select and
download the most closely related sequences in
Cortinarius subg. Telamonia sect. Hinnulei (Lii-
matainen et al. 2020). Sequences were aligned in
MEGA5 (Tamura et al. 2011) using ClustalW and
then corrected manually. The nal alignment in-
cluded 512 characters, of which 65 were parsimony-
informative. Bayesian inference (BI) was performed
in MrBayes 3.2.6 (Ronquist et al. 2012), under the
following parameters: data not partitioned,
GTR+G+I as substitution model, two independent
runs, four chains, temperature set to 0.2, sampling
every 100 generations, until convergence parame-
ters were met after 0.74 million generations. Finally,
a maximum likelihood (ML) analysis was run using
RAxML version 8.2.12 (Stamatakis 2014) using the
GTRGAMMA model. Bootstrapping was done with
2,000 replicates. Trees were visualized in FigTree
version 1.4.3 (http://tree.bio.ed.ac.uk/software/
gtree/) and then edited in Adobe Illustrator 25.1.
Entoloma sequences were aligned with MAFFT
version 7 using the E-INS-i option (Katoh et al.
2019). The alignment was checked and edited in
SeaView version 4 (Gouy et al. 2010). Maximum
likelihood (ML) analysis was performed in PhyML
version 3.0 (Guindon et al. 2010) using the non-par-
ametric Shimodaira-Hasegawa version of the ap-
proximate likelihood-ratio test (SH-aLRT) and the
GTR+I+Γ model of evolution. The nal tree (Fig. 4)
was edited in MEGA7 (Kumar et al. 2016) and Ado-
be Illustrator CS4.
For the Suillus study, consensus sequences were
subjected to BLAST searches (https://blast.ncbi.
nlm.nih.gov/Blast.cgi). ITS sequences sharing high-
er identity with our newly generated sequences
were downloaded. All sequences were aligned using
the MUSCLE alignment tool (Edgar 2004), which is
available online through the European Bioinfor-
matics Institute (https://www.ebi.ac.uk/Tools/msa/
muscle/). Next, sequences in the aligned ITS dataset
were trimmed at the conserved motifs 5’–(…GAT)
CATTA–3’ (3’ end of the SSU) and 5’–
GACCT(CAAA…)–3’ (5’ end of the LSU) (Dentinger
et al. 2011). A maximum likelihood (ML) tree was
inferred using RAxML-HPC2 version 8.1.11
(Stamatakis 2014) with the GTRGAMMA model of
nucleotide substitution. Rapid bootstrapping was
done with 1,000 replicates. Phylogenetic analysis
was performed on the CIPRES Science Gateway
version 3.3 (Miller et al. 2010).
Crepidotus sequences that shared >87.5% iden-
tity with the newly generated ITS sequence in
BLAST were downloaded from NCBI GenBank
(https://www.ncbi.nlm.nih.gov/genbank/). Agro-
cybe pediades (Fr.) Fayod), GenBank accession no.
MG817376, was selected as outgroup. Sequences
were aligned using MAFFT version 7.450 (Katoh &
Toh 2008) with default parameters in Geneious ver-
sion 11.1.5. Maximum likelihood (ML) was inferred
with RAxML version 8.2.11 (Stamatakis 2014). The
GTR+G model was selected, and a total of 1,000
bootstrap (MLBS) replicates were used. The average
distance between clades was estimated with MEGA
version 10.0.4 (Kumar et al. 2018).
All existing ribosomal DNA sequences of Herpo-
myces species were downloaded from GenBank. Se-
quences of two species of Pyxidiophora Bref. &
Tavel, which served as outgroup, were also down-
loaded. Sequences were aligned by region (SSU,
ITS, LSU) using MUSCLE version 3.7 (Edgar 2004)
on the CIPRES Science Gateway (Miller et al. 2020).
Aligned ITS sequences were trimmed with the con-
served motifs 5’–CATTA–3’ (3’ end of SSU) and 5’–
GACCT–3’ (5’ end of LSU); the alignment portion
between these motifs was included in subsequent
analysis (Dentinger et al. 2011). Aligned LSU se-
quences were trimmed with the 5’–GACCT–3’ motif;
the alignment portion upstream from and including
this conserved motif were included in subsequent
analysis. Next, aligned sequences were trimmed us-
ing TrimAl version 1.3 (Capella-Gutiérrez et al.
2009) with gap threshold (-gt) of 0.6 and minimal
coverage (-cons) of 0.5. Appropriate models of nu-
cleotide substitution were selected under the Akai-
ke Information Criterion corrected for small sample
size (AICc) using the IQ-TREE built-in ModelFind-
er (Kalyaanamoorthy et al. 2017). Selected models
Sydowia 73 (2021) 287
Dima et al.: FUSE 7
were GTR+F+R2 (SSU, -lnL=2066.978), TVM+F+G4
(ITS, -lnL=4635.696), and TIM3+F+R2 (LSU,
-lnL=2729.742). The data for each region were com-
bined using MEGA version 7.0.26 (Kumar et al.
2016). We used the command-line version of IQ-
TREE to perform a maximum likelihood (ML) anal-
ysis of the concatenated SSU–ITS–LSU dataset,
under multiple partitions (Nguyen et al. 2015, Cher-
nomor et al. 2016) and with branch support calcu-
lated from 1,000 ultrafast bootstrap replicates
(Hoang et al. 2018). The nal tree with ML boot-
strap (MLBS) support values was visualized in
FigTree version 1.4.3 (http://tree.bio.ed.ac.uk/soft-
ware/gtree/) and edited in Adobe Illustrator ver-
sion 25.1.
Selected ITS sequences of Leucoagaricus spp.
were aligned with MAFFT online version 7 using
the E-INS-i option (Katoh et al. 2019). The align-
ment was checked and edited in SeaView version 4
(Gouy et al. 2010). A maximum likelihood (ML)
phylogenetic analysis was performed in raxmlGUI
version 1.5 (Silvestro & Michalak 2012) using the
GTRGAMMA nucleotide substitution model.
Branch support was evaluated using 1,000 boot-
strap replicates. The best scoring tree was edited
with MEGA7 (Kumar et al. 2016).
ITS and LSU datasets with sequences for 51
Phytopythium isolates were used for the phyloge-
netic analyses, with two isolates of Pythium takay-
amanum Senda & Kageyama as outgroup. The data-
set was aligned using MAFFT 7 with default set-
tings (Katoh et al. 2019) and adjusted manually in
BioEdit (Hall 1999). The best-tting model of evolu-
tion was selected using the Akaike Information Cri-
terion in jModelTest version 0.1.1 (Posada 2008).
The aligned sequences were concatenated using Se-
quenceMatrix 1.8 (Vaidya et al. 2010), resulting in a
nal length of 2,140 characters. Maximum likeli-
hood (ML) and Bayesian inference (BI) analyses
were performed using RAxML 8.2.10 (Stamatakis
2014) and MrBayes 3.2.2 (Ronquist et al. 2012) in
the CIPRES Science Gateway. The ML analysis was
performed with 1,000 bootstraps (MLBS) replicates
using the GTRGAMMAI model. The BI used the
Markov Chain Monte Carlo (MCMC) methodology
to calculate posterior probabilities (BIPP). Four
MCMC chains were run for ve million generations,
with sampling every 1,000 generations, and a burn-
in of 10 %. Phylogenetic trees were viewed in
FigTree version 1.4.2 (http://tree.bio.ed.ac.uk/soft-
ware/gtree/) and edited in Inkscape version 0.92
(https://inkscape.org/).
For the Pluteus study, two ITS and two LSU se-
quences were newly generated. Another 51 related
sequences—31 ITS and 12 LSU—were downloaded
from GenBank and UNITE. Pluteus seticeps (G.F.
Atk.) Singer, GenBank accession number HM562192,
was selected as the outgroup. All sequences were
aligned by MAFFT version 7.110 (Katoh et al. 2019).
Phylogenetic analyses were performed for the con-
catenated ITS–LSU dataset with both maximum
likelihood (ML) and Bayesian inference (BI) meth-
ods. The ML analysis was run through the CIPRES
Science Gateway online interface (Miller et al. 2010)
using RAxML version 8.2.10 (Stamatakis 2014) un-
der the GTRGAMMA model and with 1,000 boot-
strap replicates. BI was performed using MrBayes
version 3.2.5 (Ronquist et al. 2012). Four Markov
chain Monte Carlo (MCMC) chains were run for 1
million generations, sampling trees every 100 gen-
erations. The rst 25% of sampled trees were dis-
carded as burn-in.
The newly generated Rhodocollybia sequences
were compared with those deposited in public data-
bases (GenBank, UNITE) using the BLAST search
tool (Altschul et al. 1990). Identical or most similar
sequences were downloaded. Sequences belonging
to the R. maculata (Alb. & Schwein.) Singer com-
plex were selected as outgroup. Altogether 24 se-
quences were included in our nal dataset. Multiple
sequence alignment was done by PRANK (Löytyno-
ja & Goldman 2005) as implemented in its graphi-
cal interface (PRANKSTER) with default settings.
After manual adjustment in SeaView version 4
(Gouy et al. 2010), the alignment included 783 char-
acters. The phylogenetically informative indels were
coded following the simple indel coding algorithm
(Simmons et al. 2001) with the program FastGap 1.2
(Borchsenius 2009). Adding indel characters to the
nucleotide alignment of ITS sequences increases
the robustness of phylogenetic analyses (Nagy et al.
2012). The nal matrix including nucleotide + bi-
nary data was 829 characters in length. Bayesian
inference (BI) analyses were performed with Mr-
Bayes version 3.1.2 (Ronquist & Huelsenbeck 2003).
The nucleotide and indel characters were split into
two partitions to which the GTR+G and two-state
Markov model, respectively, were applied. Four
Markov chains were run each for 5,000,000 genera-
tions, sampling every 100 generations. A burn-in of
12,500 trees was established. Sampled trees from
both runs were combined in a 50 % majority rule
consensus phylogram and posterior probabilities
(BIPP) were calculated (Fig. 34). In addition, maxi-
mum likelihood (ML) analysis was carried out using
RAxML version 7 (Stamatakis et al. 2008) in raxm-
lGUI (Silvestro & Michalak 2012) with 1,000 boot-
strap replicates under the GTRGAMMA substitu-
288 Sydowia 73 (2021)
Dima et al.: FUSE 7
Fig. 1. Laboulbeniales spp. nov. A–C. Cantharomyces paschalis, slide WR4370 (holotype). D–E. Cryptandromyces pinguis, slide
SS·E534 (holotype). F–G. Cryptandromyces tricornis, slide SS·E482a (holotype). Scale bar 50 μm.
Sydowia 73 (2021) 289
Dima et al.: FUSE 7
tion model for DNA and the default set for binary
(indel) characters. The nal tree was edited in
MEGA7 (Kumar et al. 2016) and Adobe Illustrator
CS4.
Taxonomy
Ascomycota, Laboulbeniomycetes, Laboulbeniales,
Laboulbeniaceae
Cantharomyces paschalis
W. Rossi & Santam., sp.
nov. – Fig. 1
MycoBank no.: MB 839553
H o l o t y p u s . – CHILE. Rapa Nui Island, Maugaoa, on
Mimopaederus insularis Cameron, 1936 (Coleoptera, Staphyli-
nidae, Oxytelinae), 11 December 1963, leg. J.F.G. & T.M. Clarke,
slide WR4370 (4 mature and 11 immature or incomplete thalli;
holotype at FI).
D es cr ip ti on . – Cells I and VI almost hyaline;
the rest of the thallus yellowish brown, with a large,
contrasting, dark spot on the upper, inner side of
cell III and a second spot, lighter and smaller than
the previous, on the ventral, lower angle of cell II. –
C e l l I obtrapezoidal in section, slightly longer
than maximum width. – C e l l I I larger, irregularly
shaped or broadly pentagonal. – C e l l I I I quad-
rangular, slightly broader than long. – A p p e n d -
a g e with the lower cell larger than cell III, irregu-
larly quadrangular, the inner side longer than the
outer, with the elongate antheridium occupying
about 1/3 of the cell on the inner side; the second
cell of the appendage is slightly narrower than the
rst but much shorter, bearing the hemispherical
third cell that gives rise to two long, diverging
branches, the outer of which bifurcates above its
lower cell. Stalk cell of the peritheci-
u m relatively short, slightly broader distally. –
P e r i t h e c iu m ovoid, very slightly asymmetrical,
ending in a truncate tip with almost attened and
hyaline apex. – Length from foot to perithecial apex
110–140 μm; longest branch of the appendage
125 μm; perithecium, including basal cells 50–63 ×
25–30 μm.
E t y m o l o g y. – From Latin Pascha (Easter in
English, Pascua in Spanish), referring to the holo-
type locality, Easter Island (or Isla de Pascua).
Hosts and distribution. Thus far only
known from Mimopaederus insularis (Coleoptera,
Staphylinidae, Oxytelinae) on Rapa Nui Island (also
known as Easter Island or Isla de Pascua), Chile.
Additional material examined. Ibid., slide
WR4371 (1 mature thallus; paratype at FI).
N o t e s . – To date, 27 species have been de-
scribed in the genus Cantharomyces, most of which
(19) are associated with Staphylinidae; the remain-
ing species are found on riparian insects of the fam-
ilies Dryopidae, Limnichidae, and Hydrochidae
(Picard 1912, Thaxter 1931, Haelewaters & De Ke-
sel 2013). Cantharomyces paschalis might be com-
pared with C. chilensis Thaxt., described from Thi-
nodromus signatus (Erichson, 1834) [as Tro-
gophloeus signatus Sol.] from Chilean mainland,
which is more than 3,600 km away from Easter Is-
land. The latter fungus differs from the new species
mainly for the blackish color of a large portion of
the thallus, for the more numerous and shorter
branches of the appendage and for the distinctly
larger antheridium (Thaxter 1931).
