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Abstract and Figures

Family Cortinariaceae currently includes only one genus, Cortinarius, which is the largest Agaricales genus, with thousands of species worldwide. The species are important ectomycorrhizal fungi and form associations with many vascular plant genera from tropicals to arctic regions. Genus Cortinarius contains a lot of morphological variation, and its complexity has led many taxonomists to specialize in particular on infrageneric groups. The previous attempts to divide Cortinarius have been shown to be unnatural and the phylogenetic studies done to date have not been able to resolve the higher-level classification of the group above section level. Genomic approaches have revolutionized our view on fungal relationships and provide a way to tackle difficult groups. We used both targeted capture sequencing and shallow whole genome sequencing to produce data and to perform phylogenomic analyses of 75 single-copy genes from 19 species. In addition, a wider 5-locus analysis of 245 species, from the Northern and Southern Hemispheres, was also done. Based on our results, a classification of the family Cortinariaceae into ten genera—Cortinarius, Phlegmacium, Thaxterogaster, Calonarius, Aureonarius, Cystinarius, Volvanarius, Hygronarius, Mystinarius, and Austrocortinarius—is proposed. Seven genera, 10 subgenera, and four sections are described as new to science and five subgenera are introduced as new combinations in a new rank. In addition, 41 section names and 514 species names are combined in new genera and four lecto- and epitypes designated. The position of Stephanopus in suborder Agaricineae remains to be studied. Targeted capture sequencing is used for the first time in fungal taxonomy in Basidiomycetes. It provides a cost-efficient way to produce -omics data in species-rich groups. The -omics data was produced from fungarium specimens up to 21 years old, demonstrating the value of museum specimens in the study of the fungal tree of life. This study is the first family revision in Agaricales based on genomics data and hopefully many others will soon follow.
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Fungal Diversity
https://doi.org/10.1007/s13225-022-00499-9
Taming thebeast: arevised classification ofCortinariaceae based
ongenomic data
KareLiimatainen1 · JanT.Kim2 · LisaPokorny1,3 · PaulM.Kirk4 · BrynDentinger5 · TuulaNiskanen1,6
Received: 20 October 2021 / Accepted: 20 January 2022
© The Author(s) 2022
Abstract
Family Cortinariaceae currently includes only one genus, Cortinarius, which is the largest Agaricales genus, with thousands
of species worldwide. The species are important ectomycorrhizal fungi and form associations with many vascular plant gen-
era from tropicals to arctic regions. Genus Cortinarius contains a lot of morphological variation, and its complexity has led
many taxonomists to specialize in particular on infrageneric groups. The previous attempts to divide Cortinarius have been
shown to be unnatural and the phylogenetic studies done to date have not been able to resolve the higher-level classification
of the group above section level. Genomic approaches have revolutionized our view on fungal relationships and provide a
way to tackle difficult groups. We used both targeted capture sequencing and shallow whole genome sequencing to produce
data and to perform phylogenomic analyses of 75 single-copy genes from 19 species. In addition, a wider 5-locus analysis
of 245 species, from the Northern and Southern Hemispheres, was also done. Based on our results, a classification of the
family Cortinariaceae into ten genera—Cortinarius, Phlegmacium, Thaxterogaster, Calonarius, Aureonarius, Cystinarius,
Volvanarius, Hygronarius, Mystinarius, and Austrocortinarius—is proposed. Seven genera, 10 subgenera, and four sec-
tions are described as new to science and five subgenera are introduced as new combinations in a new rank. In addition, 41
section names and 514 species names are combined in new genera and four lecto- and epitypes designated. The position of
Stephanopus in suborder Agaricineae remains to be studied. Targeted capture sequencing is used for the first time in fungal
taxonomy in Basidiomycetes. It provides a cost-efficient way to produce -omics data in species-rich groups. The -omics data
was produced from fungarium specimens up to 21years old, demonstrating the value of museum specimens in the study
of the fungal tree of life. This study is the first family revision in Agaricales based on genomics data and hopefully many
others will soon follow.
Keywords Agaricales· Fungariomics· Fungi· HybPiper· Museomics· Targeted capture sequencing· Whole genome
sequencing
Handling Editor: Ruilin Zhao.
* Kare Liimatainen
k.liimatainen@kew.org
Jan T. Kim
j.t.kim@herts.ac.uk
Lisa Pokorny
l.pokorny@kew.org
Paul M. Kirk
p.kirk@kew.org
Bryn Dentinger
bdentinger@nhmu.utah.edu
Tuula Niskanen
tuula.niskanen@cortinarius.fi
1 Jodrell Laboratory, Royal Botanic Gardens, Kew,
SurreyTW93AB, UK
2 Department ofComputer Science, School ofPhysics,
Engineering andComputer Science, University
ofHertfordshire, Hatfield, HertfordshireAL109AB, UK
3 Present Address: Institut Botànic de Barcelona (IBB,
CSIC-Ajuntament de Barcelona), 08038Barcelona,
Catalonia, Spain
4 Biodiversity Informatics andSpatial Analysis, Jodrell
Laboratory, Royal Botanic Gardens, Kew, SurreyTW93AB,
UK
5 Natural History Museum ofUtah andSchool ofBiological
Sciences, University ofUtah, SaltLakeCity, UT, USA
6 Botanical Museum, University ofHelsinki, P.O. Box7,
00014Helsinki, Finland
Fungal Diversity
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Introduction
Genomic-level data have revolutionized our views on
fungal relationships and helped us create better phylog-
enies for previously unresolved lineages (e.g., Chang etal.