Authors: S. Santamaria & W. Rossi
Basidiomycota, Agaricomycetes, Agaricales, Corti-
nariaceae
Cortinarius squamosus
E. Sesli, sp. nov. – Figs. 2, 3
Index Fungorum no.: IF 557116
D i a gn o s i s . – Characterized by a reddish brown, quite
scaly, irregular, hygrophanous, broadly umbonate pileus; ad-
nate, distant and reddish brown lamellae; very thin content;
cylindrical or clavate, curved, reddish brown and brous stipe,
almond-like odor; and growing under Castanea.
H o l o t y p u s . – TURKEY. Trabzon Province, Çilekli,
40°58’01.99”N, 39°46’16.68”E, 406 m a.s.l., in forest of Cas-
tanea sativa (Fagales, Fagaceae), 19 September 2014, leg. E.
Sesli (KATO Fungi 3386; holotype). Sequences ex-holotype:
MW314263 (ITS), MW314262 (LSU).
Description. Pileus conical then con-
vex to plane, broadly umbonate, 15–30 mm in diam.,
margin crenate, ± incurved, reddish brown, distinct-
ly scaly, irregular, hygrophanous, darker reddish
brown when moist, lighter when dry, remaining red-
dish in some parts; edge lighter than the pileus sur-
face. – L a me l l a e adnate, distant, reddish brown,
broad, thick, edges entire to rarely crenate, L = 20–
30, I = 14. Context of pileus very thin,
nearly as thick as a lamella except where it is at-
tached to the pileus. – T a s t e indistinct. – O d o r
resembling almonds. – S t i p e cylindrical to clavate,
sometimes attened, generally curved, 30–40 ×
3–5 mm, hygrophanous, reddish brown, lighter in
places, brous, with a mass of white mycelium at
the base. – B a s i d i o s p o r e s (7.2–)7.5–9.0(–9.6) ×
(4.8–)5.5–6.5(–7.2) μm, average 8 × 6 μm [n = 65],
Q=1.2–1.4, light ochre or greenish-yellow; ellipsoid
to subglobose, typically verrucose. – B a s i d i a
clavate, (20–)25–30(–35) × (6.3–)7.0–9.0(–10.7) μm
[n = 40], with 4(2) sterigmata and clamped, some
with granular contents. – M a r g i n a l c e l l s
clavate. – E p i c u t i s composed of cylindrical, 3.3–
7.6 μm hyphae. – H y po c u t is composed of broad-
ly cylindrical to enlarged, 35.5–63.7 × 13.1–18.9 μm
290 Sydowia 73 (2021)
Dima et al.: FUSE 7
Fig. 2. Phylogeny of Cortinarius subgen. Telamonia sect. Hinnulei, reconstructed from an ITS dataset. The topology is the result
of ML inference performed with RAxML. For each node, BIPP (if 0.95) and MLBS (if 70) are presented above the branch lead-
ing to that node. Newly generated sequence highlighted in boldface.
Sydowia 73 (2021) 291
Dima et al.: FUSE 7
Fig. 3. Cortinarius. squamosus, collection KATO Fungi 3386
(holotype). a–b. Basidiomata. c. Section through the hymeni-
um. d. Pileipellis. e. Stipitipellis. f. Basidiospores. Scale bars
a–b 10 mm, c–f 10 μm.
encrusted hyphae. – C l a m p c o n n e c t io ns pre-
sent.
Etymology. from squamosus (Latin),
meaning covered with or characterized by scales,
referring to the distinctly scaly pileus.
Habitat and distribution. Solitary to
gregarious under Castanea sativa. Most probably
saprotrophic, fruiting in autumn. Known only from
the East Black Sea Region of Turkey so far.
Additional material examined. TURKEY.
Trabzon Province, Maçka, Sevinç neighborhood, in mixed de-
ciduous forest under Castanea sativa (Fagales, Fagaceae),
2 September 2020, leg. E. Sesli (KATO fungi 4325; paratype).
N o t e s . – The ITS sequence obtained from the
holotype collection KATO Fungi 3386 shared 98%
identity with C. subrigidipes M.M. Moser
(MT935523) and C. paraphaeochrous M.M. Moser
(MT935282) of Cortinarius subgenus Telamonia
section Hinnulei. A threshold of 99 % is sufcient to
recognize species in Cortinarius using the ITS re-
gion (Garnica et al. 2016). Our phylogenetic analy-
sis of selected sequences from this section (Fig. 2)
placed C. squamosus sister to C. scabridipileus Ky-
tov., Liimat. & Niskanen, with quite some evolu-
tionary distance between both sequences. Morpho-
logically, C. scabridipileus differs by the following
characteristics: slightly larger, unequally plano-
convex to very shallowly plano-depressed, cacao
brown to hazel brown, slightly squarrose pileus;
somewhat tapering or thickened, whitish silky--
brillose stipe; and ellipsoid to obovoidly ellipsoid or
subglobose basidiospores (Niskanen 2020).
Cortinarius conicoumbonatus E. Sesli, Liimat.
& K. Demirel is a similar species that differs by its
coniferous habitat (associated with Picea). Mor-
phologically, it differs from C. squamosus in having
a cream-colored, light ochre to beige brown, smooth
pileus; a somewhat whitish stipe; light ochre-
brown to rust-brown lamellae; an unpleasant and
sharp odor; and broadly ellipsoid to obovoid ba-
sidiospores measuring 7.9 × 5.2 μm (Sesli & Lii-
matainen 2018, Sesli et al. 2020). Another close
species, C. hinnuleus Fr., has a 25–70 mm in diam.,
hemispherical to conical, pale reddish to orange
brown pileus; 30–120 mm, whitish brillose to red-
dish brown stipe; earthy smell; and subglobose to
obovoid basidiospores measuring 7–9 × 5–6.5 μm
(Breitenbach & Kränzlin 2000, Niskanen & Kytö-
vuori 2008, Soop 2018). Cortinarius hinnuleoarmil-
latus Reumaux differs by its larger pileus (30–80
mm in diam.), raphanoid smell, longer stipe (60–
100 mm), and longer basidiospores (8.5–10 μm)
(Niskanen & Kytövuori 2008, Soop 2018). Corti-
narius roseonudipes Rob. Henry & Moënne-Locc. is
different from the new species by its yellowish or
dark brown and larger pileus (25–60 mm in diam.),
longer stipe (40–120 mm), and slightly longer ba-
sidiospores (8.5–10 μm) (Niskanen & Kytövuori
2008, Soop 2018). A Salix-associated species, C.
minutalis Lamoure, also placed in sect. Hinnulei,
has a hemispherical to conical, red brown to yel-
lowish brown pileus, and slightly larger basidio-
spores (Breitenbach & Kränzlin 2000, Niskanen &
Kytövuori 2008).
A number of phylogenetically distantly related
species placed in different sections (Liimatainen et
al. 2020) are similar to C. squamosus as well. Cor-
tinarius safranopes Rob. Henry (sect. Rubricori =
sect. Safranopedes) differs from C. squamosus by
its yellowish brown pileus and lamellae; longer
stipe (40–110 mm); obovoid-ellipsoid to dacryoid
and longer basidiospores (9–10 μm) (Niskanen &
Kytövuori 2008). Cortinarius umbrinolens P.D. Or-
292 Sydowia 73 (2021)
Dima et al.: FUSE 7
ton is a small species with dark brown, bell-shaped
to expanded pileus; yellowish or grey brown lamel-
lae; greyish brown and brous stipe; and ellipsoid
to amygdaloid and longer basidiospores (8–9.5 μm)
(Breitenbach & Kränzlin 2000, Lindström 2012,
Soop 2018). Cortinarius aavae Liimat. & Niskanen
(sect. Leiocastanei), a conifer-associated species,
has a reddish brown or brown pileus measuring
15–40 mm in diam., an indistinct smell, and amyg-
daloid basidiospores that are smaller compared to
C. squamosus (6.5–7.5 × 4.0–4.5 μm) (Niskanen et
al. 2012).
Author: E. Sesli
Ascomycota, Laboulbeniomycetes, Laboulbeniales,
Laboulbeniaceae
Cryptandromyces pinguis
Santam. & W. Rossi, sp.
nov. – Fig. 1
MycoBank no.: MB 839555
H ol o ty p us . NICARAGUA. Departamento de Estelí,
Mesas de Moropotente, on left epipleuron of Euconnus (Napo-
chus) sp. (Coleoptera, Staphylinidae, Scydmaeninae), pitfall
trap, 23–30 August 2007, leg. P. Andrés, slide BCB SS·E534
(holotype).
D e s c r i p t i o n . – Thallus hyaline. – L o w e r
r e c e p t a c l e slender and elongate, gradually en-
larging from below upwards, with the suprabasal
cell distinctly shorter and broader than the basal.
A p p e n d a g e diverging from the perithecium,
consisting of nine cells, gradually widening up to
the third or the fourth cell, after which it tapers
rather abruptly, never exceeding the perithecial
apex. Stalk cell of the perithecium
about twice longer than broad, with the lower por-
tion joined to inner side of the basal cell of the ap-
pendage. One of the basal cells is much larger than
the others and bulges outwards. – P er it he ci um ,
including basal cells, symmetrical, broadly ovoid,
slightly more than twice longer than maximum
width, regularly tapering to the tip, which ends in
four small, subequal lips. – Length from foot to per-
ithecial apex 124–147 μm; length from foot to the
apex of the appendage 120–144 μm; perithecium,
including basal cells 58–67 × 24–29 μm.
E t y m o l o g y. – From Latin pinguis, which
means fatty, because of the stout habitus of the fun-
gus.
Hosts and distribution. Only known
from Euconnus (Napochus) sp. (Coleoptera,
Staphylinidae, Scydmaeninae) in Nicaragua.
N o t e s . – At present, the genus Cryptandromy-
ces includes 17 species, 9 of which are found on
members of subfamily Scydmaeninae (Staphylini-
dae). Although lacking any striking feature, except
for the large and bulging secondary stalk cell, C.
pinguis is quite different from any other Cryptan-
dromyces. It may only be compared to C. genicula-
tus Thaxt., described on Euconnus sp. [as Con-
nophron] from Argentina (Thaxter 1931) and subse-
quently reported from China on Euconnus sp. (Shen
& Ye 2006), which however has a laterally bent peri-
thecium and a much longer, slender appendage.
Authors: S. Santamaria & W. Rossi
Ascomycota, Laboulbeniomycetes, Laboulbeniales,
Laboulbeniaceae
Cryptandromyces tricornis
Santam. & W. Rossi, sp.
nov. – Fig. 1
MycoBank no.: MB 839566
H ol o ty p us . – NICARAGUA. Departamento de Estelí,
Mesas de Moropotente, on elytra of Euconnus (Napochus) sp.
(Coleoptera, Staphylinidae, Scydmaeninae), pitfall trap, 23–
30 August 2007, leg. P. Andrés, slide SS·E482a (holotype at BCB).
D e s c r i p t i o n . – Thallus short and hyaline. –
R e c e pt a c l e with the basal cell broadly trapezoi-
dal, about one and a half times longer than broad;
suprabasal cell triangular or pentagonal, distinctly
shorter than the previous cell. – A p p e nd a g e con-
sisting of up to ten subequal cells including the ba-
sal, slightly tapering only near the apex. – S t a l k
cell of the perithecium about twice longer
than broad, narrower in the middle. – Pe r i t h e -
c i u m asymmetrical, with the dorsal side almost
straight and the ventral distinctly convex, about
twice longer than maximum width, bearing apically
three elongate, straight, unequal projections, the
median of which is distinctly longer than the two
others. – Length from foot to the tip of the longest
projection 94–114 μm; length from foot to the tip of
the longest appendage 92 μm; perithecium, includ-
ing basal cells but not the apical projections 42–48
× 20–25 μm; longest outgrowth 26–29 μm.
E t y m o l o g y. – From Latin tria (= three) and
cornua (= horns), referring to the three, long, horn-
like perithecial projections.
Hosts and distribution. Only known
from Euconnus (Napochus) sp. (Coleoptera,
Staphylinidae, Scydmaeninae) in Nicaragua.
Additional material examined. Ibid., SSE475
(paratype at BCB).
N o t es . – The three long and erect projections of
C. tricornis are unique among the species of the
same genus and make it possible to distinguish this
new species at rst glance.
Authors: S. Santamaria & W. Rossi
Sydowia 73 (2021) 293
Dima et al.: FUSE 7
Fig. 4. Phylogeny of Entoloma subgen. Cyanula, reconstructed from an ITS dataset. The topology is the result of ML inference
performed in PhyML 3.0. For each node, the SH-aLRT support value (if >50) is presented on the branch leading to that node. New
species are marked in blue; newly generated sequences are highlighted in boldface; labels in smaller font size represent geo-
graphic origin except Europe. Bar indicates 0.05 expected changes per site per branch. Part A.
294 Sydowia 73 (2021)
Dima et al.: FUSE 7
Fig. 4. Continued. Part B.
Sydowia 73 (2021) 295
Dima et al.: FUSE 7
Fig. 4. Continued. Part C.
296 Sydowia 73 (2021)
Dima et al.: FUSE 7
Basidiomycota, Agaricomycetes, Agaricales, Ento-
lomataceae
Entoloma brunneicoeruleum
O.V. Morozova,
Noordel., Brandrud, J.B. Jordal & Dima, sp. nov.
Figs. 5a, 6
MycoBank no.: MB 839631
H o l o t y p u s . – RUSSIA. Pskov region, Pechorsky dis-
trict, eastern shore of Lake Velje, 57°38’16”N, 27°47’35”E, 90 m
a.s.l., on soil in wet AlnusBetula forest, 19 July 2007, leg. O.
Morozova (LE 302098; holotype). Sequences ex-holotype:
MZ145170 (ITS).
Description. – Basidiomata medium-
sized, collybioid. – P i l e u s 13–30 mm in diam., con-
Fig. 5. In situ photos of the new Entoloma species. A. Entoloma brunneicoeruleum, collection LE302098 (holotype). B. Entoloma
callipygmaeum, collection LE253784 (holotype). C. Entoloma minutigranulosum, collection LE 302096 (holotype). D. Entoloma
perasprellum, collection GC01100310 (holotype), E. Entoloma pulchripes, collection LE311808 (holotype). F. Entoloma tigrinum,
collection LE312493 (holotype). G. Entoloma timidum, collection LE312480 (holotype). H. Entoloma violaceoserrulatum, collec-
tion TEB 128-19. Photos O.V. Morozova (A–C, E–G), G. Corriol (D), B. Dima (H).
Sydowia 73 (2021) 297
Dima et al.: FUSE 7
ico-convex or hemispherical with more or less de-
pressed center, soon expanding to plano-convex and
applanate with slightly umbilicate center, with de-
exed and then straight margin, hygrophanous,
translucently striate almost up to the center, minute-
ly radially brillose-squamulose all over, more
densely in the center, initially greyish brown with
bluish or purplish tinge (17E4, 19E4–5) and only
slightly darker center, then greyish-brown or yellow-
brown with a darker, almost black central spot (um-
bilicus). – L a m e l l a e moderately distant, emargin-
ate or adnexed with short tooth, whitish or pale, be-
coming pinkish, with irregular to serrulate, con-
colorous edge. – St i p e 30–80 × 1.5–3 mm, cylindri-
cal or slightly broadened towards the base, some-
times compressed with longitudinal groove, dark
blue, greyish blue, steel-blue or purplish grey (20D4–
5, 20E4–6, 19D3–5), initially minutely longitudinally
striate(-brillose), but soon smooth and polished,
white tomentose at base. – C o n t e x t greyish.
S m e l l indistinct, sweetish on drying, taste not re-
ported. – Ba s i d i o s p o r e s 10.5–16 × 6.8–9.5 μm,
average 12.0–13.0 × 7.5–8.7, Q=1.3–2.0 μm, Qav=1.4–
1.7, heterodiametrical, of two types, (i) with 5–8 an-
gles in side-view, (ii) or almost nodulose in some col-
lections. – B a s i d i a 30–45 × 10.5–13.5 μm, 4-spored,
narrowly clavate to clavate, clampless. – L a m e l l a
e d g e sterile or heterogeneous. Cheilocystid-
i a 20–30(–64) × 8.5–11.5 μm, cylindrical or narrowly
clavate, often septate, usually intermixed with ba-
sidia. Caulocystidia absent. Pileipellis
a cutis with transition to a trichoderm of 4–16 μm
wide cylindrical hyphae, with ascending, cylindrical
to fusiform terminal elements 23–80 × 11–25 μm. Pig-
ment intracellular, sometimes agglutinate, brownish
in KOH. Clamp connections absent.
Etymology. Frombrunneus’ (Latin, mean-
ing brown), and ‘coeruleus (Latin, meaning blue),
referring to the brown pileus and blue stipe.
Habitat and distribution. In small
groups on soil in damp grasslands (hayeld, pas-
ture), Sphagnum bog, and damp forests with Betula
and Alnus, on richer to calcareous soil. Known from
Russia (European part), the Netherlands, Norway,
and Estonia (from soil sample).
Additional material examined. NORWAY.
Møre og Romsdal, Fjord, Valldal, Heimsetra, 62°21’00.00”N,
7°20’51.72”E semi-natural grassland (pasture), 3 September
2008, leg. J.B. Jordal, O-F-291139; Nordland, Grane,
Holmvassdalen Nature Reserve, 65°17’07.44”N, 13°18’48.58”E,
in Sphagnum in old calcareous pine forest, leg. J. Lorås &
M. Eidissen, JL 124-16; Rogaland, Strand, Strandalia,
59°02’33.36”N, 5°55’51.96”E, old wooded pasture with Betula,
in Sphagnum, 26 July 2015, leg. A.K. Svensen, AKS E1-15. –
THE NETHERLANDS. Gelderland province, Ratum, Willinks
Weust, 51°57’52.93”N, 6°46’45.94”E, 19 August 2016, leg. G.M.
Jansen (L-0608198).
Notes. Entoloma brunneicoeruleum is char-
acterized by the brown pileus, initially with a slight
bluish or purplish tinge, dark blue to steel blue
stipe, usually heterogeneous lamellae edge with un-
differentiated, cylindrical or almost cylindrical
cheilocystidia, and spores of two types, (i) medium
sized with 5–7 normally developed angles and
Fig. 6. Entoloma brunneicoeruleum, micromorphological fea-
tures drawn from the holotype (LE302098). A. Basidiospores.
B. Basidium. C. Cheilocystidia. Scale bars 10 μm, del. O.V. Mo-
rozova.
Q=1.5–1.7; (ii) large, almost nodulose with Q=1.8–
2.0. Entoloma brunneicoeruleum belongs to a group
of closely related, but phylogenetically well-sepa-
rated species around E. atrocoeruleum Noordel.,
and may be considered an E. atrocoeruleum with
more brownish pileus, hence the name. They also
differ in basidiospore size. From the other species in
this group, E. anatinum (Lasch) Donk and E. per-
asprellum and E. tigrinum (both described below),
it differs by having by far the largest basidiospores.
Entoloma anatinum differs additionally by the ab-
sence of cheilocystidia, a more distinctly brillose
stipe, and a stouter habit. Entoloma tigrinum has
ochre-tinged basidiocarps, and a very different pi-
leus surface with small, granulose squamules,
whereas E. perasprellum differs among other things
by a completely sterile lamella edge of the serrula-
tum type. The poorly known E. nigroviolaceum (P.D.
Orton) Hesler, with similar large basidiospores, has
more pronounced violaceous tinges, and a strongly
brillose stipe.
Authors: B. Dima, O.V. Morozova, M.E.
Noordeloos, T.E. Brandrud, J.B. Jordal, G.M. Jansen
& J. Lorås
298 Sydowia 73 (2021)
Dima et al.: FUSE 7
Basidiomycota, Agaricomycetes, Agaricales, Ento-
lomataceae
Entoloma callipygmaeum
O.V. Morozova, Noordel.
& Dima, sp. nov. – Figs. 5b, 7
= Entoloma gomerense sensu Noordeloos & Mo-
rozova in Mycotaxon 112: 249 (2010).
MycoBank no.: MB 839632
H ol ot y pu s. – RUSSIA. Primorsky Territory, Kedrovaya
Pad Nature Reserve, right bank of Kedrovaya River,
43°05’56”N, 131°33’21”E, on soil in ood-plain forest with
Quercus mongolica, Tilia amurensis, Acer spp., 17 August
2005, leg. O. Morozova (LE 253784; holotype). Sequences ex-
holotype: MZ145207 (ITS).
Description. – Basidiomata small, col-
lybioid. – Pi le us 7–10 mm in diam., plano-convex
with depressed center, slightly hygrophanous, deep-
ly translucently striate, ground color rather pale
bluish, blue-grey or violaceous-grey, but covered
with darker squamules, more dense at center, more
spread towards margin; squamules and striae grey-
ish blue, greyish purple, or greyish brown (18E5–7,
18F5–7, 20E5–7). – L am el l a e adnate-emarginate
with a small decurrent tooth, whitish to greyish pink
with more or less serrulate blackish purple edge. –
S t i p e 22–35 × 1–2 mm, cylindrical, dark greyish
blue (19D4–5, 19E4–5, 20D4–5), polished, glabrous,
base with white tomentum. – C o n t e x t concolor-
ous with the surface, whitish in the inner part.
Smell indistinct, taste indistinct. – B a s i d i o -
s p o r e s 8.5–11 × 6.5–9.0 μm, average 10.0 × 7.5 μm,
Q=1.3–1.5, Qav=1.4, heterodiametrical, with 5–6 an-
gles in side view. – B a s i d i a 22–32 × 8–13 μm,
clavate to elongate, clampless. – L a m e l l a e d g e
sterile. – C h e i l o c y s t i d i a 20–85 × 11–22 μm,
broadly clavate or lageniform, including those with
a rather long neck, but in some specimens, only
clavate cystidia present, with dark intracellular pig-
ment. – P i l e i p e l l i s a cutis with transition to a
trichoderm with inated, narrowly clavate or elon-
gated terminal cells 45–70 × 12–20 μm, with intracel-
lular pigment. Clamp connections absent.
Etymology. From Greek κάλλος (beauty)
and ‘πυγµαῖος (dwarsh), referring to the nice color
and small size of basidiomata.
Habitat and distribution. Sapro-
trophic on soil, among living mosses and on decayed
wood in oodplain forests. Thus far only been record-
ed from the Russian Far East (Primorski Territory).
Additional material examined. RUSSIA. Pri-
morski Territory, Sikhote-Alin Nature Reserve, vicinities of
Maisa, on living mosses in the road in mixed forest of Quercus
mongolica, Acer mono, Tilia amurensis, Pinus koreana,
45°14’19.80”N, 136°30’40.02”E, 22 August 2013, leg. O. Mo-
rozova (LE 312487); Ibid., along stream, in ood-plain forest
of Quercus mongolica, Acer mono, Tilia amurensis, Pinus ko-
reana, 27 August 2013, leg. O. Morozova (LE 312488).
N o t e s . – The tiny violaceous or blue basidio-
carps with deeply striate pileus and purplish ser-
rulate lamella edge are reminiscent of E. gomerense
Wölfel & Noordel., a Mediterranean species with a
slightly different pileus surface, and smaller, simple
basidiospores. It is also similar to E. cyanulum
(Lasch) Noordel., which, however, never has a blue-
purple serrulate lamella edge. Phylogenetically the
closest species is E. carneogriseum (Berk. & Broome)
Noordel., another species with serrulatum-type la-
mellae edge, which, however, has predominantly
yellow-brown colored basidiomata with only traces
of blue tinge near the pileus margin and in the stipe.
Authors: B. Dima, O.V. Morozova & M.E. Noorde-
loos.
Basidiomycota, Agaricomycetes, Agaricales, Ento-
lomataceae
Entoloma minutigranulosum
O.V. Morozova,
Noordel., Brandrud & Dima, sp. nov. – Figs. 5c, 8
= Entoloma ochromicaceum sensu Morozova et al.
in Novosti Sist. Nizsh. Rast. 49: 196 (2015).
MycoBank no.: MB 839633
Fig. 7. Entoloma callipygmaeum, micromorphological fea-
tures drawn from the holotype (LE253784). A. Basidiospores.
B. Cheilocystidia. C. Basidium. D. Pileipellis. Scale bars
10 μm, del. O.V. Morozova.
Sydowia 73 (2021) 299
Dima et al.: FUSE 7
H o l o t y p u s . – RUSSIA. Pskov region, Pechorsky dis-
trict, Izborsk village, Zmeinaya ravine, on calcareous grass-
land, 57°43’N, 27°51’E, 23 August 2011, leg. O. Morozova (LE
302096; holotype). Sequences ex-holotype: MZ145214 (ITS).
Description. – Basidiomata small,
collybioid. – Pi l e u s 10–20 mm in diam., hemi-
spherical to convex, expanding to plano-convex,
with blunt, slightly umbonate, or umbilicate center,
with deexed margin, not distinctly hygrophanous,
dark yellow-brown to warm ochre-brown (5B4–6,
5C4–6), with darker center (up to 6D4), entirely mi-
nutely granulose or nely squamulose, sometimes
glabrescent with age, initially not striate but be-
coming translucently striate with age. – L a m e l -
l ae moderately distant, emarginate or adnate with
a decurrent tooth, subventricose, white then pale
pink, with minutely serrulate, (in part) brown edge.
– S t i p e 20–50 × 2–3 mm, cylindrical or slightly
broadened towards base, pale brown with distinct
metallic-grey to bluish grey tinge, particularly in
the lower half, which fades with age, when very
young often with distinct greyish blue tinge on the
entire stipe, smooth, polished, base with some white
tomentum. – C o nt e x t very thin, brittle. – S me l l
a n d t a s t e inconspicuous. Basidiospores
7–12 × 5.5–7.5 μm, average 10.0 × 6.5 μm, Q=1.3–1.7,
Qav=1.5, heterodiametrical, with 5–7 distinct angles
in side-view. – B a s i d i a 31–37 × 9.5–11.5 μm,
4-spored, clavate, clampless. – L a m e l l a e d g e
sterile. – C h e i l o c y s t i d i a 35–75 × 8–14 μm,
clavate or narrowly clavate, sometimes septate,
sometimes with brownish intracellular pigment in
KOH. Hymenophoral trama regular, made
up of cylindrical elements, 50–160 × 5–22 μm.
P i l e i p e l l i s a cutis with transition to a tricho-
derm made up of 5–12 μm wide cylindrical hyphae,
and clavate, broadly clavate or sphaeropedunculate
terminal elements, 40–100 × 14–45 μm, with brown
granular intracellular pigment. – C a u l oc y s t i d -
ia absent. – Clamp connections absent.