2021; Li etal. 2021). These data have been used to tackle
macroevolutionary events, e.g., mushroom morphological
evolution (Varga etal. 2019, Sánchez-García etal. 2020)
or the evolution of symbiotic traits (Miyauchi etal. 2020).
High-throughput sequencing (HTS) techniques have also
allowed genomic data to be generated from fungarium
specimens (Dentinger etal. 2016).
Fungal genomes are small, ranging from 7.67 to 720.2
Mbp/1C (Kullman etal. 2005), with an average size of ~ 63
Mbp/1C in Ascomycota (Hill etal. 2021) and ~ 50 Mbp/1C
in Basidiomycota (Mohanta and Bae 2015; Li etal. 2018),
compared to those of plants and animals, e.g., ranging 64
Mbp/1C to 140 Gbp/1C in angiosperms (Pellicer etal.
2018) and 1.6 to 6.3 Gbp/1C in mammals (Kapusta etal.
2017). Therefore, in phylogenomics studies of fungi, in
depth or shallow whole genome sequencing (WGS) have
been an affordable option to generate HTS data, with the
vast majority of the fungal genomic studies to date hav-
ing relied on this approach. However, for species-rich
groups where hundreds to thousands of samples might
be included, targeted capture sequencing provides a more
cost-effective alternative (Hale etal. 2020). Enrichment
methods have been widely used to study the systematics
of plants and animals (e.g., Johnson etal. 2019; Faircloth
2017) and they have recently been applied in fungal sys-
tematics to study lichen-forming Ascomycota families
Lobariaceae (Widhelm etal. 2019) and Parmeliaceae
(Grewe etal. 2020), as well as the Peltigeralean backbone
(Widhelm etal. 2021).
The family Cortinariaceae belongs to suborder Agari-
cineae, which contains mainly the brown and dark-spored
Agaricales with thick-walled and pigmented basidiospores
(Matheny etal. 2015; Dentinger etal. 2016). According to
the most recent phylogenetic studies, the family Cortinari-
aceae only includes one genus, Cortinarius (Pers.) Gray.
Several genera formerly placed in the Cortinariaceae,
e.g., Phaeocollybia R. Heim, Hebeloma (Fr.) P. Kumm.,
and Galerina Earle, have been moved to other families
in the Agaricineae (Matheny etal. 2015). On the other
hand, many taxa previously treated as separate genera are
currently included into Cortinarius: Bulbopodium Earle,
Cuphocybe R. Heim, Dermocybe (Fr.) Wünsche, Inoloma
(Fr.) Wünsche, Myxacium (Fr.) P. Kumm., Phlegmacium
(Fr.) Wünsche, Rapacea E. Horak, Rozites P. Karst., and
Telamonia (Fr.) Wünsche, as well as sequestrate genera
Gigasperma E. Horak, Hymenogaster Vittad. p.p., Proto-
glossum Massee, Quadrispora Bougher & Castellano, and
Thaxterogaster Singer (Peintner etal. 2001, 2002, 2004;
Garnica etal. 2005; Soop etal. 2019; Nouhra etal. 2021).
Other than Cortinarius, the only genus currently placed in
the family Cortinariaceae is the South American Stepha-
nopus M.M. Moser & E. Horak, a small genus with five
described species associated with Nothofagaceae, but it
is lacking any sequence data and thus its position in the
suborder Agaricineae remains unresolved.
As currently delimited, Cortinarius is by far the largest
genus of Agaricales, and its representatives are found from
the tropics to arctic habitats in the Northern and Southern
Hemispheres. To date, over 5000 taxa, including subspe-
cies and varieties, have been recorded in Index Fungorum
(2021). ITS sequence data exist for close to 3000 species
(UNITE 2021, using an SH threshold of 1.5%), including
both described and undescribed taxa. However, sequence
data from many regions of the world are still lacking and,
thus, many more species are in urgent need of sequencing
and/or description.
Cortinariaceae species are important ectomycorrhizal
fungi and form associations with many vascular plants,
including mainly woody species (trees and some shrubs) in
the gymnosperms (Pinaceae) and the rosid angiosperms—
orders Fabales (Fabaceae), Fagales (Betulaceae, Fagaceae,
Nothofagaceae), and Rosales (Rhamnaceae, Rosaceae), in
the so-called nitrogen fixing clade; core Malvales (Malva-
ceae, plus the Cistaceae, Dipterocarpaceae, and Sarco-
laenaceae clade); and, even, orders Malpighiales (Phyllan-
thaceae, Salicaceae) and Myrtales (Myrtaceae)—as well
as some herbaceous angiosperms, both in the monocots
(Cyperaceae and Orchidaceae) and in the caryophyllids eud-
icots (Polygonaceae) (Moser and Horak 1975; Harrington
and Mitchell 2002; Frøslev etal. 2005; Garnica etal. 2005;
Jacquemyn etal. 2010; Tedersoo etal. 2011; Harrower etal.
2015a; Thoen etal. 2019). The species of Cortinariaceae
provide access to nitrogen for vascular plants in nutrient
poor habitats, and their role in carbon cyling has been stud-
ied for northern European (Bödeker etal. 2014; Lindahl
etal. 2021) and North American (Fernandez etal. 2020)
boreal forests.