E t y m o l o g y. – From Latin, minutum (very
small) and ‘granulosum’, referring to the nely
granulose pileus.
Habitat and distribution. In small
groups on soil on calcareous semi-natural grass-
lands (pastures), once also recorded from alpine
heath. Known from Austria, Denmark, France, Ger-
many, the Netherlands, Norway, Sweden, and Rus-
sia (European part and Siberia).
Additional material examined. RUSSIA. Al-
tay Republic, Altaiskiy Nature Reserve, cordon Chelyush,
46°07’07.97”N, 11°06’56.76”E, grassland near farm, 28 August
2018, leg. O. Morozova, LE 312483; Ibid., LE 312484. – AUS-
TRIA. Vorarlberg, Lechtal, Dalaas, Tannlägeralpe,
47°09’20.91”N, 9°54’16.98”E, August 2004, leg. A. Hausknecht
Fig. 8. Entoloma minutigranulosum, micromorphological fea-
tures drawn from the holotype (LE 302096). A. Basidiospores.
B. Basidium. C. Cheilocystidia. D. Pileipellis. Scale bars 10
μm, del. O.V. Morozova.
(WU25057, as E. glaucobasis). – GERMANY. Bayern, Leng-
gries, Hohenwiesen, Hohenwiesener Berg, Hochalm,
47°36’32.29”N, 11°35’11.49”E, 2 August 2014, leg. M. Dondl,
MD-2014-7 (L0607941). – THE NETHERLANDS. Gelderland
Province, Staverden, Leemputten, 52°02’40.74”N,
6°40’12.70”E, October 2019, leg. Finy Salzman, L0607681. –
NORWAY, Finnmark, Porsanger, Kolvik N, 70°17’33.00”N,
22°06’52.20”E, calcareous seminatural pasture, 12 August
2015, leg. G. Gaarder, GG 7142; Nordland, Grane, Ner-Laks-
fors, 65°37’47.64”N, 13°16’11.64”E, calcareous meadow along
the river, 14 August 2003, leg. G. Gaarder, GG 3568 (O-F-
175913); Nordland, Grane, Holmvassdalen Nature Reserve,
65°18’52.32”N, 13°18’08.49”E, low herb vegetation in calcare-
ous spruce forest, 22 August 2011, leg. J. Lorås & S. Eidissen,
JL 99-11; Ibid., 65°33’19.43”N, 13°45’52.18”E, 4 September
2014, JL 54–14; Ibid., 65°18’57.08”N, 13°19’12.87”E, JL 99-16;
Nordland, Grane, Danielåsen Nature Reserve, 65°33’11.42”N,
13°38’26.33”E, tall herb vegetation in calcareous spruce for-
est, 28 August 2017, leg. J. Lorås & S. Eidissen, JL 69–17; Opp-
land, Nord-Aurdal, Øystre Slidre, Skrautvål, Jordet (Bundli
Ø), 61°02’22.92”N, 9°12’25.92”E, calcareous pasture (shallow
soil), 6 August 2016, leg. T.E. Brandrud & G. Gulden, TEB 121-
16 (O-F-304886). – SWEDEN. Lule lappmark, Jokkmokk, Pad-
jelanta, Ajajaure, 67°24’28.72”N, 16°44’40.94”E, alpine heath
with Betula nana and Salix spp., 16 August 2016, leg. G. Gul-
den (GB-0204540).
N o te s . – Typically the stipe of this species ap-
pears to be bicolored: ochre-brown at the apex, and
steel-grey with a glaucous-bluish tinge at the base.
300 Sydowia 73 (2021)
Dima et al.: FUSE 7
This is reminiscent of E. glaucobasis Huijsman ex
Noordel., which, however, has a distinctly brillose-
striate stipe surface, and larger basidiospores. When
very young the entire stipe can be bluish, and in
such stages this species may be misidentied as E.
poliopus var. parvisporigerum Noordel., but this
species has a more bluish, darker pileus, and is a
member of the /chalybeum clade. Entoloma per-
asprellum may appear similar, but has larger, nodu-
lose-angled basidiospores. Entoloma minutigranu-
losum was erroneously referred to as E. ochromica-
ceum Noordel. & Liiv by Morozova et al. (2015), but
the latter species lacks grey-bluish-metallic tinges
in the stipe, and has larger spores (Noordeloos
1992). We note that E. minutigranulosum was also
present in an environmental soil sample from cal-
careous grassland in the province of Limburg, the
Netherlands (data not shown).
Authors: B. Dima, O.V. Morozova, M.E.
Noordeloos, T.E. Brandrud, J.B. Jordal, G.M. Jansen,
E. Larsson & J. Lorås
Basidiomycota, Agaricomycetes, Agaricales, Ento-
lomataceae
Entoloma perasprellum
Corriol, Dima, O.V. Mo-
rozova, J.B. Jordal & Noordel., sp. nov. – Figs. 5d, 9
MycoBank no.: MB 839634
H o l o t y p u s . – FRANCE. Seine-et-Marne, Fontaineb-
leau, near the Hippodrome de la Solle, 3 October 2001, about
thirty basidiomata, leg. G. Corriol (GC01100310; holotype in
BBF). Sequences ex-holotype: MZ145177 (ITS).
Description. – Basidiomata medium-
sized, collybioid. – P i l e u s 10–40 mm in diam.,
campanulate then expanding to convex or plano-
convex, usually distinctly umbilicate, with deexed
margin, which is often crenulate, and sometimes
undulating-lobed, opaque, not hygrophanous, in-
distinctly translucently striate at rst, soon clearly
translucently striate, grey-blue at rst, but blue
tinge quickly disappearing (Mu 10YR 5/4), entirely
brillose to squamulose, then with neat, small, grey-
brown (10YR 3/3) squamules on a paler yellow-
brown (10YR 3/3) background. – L am el l a e rath-
er distant, adnate-emarginate, white to greyish,
with an irregular edge that is often brown pigment-
ed, but non-pigmented forms occur frequently. –
S t i p e 25–50 × 2–5 mm, cylindrical to compressed,
glabrous, polished, or with ne pruinose-brillose
covering, when fresh often bright blue or blue-grey,
quickly pallescent to pale grey or pale greyish
brown, paler than pileus, with white mycelium at
base. – C o n t e x t white. – S m e l l indistinct or
somewhat sweet, like owers. – B a s i d i os po r e s
(8.9–)9.4–11.0(–12.4) × (6.0–)6.4–7.6(–8.3) μm, aver-
age 10.0 × 7.0 μm; Q = (1.25–)1.30–1.60(–1.81),
Qav=1.45, heterodiametrical, with 6–7(–8) fairly pro-
nounced angles, but with a tendency to be nodulose.
– B a s i d i a 28–40 × 11–12 μm, cylindrical-clavate,
4-spored, clampless. – L a m e l l a e d g e sterile. –
C h e i l o c y s t i d i a 19–38 × 3–6 μm, well-devel-
oped, subcylindrical to narrowly clavate, with
brown, intracellular pigment. – C a u l o c y s t i d i a
not observed. – P i l e i pe ll is a transition between
a cutis and a trichoderm, made up of inated-
clavate terminal elements, up to 25 μm. – P ig m e nt
brown, intracellular. – C l a m p c o n n e c t i o n s
absent.
E t y m o l o gy. – Referring to the resemblance to
E. asprellum.
Habitat and distribution. Found
mainly in calcareous alpine heaths and subalpine
grasslands (pastures), but also in temperate/nemor-
al xerophytic grassland on base-rich sandy soil, with
Clitocybe collina (Velen.) Klán, C. costata Kühner &
Romagn., Lycoperdon lividum Pers., Marasmius
oreades (Bolton) Fr., Hebeloma cistophilum Maire
(holotype site). Known from France, Norway, Swe-
den, and Russia (Caucasus and Kamchatka).
Additional material examined. Ibid. (L, iso-
type). – NORWAY. Trøndelag, Rennebu, Jøldalen, Bortstuguse-
tra, 62°52’15.96”N, 9°32’52.80”E, seminatural pasture, 23 Au-
gust 2018, leg. J.B. Jordal, JBJ 18-004 (O-F-256732). – RUS-
Fig. 9. Entoloma perasprellum, micromorphological features
drawn from the holotype (GC01100310). A. Basidiospores.
B. Basidium and basidioles. C. Lamella edge. Scale bars
10 μm, del. G. Corriol.
Sydowia 73 (2021) 301
Dima et al.: FUSE 7
SIA. Karachaevo-Cherkesia Republic, Teberda Biosphere Re-
serve, Klukhor pass, 43°21’06.97”N, 41°41’46.22”E, ~2700 m
a.s.l., on alpine grassland, 23 August 2012, leg. O. Morozova
(LE 312501); Kamchatka Region, vicinities of Esso, near the
village, 55°55’37.12”N, 158°41’04.81”E, 470 m a.s.l., in grass-
land, 5 August 2005, leg. O. Morozova (LE 312499); Ibid., on
hill, ~800 m a.s.l., in subalpine grassland, 5 August 2005, leg. O.
Morozova (LE 312500). – SWEDEN. Åsele lappmark, Vilhelmi-
na, Lasterfjället, Tjårronunjes, 65°15’27.40”N, 14°37’38.24”E,
on alpine meadow on calcareous ground, 19 August 2019, leg.
E. Larsson, EL-28-19 (GB-0204546); Ibid., 65°15’27.39”N,
14°37’40.86”E, on alpine meadow on calcareous ground, 19
August 2019, leg. J.B. Jordal, JBJ 19-107 (GB-0204547); Åsele
lappmark, Vilhelmina, Murfjället, 65°10’20.15”N,
15°08’03.97”E, on alpine heath on calcareous ground, 20 Au-
gust 2019, leg. J.B. Jordal, JBJ 19-119 (GB-0204550); Ibid., JBJ
19-122 (GB-0204548); Jämtland, Frostviken, NW slope of
Raavre, 64°09’55.35”N, 17°21’17.57”E, in low alpine calcare-
ous heath, 23 August 2019, leg. J.B. Jordal, JBJ 19-180 (GB-
0204549).
Notes. Entoloma perasprellum resembles E.
asprellum (Fr.) Fayod, with its usually entirely nely
squamulose pileus and blue-grey polished stipe.
The new species differs by the almost nodulose ba-
sidiospores, and having a distinctly sterile lamella
edge of the serrulatum-type. Initially, Corriol (2016)
gave it the provisional name E. cyaneoturci, as the
type specimens did have some resemblance to E.
turci (Bres.) M.M. Moser, differing from it by the ba-
sidiospore shape and blue tinges in the basidioma-
ta. However, now we have more collections from a
rather wide geographic area, the variability appears
much bigger, and the resemblance with E. turci is
less striking. On the contrary, this new species is
much more morphologically similar to E. asprellum,
hence the name.
Authors: B. Dima, O.V. Morozova, M.E.
Noordeloos, T.E. Brandrud, J.B. Jordal, G. Corriol,
G.M. Jansen & E. Larsson
Basidiomycota, Agaricomycetes, Agaricales, Ento-
lomataceae
Entoloma pulchripes
O.V. Morozova, Noordel.,
Brandrud & Dima, sp. nov. – Figs. 5e, 10
MycoBank no.: MB 839635
H o l o t y p u s . – RUSSIA. Tver region, Zubtsov district,
2 km SE from Mozgovo village, open place in calcareous pine
forest, right bank of Derzha river, 56°12’49.90”N,
034°48’09.00”E, 11 September 2015, leg. O. Morozova (LE
311808; holotype). Sequences ex-holotype: MZ145188 (ITS).
Description. Basidiomata small to
medium-sized, collybioid. – P i l e u s 10–25 mm in
diam., obtuse-conical, conico-convex or hemispher-
ical with slightly depressed center, soon expanding
to plano-convex with convex or slightly umbilicate
center, with deexed then straight margin, hy-
grophanous, translucently striate up to the center,
minutely radially brillose-squamulose all over,
more densely in the center, initially greyish brown
(8D3, 8E3–4) with blackish-purple center, changing
color with age to greyish-brown or yellowish-brown
with contrasting brownish-black center, sometimes
radially cracked when drying out, showing white
context. – L a me ll a e moderately distant, adnate-
emarginate, segmentiform to decurrent with short
tooth, whitish, pale or greyish, becoming pinkish-
grey, with irregular, more or less serrulate brown
edge (sometimes almost concolor, hard to see).
S t i p e 30–80 × 1.5–3 mm, cylindrical or slightly
broadened towards the base, sometimes compressed
with longitudinal groove, smooth, polished or mi-
nutely longitudinally striate, dark blue, greyish
blue or steel-blue (20D4–5, 20E4–6), white tomen-
tose at the base. – C o n t e x t white, greyish under
the surface. – S m e l l indistinct, sweetish on dry-
ing, taste not reported. – B a s id io s p o r e s 7–11.5
Fig. 10. Entoloma pulchripes, micromorphological features
drawn from the holotype (LE311808). A. Basidiospores.
B. Cheilocystidia. C. Basidium. D. Pileipellis. Scale bars
10 μm, del. O.V. Morozova.
302 Sydowia 73 (2021)
Dima et al.: FUSE 7
× 5–7.5 μm, average 9.0 × 6.5 μm, Q=1.3–1.6, Qav =
1.45, heterodiametrical, with 5–7 angles in side-
view, relatively simple. – B a s i d i a 27–34 × 10.5–
11.5 μm, 4-spored, narrowly clavate to clavate,
clampless. – L a m e l l a e dg e sterile. – C he i l o -
c ys t id i a 40–90 × 16–31.5 μm, broadly clavate, ve-
siculose or spheropedunculate, consisting of cylin-
drical basal and spherical to oblong apical cells,
14–90 × 9–32 μm, often septate, with or without
brown intracellular pigment in KOH. – P i le ip el -
l is a cutis with transition to a trichoderm of cylin-
drical 8–13 μm wide hyphae, with ascending termi-
nal elements, 40–85 × 16–26 μm, and brownish in-
tracellular pigment in KOH. – C a u l o c y s t i d i a
absent. Clamp connections absent.
E t y mo l o gy. – From Latin, pulchrum (= beau-
ty), referring to the beautifully colored stipe.
Habitat and distribution. In small
groups on soil in grasslands and calcareous pine
forest. Known from Russia (European part).
Additional material examined. RUSSIA.
Tver region, Zubtsov district, 2 km SE from Mozgovo village,
56°12’49.90”N, 34°48’09.00”E, open place in calcareous pine
forest, right bank of the Derzha river, 11 September 2015, leg.
O. Morozova (LE 311809); Novgorod region, Valgay district,
Valdaysky National Park, bank of Msta river, 58°16’49.19”N,
31°03’55.39”E, calcareous grassland, 23 September 2011, leg.
O. Morozova (LE 312485).
Notes. Entoloma pulchripes is characterized
by the brown pileus, sometimes having a minute
purplish tinge; dark blue to steel blue stipe; small
basidiospores; and distinctive vesiculose or
spheropedunculate cheilocystidia. It resembles E.
poliopus (Romagn.) Noordel., from which it differs
by the smaller basidiospores and the shape of
cheilocystidia. Due to the brown lamella edge, E.
pulchripes could be mistaken for E. brunneiserrula-
tum Eyssart. & Noordel., which, however, is distin-
guished by the brown stipe, large basidiospores, and
elongate terminal cells of cheilocystidia. Entoloma
sodale Kühner & Romagn. ex Noordel., a species
with subglobose and spheropedunculate cheilocyst-
idia, differs by the warm brown color of the pileus
and lager basidiospores. According to our phyloge-
netic analysis (Fig. 4), the closest related species is
E. chalybaeum (Pers.) Noordel. However, this spe-
cies differs by its dark blue pileus (vs. brown pileus
in E. pulchripes). One sequence from California,
USA (GenBank accession no. MG966312, Mushroo-
mObserver.org/240552) is very close to E. pul-
chripes, sharing 99% identity. This potential con-
specicity needs to be studied further.
Authors: B. Dima, O.V. Morozova, M.E.
Noordeloos, T.E. Brandrud & G.M. Jansen
Basidiomycota, Agaricomycetes, Agaricales, Ento-
lomataceae
Entoloma tigrinum
Noordel., O.V. Morozova, Bran-
drud, J.B. Jordal & Dima, sp. nov. – Figs. 5f, 11
MycoBank no.: MB 839636
H o l o t y p u s . – NORWAY. Nordland, Steinkjer, Kvams-
fjellet north of Lystjørna (Vesterolsenget), 64°12’46.08”N,
11°49’03.72”E, edge of rich fen, 20 August 2016, leg. M.E.
Noordeloos, O. Morozova & J.B. Jordal (O-F-304580; holo-
type). Sequences ex-holotype: MZ145176 (ITS).
Description. – Basidiomata small,
collybioid. – Pi l e u s 14–20 mm, convex, expand-
ing, umbilicate, with deexed to straight margin,
hygrophanous, background colour rather vivid
ochre-brown (“golden”) to brown, covered with
ne, darker sepia to blackish brown dots and radi-
ally arranged squamules, densest at the center, ob-
scurely translucently striate up to half the radius or
to umbilicus, – L a m e l l a e L=34–40, l=1–5, moder-
ately distant, adnate, slightly emarginate with a de-
current tooth, whitish, turning pale pink, with an
entire, concolorous edge. – S t i p e 30–50 × 2–4 mm,
cylindrical, pale ochre brown or greyish brown,
much paler than the pileus, polished or innately -
brillose, with white basal mycelium, rarely slowly
staining reddish at the base. – Co n t e x t pale. –
Fig. 11. Entoloma tigrinum, micromorphological features
drawn from the holotype (LE312493). A. Basidiospores. B. Ba-
sidium C. Pileipellis. Scale bars 10 μm, del. O.V. Morozova.
Sydowia 73 (2021) 303
Dima et al.: FUSE 7
S m e l l indistinct, taste not reported. – B as i d i o -
s po r e s 10–13 × 7.0–8.5 μm, average 11.0 × 7.8 μm,
Q=1.3–1.7, Qav=1.5; heterodiametrical, 5–8-angled in
side-view. – B a s i d i a 35–52 × 10–13 μm, clavate,
4-spored, clampless. – L a m e l l a e d g e fertile. –
Cheilocystidia absent. Hymenophoral
t ra m a regular, made up of cylindrical to inated
elements, up to 25 μm wide. – P i l e i pe l l i s a hy-
meniderm of broadly clavate, broadly fusiform or
ellipsoid terminal elements, 50–130 × 10–28(–50)
μm. – P i g m e n t brown, intracellular, diluted and
in the form of golden brown granules. – P i l e i t -
r am a of cylindrical to inated hyphae, up to 23 μm
wide with abundant brilliant granules. – S t i p i t i -
p e l l i s a cutis of cylindrical 4–8 μm wide hyphae,
with a few sub-clavate terminal 7–12 μm wide end-
ings, with brown, hyaline, intracellular pigment. –
Clamp connections absent.
E t y m o l o gy. – Referring to the resemblance of
the pileal surface to that of Lentinus tigrinus.
Habitat and distribution. Calciphil-
ous, terrestrial in alpine meadows and heaths, but
also in calcareous fen in a rather open Picea forest.
Thus far only reported from northern Norway and
Swedish Lapland.
Additional material examined. Ibid. (LE
312493; isotype). – SWEDEN. Lule lappmark, Jokkmokk, Pad-
jelanta, Arranoaijvve, 66°36’19.46”N, 19°49’46.78”E, on alpine
heath on calcareous ground, 11 August 2016, leg. J. Olsson
(GB-0204539); Ibid., Vielggisbakte, 66°36’19.46”N, 19°49’46.78”
E, on alpine meadow on calcareous soil, 12 August 2016, leg. J.
Olsson (GB-0204538); Åsele lappmark, Vilhelmina, Lasterfjäl-
let, Tjårronunjes, 64°09’55.47”N, 17°19’11.50”E, on alpine
heath on calcareous ground, 19 August 2019, leg. J.B. Jordal,
JBJ 19-109 (GB-0204537); Ibid., Murfjället, 65°10’20.15”N,
15°08’03.97”E, on alpine heath on calcareous ground, 20 Au-
gust 2019, leg. J.B. Jordal, JBJ 19-128 (GB-0204536); Pite
lappmark, Arjeplog, NE side of Mt. Ákháris, 66°02’45.45”N,
17°50’23.73”E, on alpine heath calcareous soil, 14 August
2018, leg. E. Larsson, EL156-18 (GB-0204535).
Notes. Entoloma tigrinum is distinctive be-
cause of the warm ochre-brown pileus with darker
greyish brown center, which often has a granulose-
squamulose surface alternating with lighter patch-
es—resembling a bit the pileal surface of Lentinus
tigrinus (Bull.) Fr., and the fertile lamella edge. It is
reminiscent of E. leochromus Noordel. & Liiv, a
more or less similarly colored species with fertile
lamella edge. However, this species is phylogeneti-
cally distant. Entoloma minutigranulosum has a
similarly colored pileus, but differs in having glau-
cous to blue tinges in the stipe and a sterile lamella
edge.
Authors: B. Dima, O.V. Morozova, M.E.
Noordeloos, T.E. Brandrud, J.B. Jordal, G.M. Jansen
& E. Larsson
Basidiomycota, Agaricomycetes, Agaricales, Ento-
lomataceae
Entoloma timidum
O.V. Morozova, Noordel., Bran-
drud, J.B. Jordal & Dima, sp. nov. – Figs. 5g, 12
MycoBank no.: MB 839637
H o lo t y pu s . – RUSSIA. Karachaevo-Cherkesia Repub-
lic, Teberda Nature Reserve, near Teberda town, on grassland
in broad-leaved forest, 43°26’19.92”N, 41°43’52.20”E, 1350 m
a.s.l., 21 August 2012, leg. O. Morozova (LE 312480; holotype).
Sequences ex-holotype: MZ145197 (ITS).
Description. – Basidiomata small,
collybioid. – Pi l eu s 15–35 mm in diam., abruptly
conical or hemispherical, then expanding to conico-
convex, convex, and plano-convex with slightly de-
pressed center, with deexed then straight margin,
hygrophanous, translucently striate up to the cent-
er, minutely squamulose, more densely in center,
typically pallid; pale beige, yellowish brown, or pale
grey with a darker central spot (4A2–3, 5B3–4, 5C3–
4), initially often almost whitish towards margin.
L a m e l l a e moderately distant, adnate-emargin-
ate, decurrent with a tooth or arcuate, white, be-
coming pale or pinkish, with a concolorous entire
edge. – S t i p e 20–70 × 1.5–3 mm, cylindrical or
slightly broadened towards the base, sometimes
compressed with longitudinal groove, smooth, pol-
ished, either entirely pale beige or greyish, or with
distinct greyish blue tinge, then often discoloring to
Fig. 12. Entoloma timidum, micromorphological features
drawn from the holotype (LE312480). A. Basidiospores. B. Ba-
sidium. C. Cheilocystidia. D. Pileipellis. Scale bars 10 μm, del.
O.V. Morozova.
304 Sydowia 73 (2021)
Dima et al.: FUSE 7
beige with age, especially in upper part, white
tomentose at the base. – C o n t e x t white, under
the surface beige. – S m el l indistinct, taste not re-
ported. – B a s i d i o s p o r e s 8–11 × 5.5–8 μm, aver-
age 9.5 × 6.5 μm, Q=1.2–1.6, Qav=1.4, heterodiametri-
cal, with 5–7 angles in side-view. – B as i d i a 30–39
× 9–10.5 μm, 4-spored, narrowly clavate to clavate,
clampless. – L a m e l l a e d g e fertile. – C h e i l o -
cystidia absent. Hymenophoral trama
regular, made up of 4–15 μm wide, cylindrical hy-
phae. – P il e i p el l i s a cutis of cylindrical 5–10 μm
wide hyphae, with transition in the center to a
trichoderm of chains of inated cells with cylindri-
cal, inated or clavate terminal elements, 30–75 ×
10–17 μm, and yellow-brown intracellular pigment
in KOH. – C a u l o cy s t i di a as ascending bundled
cylindrical to slightly inated cells, 23–28 × 5–7 μm.
Clamp connections absent.
E t y m o l o g y. – From Latin, timidus meaning
modesty, referring to the tender habitus and expres-
sionless colors of basidiomata, especially the grey-
ish-beige stem.
Habitat and distribution. In small
groups in semi-natural grasslands and margins of
rich to calcareous Tilia–Fraxinus–Corylus and Pi-
cea forests. Known from Norway and Russia (Euro-
pean part and Caucasus).
Additional material examined. RUSSIA.
Tver region, Staritsa district, vicinities of the Krutitsy village,
bank of Volga River, 56°34’22.49”N, 33°36’25.48”E, in grass-
land, 10 September 2015, leg. O. Morozova (LE 311800); Ka-
rachaevo-Cherkesia Republic, Teberda Nature Reserve, near
Teberda town, 43°26’19.92”N, 41°43’52.20”E, in grassland in
broad-leaved forest, 1350 m a.s.l., 6 August 2009, leg. O. Mo-
rozova (LE 312481); Ibid., foot of a waterfall on the western
slope of Mount Kelbashi, 56°52’03.71”N, 35°54’56.47”E, in
grassland, 1450 m a.s.l., 25 August 2012, leg. O. Morozova (LE
312482). – NORWAY. Nordland, Grane. Holmvassdalen Nature
Reserve, 65°31’57.59”N, 13°43’33.64”E, tall herb vegetation in
calcareous spruce forest, 14 August 2004, leg. J. Lorås, JL 15-
14; Nordland, Grane, Danielåsen Nature Reserve,
65°33’11.42”N, 13°38’26.33”E, tall herb vegetation in calcare-
ous spruce forest, 9 August 2016, leg. J. Erlandsen & M. Eidis-
sen, JL 13–16; Oppland, Lunner, S. Oppdalen, Amundrud,
60°17’39.84”N, 10°40’48.72”E, ~450 m a.s.l., on calcareous,
semi-natural (mown) meadow, 19 August 2014, leg. T.E. Bran-
drud (O-F-75148); Troms, Senja, Lenvik, Senja familiepark,
Gressmyrbotn, 69°17’07.08”N, 17°53’43.44”E, 80 m a.s.l., in
base rich semi-natural pasture, 31 August 2011, leg. J.B. Jordal
(O-F-252355); Telemark, Kragerø, Grønnåsliane,
58°52’19.39”N, 9°19’48.47”E, 8 August 2020, rich TiliaFraxi-
nus forest, near ditch, leg. T.E. Brandrud, TEB 73-20; Trønde-
lag, Steinkjer, Egge church, 64°01’26.04”N, 11°28’26.76”E,
rich lawn of churchyard, 23 August 2016, leg. Ø. Weholt, OW
E10-16.