The first study based on molecular data (Høiland & Holst-
Jensen 2000) indicated that many of the traditional infrage-
neric groupings of Cortinarius were unnatural (e.g., Moser
1983; Brandrud etal. 1989; Bidaud etal. 1994). During the
following 20years many studies were conducted to gain a
better understanding of their natural relationships. Most of
these datasets were based only on sequences from the ITS
and LSU regions (Høiland & Holst-Jensen 2000; Peintner
etal. 2004; Garnica etal. 2005; Harrower etal. 2011; Lii-
matainen etal. 2014, 2020a; Stensrud etal. 2014), and only
two extensive studies of the genus also included data from
RPB1 and/or RPB2 regions (Garnica etal. 2016; Soop etal.
2019). So far, the only study to include more than a handful
Fungal Diversity
1 3
of DNA markers (ITS, nLSU, GPD, MCM7, RPB1, RPB2,
and TEF1) was conducted by Stefani etal. (2014) for the
delimitation of Australian dermocyboid Cortinarius species.
Despite these efforts, no genus-wide, revised subgeneric
classification of Cortinarius has been presented because it
has not been possible to resolve the backbone of the phy-
logeny. Existing sequence data have, however, allowed the
revision of various sections within Cortinarius, with studies
by Soop etal. (2019), Liimatainen etal. (2020a), Ammirati
etal. (2021), and Niskanen and Liimatainen (2021) present-
ing the most updated morpho-genetic, section-level classi-
fication of Cortinarius, including a total of 172 sections.
The aim of this study was to conduct the first phylog-
enomic study of family Cortinariaceae to resolve higher-
level relationships and allow for a revised genus-level clas-
sification of the group.
Materials andmethods
Molecular sampling
Sampling was designed to cover as many of the major line-
ages of Cortinariaceae as possible, based on the latest phy-
logenetic trees published for the family (Garnica etal. 2016;
Soop etal. 2019). Vouchers of 19 dried fungarium speci-
mens sampled for genomics work are deposited in the collec-
tions of the Royal Botanic Gardens, Kew, United Kingdom
(K) and/or (H) University of Helsinki, Finland.
DNA extraction andgenomic library preparation
DNA was extracted from 2 to 4mg of dried grounded
lamella with the DNeasy Plant Mini kit (Qiagen, German-
town, USA). Extracted DNA was quantified using a Quan-
tus™ fluorometer and the Quantifluor dsDNA system kit
(Promega Corporation, Madison, WI, USA). To get an esti-
mation of the average fragment size, samples were assessed
on a 2100 Bioanalyzer (Agilent Technologies, Santa Clara,
CA, USA), using the appropriate DNA chips and reagents,
or a 4200 TapeStation System (Agilent Technologies), using
the corresponding Genomic DNA ScreenTapes and reagents.
The DNA was then fragmented using an M220 Focused-
ultrasonicator™ (Covaris, Woburn, MA, USA) with varied
shearing times (30–45s) depending on the DNA fragment
size profile. The average fragment size in the specimens that
were used for WGS ranged from 680 to 745bp, and from
660 to 880bp for the specimens used for the targeted capture
sequencing.
Dual-indexed libraries for WGS were prepared using a
TruSeq® Nano DNA LT (Illumina Inc.) sample kit follow-
ing the manufacturer’s protocols. Dual-indexed libraries
for the targeted capture sequencing were prepared using
the NEBNext® Ultra™ II Library Prep kit and the NEB-
Next® Multiplex Oligos for Illumina® (Dual Index Primer
Set 1), according to the manufacturer’s protocols (New
England BioLabs, Ipswich, MA, USA), although at half
the recommended volumes. The resulting genomic librar-
ies were quantified and qualified as above (i.e. Quantus and
Bioanalyzer/TapeStation).
WGS, genome assembly, andextraction
ofsingle‑copy orthologs
For WGS, five to six libraries were pooled following Dent-
inger etal. (2016). The sequencing was performed on an
Illumina MiSeq with v3 (2 × 300bp paired-end reads) chem-
istry (Illumina, San Diego, CA, USA) at Jodrell Laboratory,
Royal Botanic Gardens, Kew.
Demultiplexed reads were quality-checked with FastQC
(Andrews 2010) before trimming with Trimmomatic v0.39
(Bolger etal. 2014) with settings: LEADING:3 TRAIL-
ING:3 SLIDINGWINDOW:4:15 MINLEN:36. The genomes
were then assembled using ABySS (Simpson etal. 2009)
with a k-mer size ranging 51–96bp, depending on the qual-
ity of the sequence data.
Next, the 208 single-copy genes identified by Dent-
inger etal. (2016) were extracted from the nine assembled
genomes using exonerate v2.2.0 (Slater and Birney 2005)
with Cortinarius glaucopus (Miyauchi etal. 2020) amino
acid (AA) sequences (for the 208 single-copy genes) as que-
ries in searches against our nine assemblies. The top-scoring
hit was retained in each case. Additionally, we included the
AA sequences of the five single-copy loci currently used in
phylogenetic studies of family Cortinariaceae, which were
not part of the 208-gene queries in the exonorate search:
RPB1 (RNA polymerase II largest subunit B220; also
RPO21; C. odorifer GenBank no. DQ083857), RPB2 (RNA
polymerase II second largest subunit B150; also RPO22;
Coprinopsis cinerea, GenBank no. XM_001829088),
MCM7 (component of the Mcm2-7 hexameric helicase
complex; also CDC47; C. basirubescens Genbank no.