Notes. Entoloma timidum is characterized
by the rather anonymous pale yellowish-brown to
beige pileus, greyish-beige polished stipe, rather
small basidiospores with 5–7 distinct angles, and
the lack of cheilocystidia. Some collections have a
bluish-grey tinge in the stipe, particularly when
young, at which time the species can be confused
with the phylogenetically distant E. lividocyanu-
lum (Kühner) Noordel. It is very likely that this spe-
cies has been described by Ludwig (2007) as E. mu-
tabilipes f. acystidiatum E. Ludw. Entoloma muta-
bilipes Noordel. & Liiv differs by the (partly) sterile
lamella edge and slightly smaller basidiospores
(Noordeloos 1992). Entoloma timidum might also in
certain stages resemble E. ochromicaceum Noordel.
& Liiv, but this species has a sterile lamellar edge
with well-developed cheilocystidia.
Authors: B. Dima, O.V. Morozova, M.E.
Noordeloos, T.E. Brandrud, J.B. Jordal, G.M. Jansen
& E. Larsson
Basidiomycota, Agaricomycetes, Agaricales, Ento-
lomataceae
Entoloma violaceoserrulatum
Noordel., Brandrud,
O.V. Morozova & Dima, sp. nov. – Figs. 5h, 13
MycoBank no.: MB 839638
= Entoloma violaceoserrulatum Noordel. in En-
toloma s.l., Fungi Europaei vol. 5a: 1038 (2004),
nom. inval., Art. 40.7 (Melbourne).
H o l o t y p us . – FINLAND. Varsinais-Suomi, Archipelago
National Park, Berghamn, Island of Böskar, Lönöholmen,
60°03’00.07”N, 21°46’56.97”E, 25 August 1993, leg. J. Vauras,
JV 8329F (L; holotype).
Description. – Basidiomata medium-
sized, collybioid. – P i l e u s 12–50 mm in diam., con-
vex to plano-convex with slightly depressed center,
with subinvolute then deexed margin, not hy-
grophanous, not translucently striate, entirely velu-
tinous or tomentose, then minutely squamulose, ini-
tially dark violet to almost violaceous black, the
bluish tinges very soon fades, and then almost black-
ish and soon discoloring to grey-brown. – L am el -
l a e L = 30–40, l = 3–5, moderately crowded, adnate-
emarginate with decurrent tooth, ventricose, ini-
tially pure white, then pink, lamellae edge mbriate
to serrulate, usually concolorous, but sometimes
developing bluish black, spots. – S t i p e 20–45 ×
2–5 mm, cylindrical, initially silvery violaceous-grey,
then discoloring more brownish grey, strongly bril-
lose striate lengthwise with darker brils and often
also minute squamules towards the apex, base with
abundant white mycelium. – C o n te x t whitish to
pale grey. – S m e l l indistinct (“leptonioid”), taste
not recorded. – B a s i di o s p o r e s 8.5–11 × 6.0–7.5
μm, average 8.7 × 7.2 μm, Q = 1.25–1.45, Qav = 1.35,
Sydowia 73 (2021) 305
Dima et al.: FUSE 7
5–6-angled in side-view, rather regularly, relatively
thin-walled. – Ba s i d i a 28–40 × 4.0–10 μm,
4-spored, clampless. – L a m e l la e d g e sterile, of
serrulatum-type, with dense clusters of cylindrical
cheilocystidia (pseudocystidia), 20–50 × 2.5–7.0 μm,
thin-walled, lled with brilliant, diffractive-hyaline
granules, very rarely with bluish pigment. – H y m e -
n o p h o r a l t r a m a regular, made up of cylindrical
elements; brilliant granules very abundant in trama
and in the lamellar edge. – P i l e i p e l l i s a tricho-
derm of very broadly clavate to vesiculose elements,
40–100 × 10–30 μm, with brown, intracellular pig-
ment. – S t i p i t i p e l l i s a cutis of loosely arranged,
cylindrical hyphae, 3.0–10 μm wide, without caulo-
cystidia. – Cl a m p c on n e ct i o n s not seen with
certainty.
E t y m o l o g y The name refers to the viola-
ceous, serrulate lamellar edge, seen in the type ma-
terial. Based on study of more material, this, how-
ever, appears to be a rare feature of the species.
Habitat and distribution. Mainly re-
corded in calcareous grasslands, both natural and
semi-natural (grazed or mown), but sometimes also in
rather open, calcareous Picea–Pinus–Betula forests.
This species seems to be northern, widely distributed
in calcareous districts of Norway (north to Troms),
and recorded also in Finland and Sweden. One recent
record from the Netherlands was from a semi-natural
grassland on calcareous, loamy-sandy soil.
Additional material examined. Ibid. (TUR;
isotype). Sequences ex-isoype: MF476913 (ITS). – NORWAY,
Telemark, Porsgrunn, Åsstranda brygge, 59°05’36.96”N,
9°38’49.56”E, near-shore, open, dry, calcareous grassland, 10
September, leg. T.E. Brandrud & B. Dima, TEB 339-15 (O-F-
260353); Oppland, Lunner, S. Oppdalen, Amundrud nordre,
60°17’39.84”N, 10°40’48.72”E, calcareous hayeld, 19 August
2014, leg. T.E. Brandrud (O-F-75151); Ibid., 9 August 2019, leg.
T.E. Brandrud & B. Dima, TEB 128-19 (O). – SWEDEN. Jämt-
land, Östersund, Torvalla, Ängsmoen västra, 63°08’45.24”N,
14°45’19.08”E, calcareous semi-natural formerly grazed/
mown grassland, 28 August 2016, leg. T.E. Brandrud,
LE312676. – THE NETHERLANDS. Gelderland Province,
Staverden, Leemputten, 52°02’40.74”N, 6°40’12.70”E, 22 Oc-
tober 2019, leg. F. & R. Salzmann (L0607704).
N o t e s . – Noordeloos (2004) indicated a holo-
type specimen for this species in both L and TUR.
This is against the rules of the International Code of
Nomenclature for algae, fungi, and plants (Turland
et al. 2018). Therefore this nomen invalidum is here
corrected, with a holotype specimen in L, an isotype
specimen in TUR, and an emended description. En-
toloma violaceoserrulatum phylogenetically be-
longs to the /serrulatum clade. Within this group, it
has some aberrant characters that make it quite dif-
ferent from related taxa; the very dark violaceous
pileus when young, which is entirely tomentose-
squamulose, not translucently striate; the initially
pure white lamellae; and the stipe with the same
shade of color as the pileus but lighter, and distinct-
ly brillose striate. Other E. corvinum (Kühner)
Noordel. lookalikes have a completely different
structure of the lamellar edge, without brilliant
granules. Entoloma mougeotii (Fr.) Hesler has some
resemblance, but it differs by the tender, grey-viol-
aceous color, the often brown lamellar edge, with-
out brilliant granules, and slightly larger basidio-
spores.
Authors: B. Dima, O.V. Morozova, M.E.
Noordeloos & T.E. Brandrud
Ascomycota, Laboulbeniomycetes, Laboulbeniales,
Laboulbeniaceae
Laboulbenia amblystomi
W. Rossi & Santam., sp.
nov. Fig. 14
MycoBank no.: MB 839568
Fig. 13. Entoloma violaceoserrulatum, micromorphological
features drawn from collection LE312676. A. Basidiospores.
B. Cheilocystidia. C. Pileipellis. Scale bars 10 μm, del. O. V.
Morozova.
306 Sydowia 73 (2021)
Dima et al.: FUSE 7
H o l o t y p u s . – THAILAND. Lamphun Province, Ban
San Ka Yom, on elytra of Amblystomus femoralis (Motschul-
sky, 1858) (Coleoptera, Carabidae, Harpalinae), light trap, 30
October 2016, leg. W. Rossi, slide WR4185 (holotype at FI).
D e s c r i p t i o n . – Thallus brownish gray, with
paler cell V and lower portion of cell I, and with
the surface of the cells of the receptacle nely dot-
ted. – C e l l I about twice longer than broad,
gradually enlarging upwards. – C e l l I I slightly
longer and distinctly broader than cell I, divided
from cell III and cell VI by oblique septa, the latter
of which is usually longer and more concave. –
C e l l I I I slightly broader than long. – C e l l I V
similar to cell III. – C e l l V small and wedge-
shaped. – O u t e r a p p e n d a g e simple and very
long when unbroken, consisting of a basal cell iso-
diametric in outline, followed by gradually longer
cells that start tapering from the fourth cell, and
with the third and fourth cells usually darker; the
upper, outer portion of the third cell shows a very
small spinous process, the remnant of the ascospore
apex; additional branches can be produced if the
main branch has been broken (Fig. 14C). – I n n e r
a p p e n d a g e with the basal cell much smaller
than the outer one, bearing two short cells, each
apically producing a short, simple branch and two
grayish antheridia, the latter replaced by short
Fig. 14. Laboulbenia amblystomi. A. Mature thallus from elytra of Amblystomus notabilis, slide WR4173. B. Mature thallus from
elytra of A. femoralis, slide WR4185 (holotype). C. Thallus from the abdominal tip of A. similis, slide WR4229. D. Detail of im-
mature thallus, slide WR4229. Arrows in all gures point to the remains of the spore apex. Scale bars A–C 50 μm, D 25 μm.
Sydowia 73 (2021) 307
Dima et al.: FUSE 7
branchlets in older thalli. – Ce l l V I from isodia-
metric to distinctly broader than long. – P e r i -
t h e c i u m oblong, about twice longer than broad,
with a broad, blackish tip not well distinguished
except by its color, the hyaline apex obliquely di-
rected outwards consisting of rounded, hardly pro-
truding, hyaline lips. – Length from foot to perithe-
cial apex 115–120 μm; length from foot to tip of
longest appendage 295 μm; perithecium 49–50 ×
22–24 μm.
E t y m o l o g y. – Referring to the host genus,
Amblystomus.
Hosts and distribution. On species of
Amblystomus (Coleoptera, Carabidae, Harpalinae)
in Italy, Spain, and Thailand.
Additional material examined. THAILAND.
Lamphun Province, Ban Lam Chan, Tambon Pratu Pa, on
elytra of Amblystomus notabilis Landin, 1955 (Coleoptera,
Carabidae, Harpalinae), light trap, 23 October 2016, leg. W.
Rossi, slide WR4173 (paratype at FI); Lamphun Province,
Mueang Lamphun District, harvested rice elds near Umong,
on various body parts of A. similis Landin, 1955, 13 February
2017, leg. M. Bernardi & W. Rossi, slide WR4229 (paratype at
FI). ITALY, Sassari Province, near Romana, on A. sardous Bau-
di di Selve, 1864, 5 December 1975, leg. G. Franzini, slide
WR1115 (paratype at FI); Ibid., slides WR1116, WR1117,
WR1118 (paratypes at FI); Oristano Province, near Abbasanta,
on A. melonii Wrase & Magrini, 2012, 6 January 2000, leg. P.
Magrini, slide WR4274 (paratype at FI); Ibid., slide WR4275
(paratype at FI); Roma Province, near Fiumicino, on A. algiri-
nus Reitter, 1887, 22 December 1975, leg. G. Franzini, slide
WR1119 (paratype at FI). SPAIN, Catalonia, Barcelona, Gual-
ba, Gualba de Baix, riera de Gualba, on A. metallescens (De-
jean, 1829), 17 April 1992, leg. S. Santamaria, slide SS1459
(paratype at BCB).
N o t e s . – The description is based on the thalli
occurring on A. femoralis (holotype). The thalli
found on the pro- and mesothorax of A. similis are
larger (up to 250 μm), but this is not unusual when
thalli of Laboulbenia grow on these parts of the
body; the thalli found on the elytra of the same in-
sect are identical with the parasites observed on the
elytra of A. femoralis and A. notabilis. The parasites
occurring on European species of Amblystomus (A.
algirinus, A. melonii, A. metallescens, and A. sar-
dous) have more or less the same length of the type
series (120–140 μm), but have a stockier habitus.
This is caused by the shape of the suprabasal cell,
which is broader than long in the European thalli
vs. longer than broad in the Asian ones. All other
features are practically identical, including obvi-
ously the remains of the spore apex. Our caution is
also due to the lack of information from the huge
geographical gap between Europe and Thailand.
Any doubt can be eliminated only by the study of
the fungi occurring on the species of Amblystomus
from this large intermediate area.
At rst sight, new species displays no striking
feature. However, the remains of the ascospore apex
have never been observed before in any of the more
than 650 species of Laboulbenia described so far.
This feature was also not detected in the detailed
studies on the development of Laboulbenia pub-
lished by Tavares (1985) and De Kesel (1989).
Laboulbenia amblystomi seems to be related to
L. polyphaga, but is distinguished from it by several
features: in the new species the perithecium is freer
from the receptacle, the perithecial tip is broader,
the antheridia are darker and fewer in number, the
appendage is made of much shorter cells, of which
the 3rd and 4th are darker. The fungus reported as
L. polyphaga Thaxt. on Amblystomus sp. from Bali
Island (Indonesia) most likely belongs to L. amb-
lystomi; the photo accompanying this record (Sugi-
yama & Majewski 1985: Fig. 5) leave very little
doubts.
Authors: S. Santamaria & W. Rossi
Basidiomycota, Agaricomycetes, Boletales, Suil-
laceae
Suillus quercinus
Sarwar, Naseer & Khalid, sp. nov.
– Figs. 15–17
MycoBank no.: MB 835423
H o l o t y p e . – PAKISTAN. Khyber Pakhtunkhwa Prov-
ince, Swat District, Toa Valley, 2100 m a.s.l., under Quercus
incana (Fagales, Fagaceae), 1 August 2018, leg. A. Naseer &
A.N. Khalid, ANK356 (LAH36421; holotype). Sequences ex-
holotype: MT361745 (ITS).