JN985546), GPD (glyceraldehyde-3-phosphate dehydroge-
nase (GAPDH), isozyme 3; also TDH3; C. austrosanguineus
JX675721), and TEF1 (Translational elongation factor EF-1
alpha; also eEF1A; C. sodagnitus GenBank no. DQ061275)
to also retrieve sequences of those regions from the genome
assemblies. These regions were compared against the assem-
blies to verify that they were truly single-copy ones in our
species.
Enrichment panel probe design
Our goal was to design a 20,000-probe custom myBaits®
enrichment panel for target capture of phylogenetically-
informative, single-copy nuclear orthologs. Four out of
Fungal Diversity
1 3
nine Cortinarius species (C. victoriaensis, C. neofurvolae-
sus, C. scaurus, and C. typicus), for which most single-copy
orthologs recovered by exonorate and representing differ-
ent lineages across Cortinariaceae, were selected for probe
design. The size of the dataset exceeded the limits of the
20,000 probe enrichment panel and we therefore discarded
20 target genes with the most missing data from all four
species. The final dataset included 188 targets, from those
identified by Dentinger etal. (2016), with the addition of
the currently used loci (RPB1, RPB2, MCM7, GPD, and
TEF1), resulting in a total of 193 targets. For the probe
design, nucleotide sequences containing both intron and
exon regions were used. Based on the visual inspection of
the alignments of each target, the intron regions were gener-
ally short (< 50bp) and largely conserved within the family
making it possible to include them in the enrichment panel.
The design and production of the probes was done by Arbor
Biosciences (Ann Arbor, Michigan, USA) based on the
sequence data provided. The probes were 120 nucleotides
long and designed with ~ 2 × tiling density.
Hybridisation andtargeted capture sequencing
Dual-indexed genomic libraries were pooled and hybridised
with our custom myBaits® enrichment panel, following v3.0
manufacturer’s protocols, with the exception of pooling four
or nine libraries per hybridisation reaction and each reaction
having a total of ¼ of the recommended volume. Hybridisa-
tions were performed at 65°C for 20h in Vapo Protect Mas-
tercycler 6325 thermocycler (Eppendorf, Arlington, UK).
Captured targets were amplified with a KAPA HiFi 2 × Hot-
Start ReadyMix PCR kit (Roche, Basel, Switzerland) for 10
cycles, and the PCR products were cleaned using Agencourt
AMPure XP magnetic beads. Final products were quantified
and qualified as above. Thirteen enriched libraries were fur-
ther pooled for sequencing on an Illumina MiSeq platform
(Illumina, San Diego, CA, USA) using v2 Nano chemis-
try (2 × 250bp paired-end reads) at the Jodrell Laboratory,
Royal Botanic Gardens, Kew.
Target retrieval andsequence assembly
Demultiplexed reads were quality-checked as above before
trimming with Trimmomatic v0.39 (Bolger etal. 2014)
with settings: LEADING:20 TRAILING:20 SLIDING-
WINDOW:4:20 MINLEN:36. The HybPiper v1.3.1 (John-
son etal. 2019) pipeline was used for downstream analy-
ses. First, quality-filtered, trimmed reads were mapped to
amino acid (AA) sequences corresponding to our target
loci using BLASTx (Altschul etal. 1990). Second, paired,
mapped reads were assembled into contigs using SPAdes
v3.13.1 (Bankevich etal. 2012), using default settings with
the exception of minimum coverage, which was set to 4x.
Third, the intronate.py script was used to generate super-
contigs (scaffolded merged SPAdes contigs containing
both complete exon and intron sequences) and retrieve_
sequences.py was used to retrieve the final supercontig
sequences (of the target loci from each of our specimens)
to build the data matricess required for subsequent phylo-
genetic analyses. Finally, summary statistics (e.g., percent
of reads mapped to target) were generated using SAMtools
(Li etal. 2009) and the hybpiper_stats.py script.
Data mining anddata matrix generation
Two different data matrices were assembled for down-
stream phylogenetic analyses. The first consisted of the
single-copy orthologs from both the shallow WGS and the
targeted capture sequencing data, for a total of 19 species.
Of the original 193 single-copy orthologs targeted, 75 of
them (including RPB1, RPB2, MCM7, GPD, and TEF1),
present at least in > 50% of the species sampled and
with > 500bp of average length recovered, were selected
for further analysis. For Cortinarius crassus, the data
matrices originating from WGS versus targeted capture
sequencing were kept separate to allow direct comparison
of these two approaches.
For the second data matrix, we mined NCBI GenBank
for RPB1 sequences from Cortinariaceae species, which we
combined with 17 newly generated RPB1 sequenced from
the WGS and targeted capture sequencing data. When avail-
able, we also mined RPB2 (from 18 genomes), MCM7 (9),
GPD (12), and TEF1 (10) for these same samples. The final
data matrix included 245 species.
Multiple sequence alignment andphylogenetic
analyses
For both data matrices, all loci were individually aligned
using MAFFT v7 with iterative refinement (i.e., E-INS-i
algorithm; Katoh and Standley 2013) and, then, manually
adjusted in SeaView (Galtier etal. 1996) following the
guidelines summarized in Morrison (2006). The individual
alignments were then concatenated in Mesquite v3.2 (Mad-
dison and Maddison 2017). Phylogenetic trees were gener-
ated from the two concatenated data matrices, with model
parameter estimation partitioned by loci, using RAxML
v8.2.12 with 1000 traditional bootstrap (BS) replicates under
the GTR + Γ model (Stamatakis 2014), as advised by Young
& Gillung (2020). For the first data matrix, Crepidotus sp.