Description. – Basidiomata medi-
um-sized, length 5.5–10 cm. – P i l eu s 4.0–9.5 cm in
diam., convex when young, broadly convex or at
with age; margins smooth, acute, entire, regular. –
P i l e us s u rf a c e dry, not viscid or slightly viscid
only when wet, brownish-yellow to yellow, discolor-
ing to pale yellow with age, brownish-red to red-
dish-brown appressed brillose throughout the
pileal surface, which becomes prominent with age.
P i l e u s c on t e x t thick in the center and thin
towards margins, soft, pale yellow to yellowish in
age, unchanging when exposed. – Ta s t e a n d
odor not recorded. Hymenium tubes up to
10 mm deep, adnate to sub-decurrent, fully peeling,
yellow when young, yellowish-brown with age,
slowly changing to dark brown when exposed. –
P o r e s irregular, 1–3 mm broad, concolorous with
the tubes. – S t i p e central, equal, curved, length
5.0–9.5 cm, 1.3–1.5 cm in diam., concolorous with
the tubes but white at the base; brown glandular
dots all over the surface that are prominent towards
the apex, annulus absent, basal mycelia white. –
308 Sydowia 73 (2021)
Dima et al.: FUSE 7
Fig. 15. Phylogeny of Suillus reconstructed from an ITS dataset. The topology is the result of ML inference performed with
RAxML. For each node, MLBS (if >50) is presented above/below the branch leading to that node. Labels consist of species names
followed by GenBank accession numbers. • indicates the new species.
Sydowia 73 (2021) 309
Dima et al.: FUSE 7
Fig. 16. Suillus quercinus. A, C, D, F. Fresh basidiomata in the eld showing distinct pileal features. B, E, G. Hymenium and stipe
surface. Scale bars A–C 1.5 cm, D–E 1.2 cm, F–G 1 cm.
310 Sydowia 73 (2021)
Dima et al.: FUSE 7
S t i p e c o n t e x t solid, whitish to slightly pale
yellow with no color change when exposed. – B a -
s i d i o s p o r e s (8.2–)8.9–10.9(–11.2) × (3.2–) 3.5–
4.1(–4.3) μm, average 9.8 × 3.9 μm, Q=2.2–2.9,
Qav = 2.5, ellipsoid or oblong, yellowish brown in
5 % KOH, thick-walled. – B a s i d i a 19.9–39.3 ×
6.3–9.9 μm, average 28.4 × 8.0 μm, narrowly utri-
form to clavate, four-spored, hyaline to brown in
5 % KOH, thin-walled. – P l eu ro c y s ti d i a 47.7–
85.5 × 4.9–6.7 μm, average 62.4 × 5.9 μm, cylindrical
to sub-clavate, pseudocystidium, hyaline to brown
in 5% KOH, thin-walled. – C h e i l o c y s t i d i a
11.9–25.7 × 3.9–9.1 μm, average 16.4 × 6.5 μm,
clavate to mucronate, hyaline in 5 % KOH, thin-
walled. – Pi l e i pe l li s 6.1–19.7 μm, average width
11.4 μm, septate, branched, hyaline in 5 % KOH,
thin-walled. – St i p i t i p e l l i s 5.9–17.6 μm, aver-
age width 9.9 μm, septate, branched, hyaline in 5 %
KOH, clamp connections present, thick-walled.
E t y m o l o g y. – Referring to the Quercus for-
ests in which this species was found, alluding to a
strict mycorrhizal association.
Habitat and distribution. Thus far
only reported from Pakistan in temperate moist for-
ests of Quercus incana.
Additional material examined. PAKISTAN.
Khyber Pakhtunkhwa Province, Shangla/ Swat boundary, Toa,
Solitary, under Quercus incana rhizosphere, 1 August 2019,
leg. M. Usman, A. Naseer & Ab.N. Khalid, ANK328 (LAH36422);
Khanspur, Ayubia, in forest of Q. incana, 24 July 2011, leg. S.
Sarwar & A.N. Khalid, B102 (LAH240711).
Notes. Suillus is an ectomycorrhizal genus of
currently some 60 described species (Wijayawardene
et al. 2020), characterized by eshy, pored basidio-
mata, sometimes a ring, wide pore openings, and
smooth spores (Bessette et al. 2000, Kuo 2004). Most
of the species of this genus are found in association
with Pinaceae and they are widely distributed with
reports from all continents, including Asia (Kirk et
al. 2008, Sarwar et al. 2018). In their preliminary
checklist of the Boletales in Pakistan, Sarwar &
Khalid (2012) reported 9 species of Suillus: S. bovi-
nus (L.) Roussel, S. collinitus (Fr.) Kuntze, S. granu-
latus (L.) Roussel, S. grevillei (Klotzsch) Singer, S.
luteus (L.) Roussel, S. placidus (Bonord.) Singer, S.
sibiricus (Singer) Singer, S. tomentosus Singer, and
S. viscidus (L.) Roussel. Since then, S. brevipes
(Peck) Kuntze (Sarwar et al. 2011), S. avidus (Fr.) J.
Presl (Sarwar et al. 2012), S. marginielevatus S. Sar-
war, Khalid & Dentinger (Sarwar et al. 2015), and S.
himalayensis B. Verma & M.S. Reddy (Sarwar et al.
2018) were added to the funga of Pakistan. With S.
quercinus, 15 species in the genus are known from
the country.
Our ITS dataset for the phylogenetic analysis in-
cluded 43 sequences and 670 characters, of which
487 were constant and 142 were parsimony-inform-
ative. The sequences from the Pakistani collections
formed a well-supported (MLBS=73) monophyletic
clade within the S. sibiricus group (Fig. 15). Mor-
phologically, S. quercinus is similar to S. sibiricus
due to its yellowish pileus surface and brownish
squamules. However, S. sibiricus has a white pileal
veil, which is absent in S. quercinus (Sarwar et al.
2018). Suillus himalayensis is also a phylogeneti-
cally closely related species. The new species can be
distinguished from S. himalayensis due to its thin
stipe, and prominent brillose pileus surface. In ad-
dition, the pileus context of S. himalayensis chang-
es color after bruising (Verma & Reddy 2014a); this
is not the case for S. quercinus. Ecologically, S.
quercinus is likely associated with Quercus inca-
na—as it was only observed in oak forests. Both S.
himalayensis and S. sibiricus are mostly associated
with conifers (Sarwar & Kahlid 2014; Sarwar et al.
Fig. 17. Suillus quercinus, line drawings of micromorphologi-
cal structures. A. Basidia. B. Basidipospores. C. Cheliocystid-
ia. D. Pleurocystidia. E. Pileipellis. F. Stipitepellis. Scale bars
A, C 3 μm; B 4.5 μm; D 4 μm; E–F 9.5 μm, del. M. Usman.
Sydowia 73 (2021) 311
Dima et al.: FUSE 7
2015, 2018). Some other Suillus species have been
found in association with oak trees but these are
not phylogenetically closely related with S. querci-
nus and mostly they were reported from mixed for-
ests so their mycorrhizal associations are as yet un-
certain (Sarwar et al. 2011, 2018; Sarwar & Khalid
2012).
Authors: A. Naseer, S. Sarwar, M. Usman & A.N.
Khalid
Interesting taxonomical notes, new hosts, and
geographical records
Basidiomycota, Agaricomycetes, Agaricales, Crepi-
dotaceae
Crepidotus malachioides
Consiglio, Prydiuk &
Setti, Il genere Crepidotus in Europa, p. 303 (2008).
– Figs. 18, 19
M a te r i al e xa m i n e d . – ITALY. Friuli Venezia Giulia,
Farra d’Isonzo, Isonzo (Soca) river, on trunks of broadleaved
trees on the ground, 13 June 2019, leg. G. Ferisin (GDOR5069).
Description. – Basidiomata gregari-
ous in large groups with reduced, lateral stem. – P i-
l e us 4–15 × 4–14 mm, convex to slightly applanate,
spathuliform, reniform, rounded abelliform to
mostly abelliform, hygrophanous, margin inexed
to straight, surface initially pubescent, later gla-
brous, generally with dense white villosity near the
point of attachment, from whitish to pale orange in
early stage, becoming ochraceous to light brown
when matured. – St ip e up to 3 mm, only visible in
very young stage, whitish, slightly pubescent.
C o n t e x t up to 1 mm thick, whitish to orange-
white, taste not recorded, smell indistinct. – La -
m el la e L = 5–20, l = 3–5, up to 2 mm wide, ventri-
cose, adnexed, pale cream when young, then ochra-
ceous orange to light brown when matured, edge
mbriate and paler than surface. – B a s i d i o -
s p o r es (4.7–)4.8–5.5(–5.6) × (4.4–)4.5–5.4(–5.5) μm,
Q = (1.00–)1.00–1.11(–1.13), globose to subglobose,
yellowish in KOH, generally with one large vacuole,
moderately echinulate. – B a s i d i a 19–30 × 6–7 μm,
4-sterigmate, clavate. – C h e i l o c y s t i d i a (20–
)25–35(–40) × (6–)8–10(–11) μm, from clavate to cy-
lindrical or exuose with subcapitate terminal part,
hyaline, thin-walled. – P i l e i p e l l i s a hymeniderm
with pileocystidia [(20–)21–30(–44) × (4–)4–7(–9) μm]
similar to cheilocystidia, non-gelatinized, hyaline.
– Tr a m a composed of parallel hyphae up to 8 μm
wide, hyaline to yellowish in KOH, thin-walled. –
Clamp connections present in all parts.
Habitat and distribution. On hard-
wood and conifer. Described from Ukraine, also re-
ported from some other European countries (Aus-
tria, Italy, Slovakia) and in China.
Notes. Crepidotus malachioides Consiglio,
Prydiuk & Setti belongs to section Sphaerula sensu
Hesler & Smith (1965) due to its globose to subglo-
bose basidiospores. This species is mainly charac-
terized by its small-sized basidiomata with whitish
to light brown tinge, globose to subglobose basidi-
ospores with “spore wall formed of rod-shaped
warts with truncate apex” under SEM (Jancˇovicˇová
et al. 2014) that appear as being nely echinulate
under the light microscope, and a hymenidermic
pileipellis with capitate pileocystidia similar to
cheilocstidia that are sometimes covered with mu-
cus. Our Italian collection (voucher GDOR5069) has
smaller basidiospores compared to the type mate-
rial [(4.7–)4.8–5.5(–5.6) × (4.4–)4.5–5.4(–5.5) μm vs.
5.2–6.2 × 5.1–5.8 μm] and Austrian and Slovakian
collections (5.1–5.9 × 4.7–5.6 μm); and cheilocystia
are shorter than those of the type (32.1–43.3 × 8.1–
11.1 μm) and Austrian and Slovakian collections
(33.9–47.2 × 7.5–10.1 μm) (Jancˇovicˇová et al. 2014).
In our observations, no mucus was found on cheilo-
cystia or pileocystidia, highlighting that this char-
acter is not constant among collections of this spe-
cies.
The Italian ITS sequence of C. malachioides
shares 100% (636/636 bp) identity with the ex-epi-
type sequence of C. malachioides (GenBank acces-
sion no. NR_132047). In the ML tree (Fig. 18), four
sequences of C. malachioides from Europe and a
sequence named “uncultured soil fungus” from Chi-
na clustered in a strongly supported clade
(MLBS = 87), highlighting the fact that C. mala-
chioides has a thus far Eurasian distribution. Eura-
sian sequences of C. malachioides are retrieved as a
sister clade (MLBS = 98) of Crepidotus cf. mala-
chioides from the USA and Argentina. In our ITS
analysis, the average distance between the Eurasian
and American clades is 3.7 %; and further studies
are needed to evaluate whether or not collections
from North and South America represent C. mala-
chioides.
Authors: F. Dovana & G. Ferisin
Ascomycota, Laboulbeniomycetes, Herpomycetales,
Herpomycetaceae
Herpomyces periplanetae
Thaxt., Proc. Am. Acad.
Arts Sci. 38(2): 13 (1902). – Fig. 20
M a t e r i a l e x a m i n e d . – BÉNIN. Atacora Province,
Natitingou, Auberge de la Montagne, bedroom, 10°12’40.80”N,
1°26’16.38”E, on antennae of Periplaneta americana Lin-
naeus, 1758 (Blattodea, Blattidae, Blattinae), ADK2088 [host
312 Sydowia 73 (2021)
Dima et al.: FUSE 7
label], 8 September 1997, leg. A. De Kesel, slide
BR5020195036610V; Atlantique Province, Calavi, house of
N.S. Yorou, near kitchen, on antennae of P. americana,
ADK6351 [host label], 3 June 2018, leg. A. De Kesel, slide
BR5020195041478V; Atlantique Province, Cotonou, Cadje-
houn, Hotel Chant d’Oiseau, corridor near kitchen,
6°21’25.60”N, 2°24’14.72”E, on antennae of P. americana,
ADK6485 [host label], 17 September 2019, leg. A. De Kesel,
slide BR5020195035583V. – DEMOCRATIC REPUBLIC OF
THE CONGO. Haut-Katanga Province, Lubumbashi, Univer-
sity guesthouse at Boulevard M’siri, on antennae of a nymph
of P. americana, ADK6326 [host label], 3 February 2018, leg. A.