(Dentinger etal. 2016) and Hebeloma cylindrosporum
(Kohler etal. 2015) were used as outgroups. For the second
data matrix, the backbone topology inferred from the first
data matrix was used as a topological constraint.
Fungal Diversity
1 3
Data availability
The nine new Cortinariaceae genomes sequenced for the
present study are deposited in the European Nucleotide
Archive (Study ID PRJEB49625) and the raw reads resulting
from the targeted capture sequencing in the NCBI GenBank
Sequence Read Archive, SRA (BioProject PRJNA791499).
The DNA sequences used to design the enrichment panel
probes are available on Dryad (https:// doi. org/ 10. 5061/
dryad. 0p2ng f238).
Molecular results
WGS andtargeted capture sequencing performance
Summary statistics for the WGS, targeted capture sequenc-
ing, and locus mining used in the phylogenomic analysis are
presented in Tables1 and 2. There was substantial variation
in the quality of the assemblies from WGS data and, thus,
in the recovery rate of the targeted single-copy orthologs.
Anywhere from 33 to 100% of the 75 target markers chosen
for the final phylogenomics analysis were recovered and the
recovery rate was > 70% for only four out of nine specimens.
The recovery rate for the targeted capture sequencing was
far better: it was > 85% (of the 75 loci) for nine out of eleven
specimens and substantially less (35% and 45%) in only two
specimens. Reads mapped to the initial 193 targets ranged
from 10,101 to 93,312. On average, over 23,000 reads were
needed to reach > 85% coverage for the 75 loci, and over
33,000 reads were needed to reach > 95% coverage. Pooling
nine specimens in one baiting reaction generally produced
good results: in 8 out of 9 specimens > 88% of the 75 target
loci were recoved, only in one specimen the recovery rate
was low, < 35%.
Phylogenomic inference andsystematic
rearrangements
The phylogeny inferred from 75 single-copy nuclear
orthologs for 20 accessions is shown in Fig.1. The results
of the wider 5-locus analysis, containing 245 species, are
presented in Fig.2. Nodal support BS values below 85% are
considered weak, between 85 and 95% moderate, between
95 and 99% BS strong, and lastly, 100% denotes full support.
Based on the results, the division of the family Cortinari-
aceae into ten putative genera is proposed and these names
are used hereon.
In the phylogenomics tree (Fig.1), Thaxterogaster (BS
68%) is sister to a clade encompassing all other genera (BS
79%), both weakly supported. This latter clade is further
divided into a strongly supported Cortinarius (BS 95%) and
a weakly supported clade containing the remaining genera
(BS 82%). The strongly supported (BS 95%) crown of this
latter clade is composed of fully supported Aureonarius (BS
100%), Phlegmacium (BS 100%), as well as Calonarius, here
represented by just one species. Austrocortinarius, also repre-
sented by one accession, and fully supported Cystinarius (BS
100%) are in a grade leading to the aforementioned crown
clade. From the sampled genera represented by more than one
species, only one, Thaxterogaster, received a suboptimal BS
value (< 95%).
The results of the phylogenetic analysis, based on the five
most-used single-copy marker genes in Cortinarius from
245 taxa, are presented in Fig.2. All genera represented by
more than one accession in this analysis received moderate
to full support: Aureonarius (BS 100%), Austrocortinarius
(BS 100%), Calonarius (BS 100%), Cystinarius (BS 99%),
Hygronarius (BS 92%), Phlegmacium (BS 88%), Thaxtero-
gaster (BS 99%), Volvanarius (BS 100%). The only exception
was genus Cortinarius s. str. that received lower support (BS
70%). The diversity, distribution, and selected morphological
characters for the proposed genera are summarized in Table3.
The infrageneric relationships were variably resolved in dif-
ferent genera (Fig.2). In genus Calonarius, three weakly to
moderately supported subgenera were recognized: C. subgen.
Calonarius (BS 63%), C. subgen. Calochroi (BS 87%), and
C. subgen. Fulvi (BS 75%). Genus Aureonarius was divided
into two strongly to fully supported subgenera: A. subgen.
Aureonarius (BS 100%) and A. subgen. Callistei (BS 98%).
In genus Phlegmacium, four moderately to strongly supported
subgenera were recognized: P. subgen. Phlegmacium (BS
91%), P. subgen. Carbonella (BS 98%), P. subgen. Bulbopo-
dium (BS 96%), and C. subgen. Cyanicium (represented by
just one species in our analysis). The genus Cystinarius has
two subgenera: C. subgen. Crassi (BS 100%) and C. subgen.
Cystinarius (represented by one species in the analysis). The
infrageneric relationships in the genus Cortinarius remained
mostly unresolved but the following lineages with more than
one species in our analysis received moderate to full support:
C. subgen. Dermocybe (BS 93%), C. subgen. Leprocybe (BS
93%), C. subgen. Iodolentes (BS 100%), C. subgen. Telamonia
(BS 100%), C. subgen. Myxacium (BS 97%), and C. subgen.
Cortinarius (BS 100%). In the genus Thaxterogaster many
relationships remained unresolved, although the following
subgenera received weak to full support: T. subgen. Multi-
formes (BS 83%), T. subgen. Cretaces (BS 96%), T. subgen.
Thaxterogaster (BS 99%), T. subgen. Scauri (BS 92%), and T.
subgen. Riederorum (BS 100%).
Taxonomy
New and emended generic descriptions are presented below,
as well as descriptions of new subgenera and short notes on
the previously existing subgenera. The diversity, distribution
Fungal Diversity
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Table 1 Cortinariaceae specimens used for WGS and summary statistics for the genome assembly and recovery rate of the single-copy nuclear orthologs curated by Dentinger etal. (2016)
The four species used to design the baits for the Cortinariaceae enrichment panel are in bold
Genus name Species name Voucher
number (fun-
garium)
Collection
year Yield of
DNA from
extr. (ng/µl)
260/280
(1.7–1.9) 260/230
(2.0–2.2) Pool of
five or
six
Number of
reads Assembly
size Number of
contigs Max contig
length (bp) N50 Number
(percent) of
single-copy
orthologs, out
of the 75 used
Cortinarius C. bovarius TN11-191
(K(M), H) 2011 17 1.92 1.49 5 4,545,983 3.59E + 07 386,493 48,684 1935 44 (59%)
Cortinarius C. alces TN11-065
(K(M), H) 2011 29 1.96 1.68 5 5,256,187 3.72E + 07 558,926 53,036 2176 54 (72%)
Cortinarius C. glandi-
color TN15-018
(K(M)) 2015 104 1.69 0.88 6 3,479,379 3.27E + 07 748,987 47,177 1519 30 (40%)
Cortinarius C. neofurvo-
laesus TN11-113
(K(M), H) 2011 21 1.95 1.74 5 5,015,654 3.70E + 07 374,514 59,009 3335 64 (85%)
Aureonarius A. tofaceus TN10-061
(K(M), H) 2010 96 1.93 2.42 5 5,867,977 2.41E + 07 2,450,611 14,723 1550 31 (41%)
Austrocorti-
narius A. victori-
aensis K(M)162,337 1995 123 1.77 1.34 6 3,542,880 4.32E + 07 113,713 94,283 5713 74 (99%)
Calonarius C. typicus TN14-281
(K(M), H) 2014 6.7 2 1.1 6 3,746,677 3.74E + 07 178,051 160,564 5178 75 (100%)
Cystinarius C. crassus TN07-305
(K(M), H) 2007 29 1.89 1.67 5 5,018,344 2.31E + 07 403,693 22,643 1593 25 (33%)
Thaxtero-
gaster T. scaurus TN15-013
(K(M)) 2015 81 1.81 1.3 6 3,508,661 2.72E + 07 1,012,788 25,636 2090 33 (44%)
Fungal Diversity
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Table 2 Cortinariaceae specimens used in targeted capture sequencing and HypPiper summary statistics, target recovery rate inclusive
Genus name Species name Voucher num-
ber (fungarium) Collection year Yield of DNA
from extr. (ng/
µl)
260/280
(1.7–1.9) 260/230
(2.0–2.2) MiSeq Nano
output file size
(Mb)
Reads
mapped to
target
Capture pool No. (%) of
single-copy
orthologs, out
of the 75 used
No. (%) of single
copy orthologs
recovered, out of
193 total
Cortinarius C. ominosus TN06-077
(K(M), H) 2006 11 1.88 2.24 14.1 36,403 4 75 (100%) 188 (97%)
Cortinarius C. rubellus TN15-009
(K(M)) 2015 44 1.74 0.94 6.2 12,003 4 30 (40%) 49 (25%)
Cortinarius C. cremeoglo-
bosus TN15-028
(K(M)) 2015 11 1.75 1.56 12.3 27,626 9 70 (93%) 167 (87%)
Cortinarius C. subtortus MT16-001
(K(M)) 2016 11 1.75 1.44 13.2 33,168 9 71 (95%) 177 (92%)
Aureonarius A. callisteus TN07-395
(K(M), H) 2007 55 1.84 1.81 6.6 10,101 9 26 (35%) 52 (27%)
Aureonarius A. limonius TN07-282
(K(M), H) 2007 44 1.92 2.17 15.2 32,781 9 74 (99%) 184 (95%)
Cystinarius C. crassus TN07-305
(K(M), H) 2007 25 1.81 1.5 15.3 40,731 9 74 (99%) 177 (92%)
Phlegmacium P. glaucopus TN12-286
(K(M), H) 2012 19 1.8 1.55 23.4 72,481 9 75 (100%) 193 (100%)
Phlegmacium P. volvatum TN12-267
(K(M), H) 2012 24 1.93 1.83 10.8 23,690 9 67 (89%) 149 (77%)
Thaxterogaster T. malachioides TN07-313
(K(M), H) 2007 17 1.87 0.98 18.9 51,199 9 74 (99%) 185 (96%)
Thaxterogaster T. variegatus TN07-252
(K(M), H) 2007 20 1.85 2.03 31.5 93,312 9 75 (100%) 193 (100%)
Fungal Diversity
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and selected morphological characteristics of the proposed
genera are summarized in Table3.
Cortinariaceae R. Heim ex Pouzar, Česká Mykol. 37(3):
174 (1983) em. Niskanen & Liimat.