De Kesel, slide BR5020195040440V. – TOGO. Maritime Region,
Lomé, Maison Guelly, city garden, on antennae of P. america-
na, ADK4870 [host label], 17 November 2010, leg. A. Guelly &
A. De Kesel, slide BR5020195039703V; Ibid., ADK4909 [host
label], 12 June 2011, leg. A. De Kesel, slides BR5020195044561V,
BR5020195043533V, BR5020195042505V; Centrale Region,
Fazao Malfakassa National Park, kitchen of abandoned Hotel
de Fazao, 8°41’41.08”N, 0°46’31.40”E, on antennae of P. ameri-
cana, ADK4943 [host label], 17 June 2011, leg. A. De Kesel,
slide BR5020195038676V, reported as Herpomyces in De Ke-
sel et al. (2011).
Hosts and distribution. Known from
Periplaneta spp. in North, Central, and South
American, Europe, Africa, and Asia (Thaxter 1908,
1931; Wang et al. 2016). On the African continent
thus far reported from Algeria (Maire 1916), Mo-
rocco (González Fragoso 1916), and Somalia
(Spegazzini 1915).
Notes. Herpomyces is the only genus in the re-
cently established order Herpomycetales. Both Her-
pomycetales and Laboulbeniales include the “thal-
Fig. 18. Phylogeny of Crepidotus reconstructed from an ITS dataset. The topology is the result of ML inference performed with
RAxML. For each node, MLBS (if 70) is presented above the branch leading to that node. Newly generated sequence highlighted
in bold.
Sydowia 73 (2021) 313
Dima et al.: FUSE 7
Fig. 19. Crepidotus malachioides, collection GDOR5069. A. Basidiospores. B. Basidiomata. C–D. Pileipellis. E–F. Cheilocystidia.
Photos: G. Ferisin. Scale bars 10 μm.
lus-forming” representatives of class Laboulbenio-
mycetes (Haelewaters et al. 2019, 2021). Both orders
produce 3-dimensional thalli resulting from deter-
minate mitotic divisions from a single, bicellular
ascospore (Blackwell et al. 2020). However, com-
pared to Herpomycetales with its single genus and
27 described species (Gutierrez et al. 2020), Laboul-
beniales are much more diverse with 2,325 described
species in 145 genera (Kirk 2019, Haelewaters et al.
2020). Herpomyces periplanetae is a widespread
314 Sydowia 73 (2021)
Dima et al.: FUSE 7
species, with reports in all continents except Oce-
ania. Our records of H. periplanetae from Bénin, D.R.
Congo, and Togo are the rst African ones in over a
hundred years. One of us (A. De Kesel) also observed
H. periplanetae on P. americana in a basement at the
University of Nairobi, Kenya in August 1998 but we
were unable to trace back the slides left at the uni-
versity. The whole African continent is largely unex-
plored for both Herpomycetales and Laboulbeni-
ales. Efforts are being undertaken to ll this distri-
butional knowledge gap, with eldwork planned in
Bénin and Mozambique.
Our three-locus dataset consisted of 33 Herpo-
myces isolates, representing 9 species, and 2,251
characters (SSU: 822, ITS: 603, LSU: 826) of which
599 were parsimony-informative (SSU: 67, ITS: 366,
LSU: 155). The ML analysis resulted in a H. peri-
planetae clade (MLBS=100) consisting of 10 iso-
lates—all removed from P. americana collected in
different localities: Massachusetts and New York in
the USA, Panama, and Bénin (Fig. 20). It may be
that the One Host One Parasite model sensu Haele-
waters & De Kesel (2020) is also applicable to spe-
cies of Herpomyces, which all form haustoria. This
hypothetical model postulates that the intertwined
interactions of Herpomycetales and haustorial La-
boulbeniales, such as species of Hesperomyces
Thaxt., with their hosts lead to an evolutionary
arms race, resulting in specialization and ultimate-
ly speciation. More data are needed to nd support
Fig. 20. Phylogeny of Herpomyces reconstructed from a three-locus dataset (SSU–ITS–LSU). The topology is the result of ML
inference performed with IQ-TREE. Pyxidiophora arvernensis and P. microspora (Pyxidiophorales) serve as outgroup taxa. For
each node, MLBS >60 is presented above or below the branch leading to that node. New isolate highlighted in bold.
Sydowia 73 (2021) 315
Dima et al.: FUSE 7
for this model. One additional nding from the tree
is that Herpomyces sp. isolate H77-1 from Parcob-
latta cf. lata (Brunner von Wattenwyl, 1865) (Blatt-
odea, Ectobiidae, Blattellinae) represents an unde-
scribed species.
Authors: A. De Kesel & D. Haelewaters
Basidiomycota, Agaricomycetes, Agaricales, Agari-
caceae
Leucoagaricus mucrocystis
(Pegler) Justo, Bizzi &
Angelini, Mycologia 113(2): 380 (2021) – Figs. 21–23
Basionym. Lepiota mucrocystis Pegler, Kew Bull.,
Addit. Ser. 9: 387 (1983).
Synonym. Lepiota besseyi H.V. Sm. & N.S. Weber,
Contr. Univ. Mich. Herb. 16: 212 (1987).
Material examined. FRENCH GUIANA. Kaw
Mountain, 4°29’58.4”N, 52°02’35.5”W, on decayed dead stand-
ing trunk of an unknown angiosperm tree, 26 December 2019,
leg. B. Dima & V. Papp, agaricoid form (ELTE:DB-FG-167-19)
and corticioid form (ELTE:DB-FG-168-19).
Description. – Pileus 2550 mm in diam.,
obtusely conical to paraboloid at rst, then plano-
convex to applanate/umbonate, velvety, brown when
young, cracking into small scales towards margin,
center remaining brown, whitish towards margin,
turning reddish brown when touched; margin not
striate. – L a m e l l a e medium spaced, lamellulae
present, ventricose, free, whitish then turning or-
ange or reddish brown when touched, becoming
dark brown on drying. – S t i p e 30–70 mm long, cy-
lindrical, with clavate to bulbous base, lower part
covered by brownish granules, whitish at the upper
part, rapidly turning orange or reddish brown when
touched; membranaceous ring present, white on the
upper surface, reddish brown on the lower surface. –
C o n t e x t thin, hollow in the stipe, white, turning
reddish brown when cut. – B a s i di o s p o r es (9.1–
)9.2–9.8(–10.2) × (6.2–)6.6–7.0(–7.2) μm, L=9.5 μm,
W=6.8 μm, Q=1.43 (n=30), ellipsoid, dextrinoid, me-
tachromatic in chresyl blue. – B a s i d i a clavate,
thin-walled, 4-spored, 21–29 × 9–12 μm. – Ch e i l o -
c y st i d i a up to 70 μm long, clavate, mucronate. –
P l e u r o c y s t i d i a similar to cheilocystida. –
P i l e i p e l l i s trichoderm, with lageniform and
mucronate terminal elements, hyphae with brown-
ish vacuolar pigmentation. – Hy p ha e o f c o rt i -
c o i d m y c e l i a cylindrical, 2.5–3.5 μm in diam.
Clamp connections present.
Habitat and distribution. Growing
both on litter and on decayed angiosperm wood. Re-
ported from the Dominican Republic, Dominica,
Hawaii, Nigeria, Trinidad, U.S. Virgin Islands, the
USA (Texas and Louisiana), and French Guiana
(this study).
N o t e s . – Our specimen (DB-FG-167-19) col-
lected in French Guiana shows similar macro- and
microscopic characteristics as Leucoagaricus mu-
crocystis by Justo et al. (2021). Based on our ITS
phylogeny (Fig. 21), the examined samples from
French Guiana (DB-FG-167-19, DB-FG-168-19)
clustered together with other La. mucrocystis (syn.
Lepiota besseyi H.V. Sm. & N.S. Weber) specimens
in a strongly supported clade (MLBS=100). Since
only ITS sequence data is available from this spe-
cies in public databases, the closest species using
the LSU sequence were Lepiota brunneoincarnata
Chodat & C. Martín (GenBank accession no.
EU416303, identities 1339/1361, 98 % similarity),
Leucoagaricus barssii (Zeller) Vellinga (GenBank
accession no. DQ911601, identities 1341/1364, 98 %
similarity), and Leucoagaricus nympharum (Kalch-
br.) Bon (GenBank accession no. EU416311, identi-
ties 1335/1358, 3 gaps, 98 % similarity).
Leucoagaricus mucrocystis was described by
Pegler (1983) from Dominica under the name Le-
piota mucrocystis Pegler. Based on morphological
and molecular examinations of the specimens col-
lected from the Dominican Republic and U.S. Vir-
gin Islands, Justo et al. (2021) transferred Lepiota
mucrocystis to the genus Leucoagaricus and
showed that this species is identical with Lepiota
besseyi, which was reported from Hawai’i, Louisi-
ana, and Texas (Smith & Weber 1987, Stallman
2019). Furthermore, they found that Lepiota bior-
nata (Berk. & Broome) Sacc. sensu Dennis (1952)
also corresponds to La. mucrocystis. The new lo-
cality of La. mucrocystis was discovered in French
Guiana, representing the rst record of this species
in South America. In the absence of detailed mor-
phological studies and available sequences, conr-
mation of the occurrence of this species in Africa
(see Adedokun et al. 2016) requires further investi-
gation.
The type of La. mucrocystis was found on a rot-
ten termite mound (Pegler 1983). Justo et al. (2021)
characterized this species as terrestrial, growing
both on litter of broadleaf woods and on wood on
the ground. In French Guiana, the examined speci-
mens were growing on a dead standing broadleaf
tree at a height of several meters (Fig. 22). Close to
the basidiomata, we also found a whitish mycelium
structure (corticioid form, Figs. 22B, E), which was
also produced by La. mucrocystis, veried by ITS
sequencing (Fig. 21). In the existing body of litera-
ture, we were unable to nd any reference to this
corticioid form.
In the Amazon, and in particular in French Gui-
ana, the distribution of the lepiotoid fungi is not
316 Sydowia 73 (2021)
Dima et al.: FUSE 7
Fig. 21. Phylogeny of Leucoagaricus reconstructed from an ITS dataset. The topology is the result of ML inference performed
with RAxML. MLBS >70 are shown at the branches. Newly generated sequences highlighted in blue.
Sydowia 73 (2021) 317
Dima et al.: FUSE 7
Fig. 22. Habitat and in-situ photo documentations of Leucoagaricus mucrocystis. A–C. Basidiomata on the substratum. D. Fresh
basidiomata, collection DB-FG-167-19. E. Corticioid form, collection DB-FG-168-19. Photos B. Dima (A, C–E), V. Papp (B).
well documented (e.g., Courtecuisse et al. 1996,
Jaouen et al. 2019). Jaouen et al. (2019) generated
and analyzed sequences of more than 5,000 speci-
mens of fungi in French Guiana, thus establishing
a vast amount of new data. Among them, the lepi-
otoid fungi were represented by only two species of
Lepiota (L. roseolamellata Dennis and L. atrodisca
Zeller), three Leucocoprinus [Lc. fragilissimus
(Ravenel ex Berk. & M.A. Curtis) Pat., Lc. birn-
baumii (Corda) Singer, and Lc. tenellus (Boud.)
Locq.] and two Leucoagaricus [La. sulphurellus
(Pegler) B.P. Akers and La. subcretaceus Bon] spe-
cies, in addition to unidentied species in all three
genera as well. Their work and our own experi-
ences and unpublished sequences show that lepi-
otoid taxa are much more diverse in French Guia-
na, and further morphological and molecular phy-
logenetic studies are needed to clarify the many
318 Sydowia 73 (2021)
Dima et al.: FUSE 7
unidentied species expected to occur in the Ama-
zonian forests.
Authors: Á. Radnóti, B. Dima & V. Papp
Basidiomycota, Ustilaginomycetes, Urocystidales,
Urocystidaceae
Melanustilospora ari
(Cooke) Denchev, Mycotaxon
87: 476 (2003). – Figs. 24, 25
Basionym. Protomyces ari Cooke, Grevillea 1(1): 7
(1872).
Synonym. Melanotaenium ari (Cooke) Lagerh., Bull.
Soc. Mycol. Fr. 15: 98 (1899).
Material examined. PAKISTAN. Khyber Pakh-
tunkhwa Province, Swat District, Kalam valley, Kargilo, 2085
m a.s.l., on leaves, petioles, and sheath of Arum jacquemontii
(Alismatales, Araceae), 2 May 2015, leg. A. Naseer & S. Hus-
sain (LAH SH-S02).
Description. Sori in leaves, petioles,
and sheaths of petioles as irregular, lead-colored
pustules, clustered in rows or scattered along mid-
rib covered by epidermis. In abaxial surface, sori
looking as aggregation of black dots, aggregations
mostly in rows. – Te l i o s p o re s (11.5–)13.5–17.0(–
20.5) × (16.0–)16.5–23.0(–26.0) μm, average 15.3 ×
19.1 μm (n = 50); mostly globose to ovoid, sometimes
irregular, ellipsoid to angular; yellowish brown to
strong brown; wall 2.0–6.8 μm thick, two-layered. –
E n d o s p or e s dark brown, evenly ca. 2 μm thick.
– E x o s p o r e s light olive brown, unevenly 0.9–
Fig. 23. Macro- and microscopic features of Leucoagaricus mucrocystis, collection DB-FG-167-19. A. Basidiomata. B. Pileocyst-
idium. C–D. Cheilocystidia. E–F. Basidia. G–I. Basidiospores. Photos B. Dima (A), Á. Radnóti (B–I).
Sydowia 73 (2021) 319
Dima et al.: FUSE 7
Fig. 24. Arum jacquemontii, the host plant for Melanustilospora ari in Pakistan, with both healthy and diseased leaves (arrow-
heads). Inset, close-up of a M. ari-infected leaf (arrows indicate places of sori).
Fig. 25. Melanustilospora ari