Nom. cons. (Art. 14)
Current name of the type species: Cortinarius violaceus
(L.) Gray, Nat. Arr. Brit. Pl. (London) 1: 628 (1821). Sanc-
tioned in Fr., Syst. mycol. 1: 217 (1821). Basionym of the
type species: Agaricus violaceus L., Sp. pl. 2: 1173 (1753).
Lectotype: Bulliard, Herbier de la France: pl. 598 Fig.2A,
1793 (lectotypus hic designatus, IF551873, as Agaricus
araneosus). Epitype: Sweden, Ångermanland, Härnösand,
Geresta, 25 Aug 1993, coll. H. Lindström etal. CFP 1197 (S,
epitypus hic designatus IF551874), GenBank No. OL958654
(ITS).
Synonyms: Gigaspermaceae Jülich, Biblthca Mycol. 85:
367 (1982) [1981]. Nom. illegit., Art. 53.1. Type genus:
Gigasperma E. Horak, N.Z. Jl Bot. 9(3): 491 (1971). Cur-
rent name of the type species: Thaxterogaster crypticus (E.
Horak) Niskanen & Liimat., comb. nov. IF551875. Basio-
nym of the type species: Gigasperma cryptica E. Horak, N.Z.
Jl Bot. 9(3): 491 (1971). Holotype: 27002 (PDD).
Currently included genera: Cortinarius, Aureonarius,
Austrocortinarius, Calonarius, Cystinarius, Hygronar-
ius, Mystinarius, Phlegmacium, Thaxterogaster, and
Volvanarius.
Description: Basidiomata small- to large-sized, aga-
ricoid or sequestrate, development type stipitocarpic
or pileocarpic. Pileus at first conical to hemispherical,
then low conical to low convex to plane, with or without
an umbo; surface smooth, innately fibrillose, tomentose
or ± scaly; ± brown, ± yellow/ochraceous, white, ± grey,
more or less purple or blackish brown to black, more rarely
orange, red, or green/olivaceous; dry, viscid or glutinous,
hygrophanous, with hygrophanous spots or streaks or non-
hygrophanous. Lamellae crowded to distant; adnate, adnexed
or emarginate; when young greyish white, pale grey, pale to
dark brown, or with a purplish tint or purple, more rarely
yellow, green/olivaceous, orange or red. Stipe cylindrical,
clavate, bulbous or rooting; usually silky-fibrillose, white,
pale to dark brown, with purplish tints or purple or ± yel-
low/ochraceous, more rarely green/olivaceous, orange, red
or blackish; dry to glutinous. Universal veil white, yellow/
ochraceous, purple, grey/brown, pink/red, or green/oli-
vaceous, in some species changing colour with age or on
exposure; sparse to abundant, in pileocarpic species found
from the margin of the bulb, in species of the genus Vol-
vanarius often forming a volva at the base of the stipe. In
stipitocarpic species forming incomplete and/or complete
girdles on the stipe, or a sock-like sheet on the lower part
of the stipe, more rarely forming a ring at the upper part
of the stipe; dry or viscid. Odour in many species indis-
tinct, when present in most species then best observed in
Fig. 1 Topology resulting from the RAxML analysis of 75 single-
copy nuclear orthologs. The tree is rooted with Crepidotus sp. and
Hebeloma cylindrosporum as the outgroup. Bootstrap (BS) val-
ues > 50% are indicated above branches. The four species used to
design the baits for the targeted capture sequencing are in purple
Fungal Diversity
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lamellae and then raphanoid, fruity, earthy, cellar-like, cedar
tree-like, perfume-like, yeast-like, farinaceous, grassy, rub-
bery, pelargonium-like, curry-like, anise or unpleasant. The
honey-like odour, typical to part of the species of Cortinar-
ius subgen. Myxacium, Thaxterogaster subgen. Multiformes
and T. subgen. Scauri, is best observed in the context of the
stipe. In part of the species of C. subgenus Iodolentes the
base of the stipe has an odour of iodorm that is best observed
when the basidiomata are slightly dried and in Aureonarius
subgen. Callistei the odour of the surface of the pileus in
some species is like a recently extinguished candle (ozone)
or apple-like. KOH/NaOH reaction useful in identification of
the species of Calonarius and Phlegmacium. Basidiospores
4.5–20 × 3–10μm, in vast majority of the species ± amygda-
loid, ± ellipsoid, ± citriform or ± subglobose, less commonly
obovoidly ellipsoid, fusoid, lacrymoid, or boletoid, finely
to strongly verrucose, somewhat to strongly dextrinoid in
Melzer’s reagent, some species non-dextrinoid. Cystidia pre-
sent in genus Cystinarius, Cortinarius subgen. Cortinarius,
C. sect. Camphorati, C. sect. Subtorti, and some species
of C. subgen. Iodolentes, C. sect. Bicolores and genus Vol-
vanarius. Pileipellis in vast majority of the taxa ± duplex
with a more or less developed hypoderm, simplex in Calon-
arius, Austrocortinarius, Phlegmacium subgen. Cyanicium,
Cortinarius subgen. Cortinarius and C. sect. Subtorti and
in part of the species of Phlegmacium subgen. Phlegma-
cium, Cortinarius sect. Delibuti. In genus Cystinarius some-
what duplex-like, the hypoderm is poorly developed but the
hyphae beneath the epicutis are hypoderm-like (elements
that are short and wide).
Ecology and Distribution: With a world-wide distribu-
tion; species occur both in Northern and Southern Hemi-
sphere from tropical to arctic-alpine habitats. The species
of Cortinariaceae are ectomycorrhizal and form associa-
tions with the trees and shrubs from Fagales, Salicaceae,
Cistaceae, Dipterocarpaceae, Myrtaceae, Fabaceae (e.g.,
Dicymbe), Rhamnaceae, Rosaceae, and Pinaceae, as well
as with some herbaceous angiosperms in the Cyperaceae,
Orchidaceae, and Polygonaceae.
Notes: Typical for the species of the family Cortinari-
aceae are ornamented basidiospores that are cinnamon
brown in deposit. Most species also have a cobweb-like
inner veil covering the young lamellae and the remnants of
it can often still been found at the upper part of the stipe in
older basidiomata. Characteristic is also the silky-fibrillose
stipe, at least easily observed at the top of the stipe. Majority
of the species lack cheilo- and/or pleurocystidia which are
only found in genus Cystinarius and some lineages of genera
Fig. 2 Topology resulting from the RAxML analysis of RPB1, RPB2,
MCM7, GPD, and TEF1. The tree is rooted with Thaxterogaster
as the outgroup. Bootstrap (BS) values > 50% are indicated above
branches. The new subgenera described are in dark blue
Fungal Diversity
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Cortinarius and Volvanarius. The structure of the pileipel-
lis in the majority of genera and subgenera is duplex with a
more or less developed hypoderm.
Variation in size and coloration of basidiomata is large.
One of the smallest species, Cortinarius bibulus, has a pileus
of 0.3–1.5cm in diam. and a stipe 1.5–5 × 0.1–0.3cm, and
one of the largest, Phlegmacium praestans, has a pileus up
to 20cm in diam. and a stipe up to 20 × 3cm. A wide vari-
ety of colored pigments can be found from the basidiomata
although brownish, ochrceous/yellow, greyish, whitish, and
purplish colours are most common.
Cortinarius (Pers.) Gray, Nat. Arr. Brit. Pl. (London) 1:
627 (1821) em. Niskanen & Liimat.
Nom. cons. (Art. 14)
Basionym: Agaricus sect. Cortinaria Pers., Syn. meth.
fung. (Göttingen) 2: 276 (1801).
Current name of the type species: Cortinarius violaceus
(L.) Gray, Nat. Arr. Brit. Pl. (London) 1: 628 (1821). Bas-
ionym of the type species: Agaricus violaceus L., Sp. pl. 2:
1173 (1753).
Synonyms: Cuphocybe R. Heim, Revue Mycol., Paris
16: 8 (1951). Current name of the type species: Cortinarius
elaiochrous E. Horak, M.M. Moser, Peintner & Vilgalys,
Mycotaxon 83: 449 (2002). Basionym of the type species:
Cuphocybe olivacea R. Heim, Revue Mycol., Paris 16: 8
(1951).
Dermocybe (Fr.) Wünsche, Die Pilze: 87, 125 (1877).
Basionym: Agaricus trib. Dermocybe Fr., Syst. mycol. (Lun-
dae) 1: 10, 227 [‘217’] (1821). Current name of the type spe-
cies: Cortinarius cinnamomeus (L.) Gray [as ‘Cortinaria’],
Nat. Arr. Brit. Pl. (London) 1: 630 (1821). Basionym of the
type species: Agaricus cinnamomeus L. 1753.
Hydrocybe (Fr. ex Rabenh.) Wünsche, Die Pilze: 87, 119
(1877). Basionym: Cortinarius a Hydrocybe Fr. ex Rabenh.,
Deutschl. Krypt.-Fl. (Leipzig) 1: 488 (1844). Current name
and basionym of the type species: Cortinarius duracinus
Fr., Epicr. syst. mycol. (Upsaliae): 304 (1838) [1836–1838].
Inoloma (Fr.) Wünsche, Die Pilze: 87, 126 (1877). Bas-
ionym: Agaricus trib. Inoloma Fr., Syst. mycol. (Lundae) 1:
216 (1821). Current name of the type species: Cortinarius
violaceus (L.) Gray, Nat. Arr. Brit. Pl. (London) 1: 628
(1821). Basionym of the type species: Agaricus violaceus
L., Sp. pl. 2: 1173 (1753).
Myxacium (Fr.) P. Kumm., Führ. Pilzk. (Zerbst): 22
(1871). Basionym: Agaricus trib. Myxacium Fr., Syst. mycol.
(Lundae) 1: 247 (1821). Current name of the type species:
Cortinarius collinitus (Sowerby) Gray [as 'Cortinaria col-
linita'], Nat. Arr. Brit. Pl. (London) 1: 628 (1821). Basionym
of the type species: Agaricus collinitus Sowerby, Col. fig.
Engl. Fung. Mushr. (London) 1(no. 2): tab. 9 (1796).
Myxopholis Locq., Fl. Mycol., 3. Cortinariales-A.: 146
(1979) [1977]. Basionym and current name of the type
Table 3 Species diversity, distribution, and summary of the selected morphological characteristics of the 10 genera of Cortinariaceae
*The largest subgenus Telamonia includes 80 sections
Genus Generic
syno-
nyms
Estimated
no. of spe-
cies
Subgen./sec-
tion no Distribution Cystidia Stipitocarpic
(S)/Pile-
ocarpic (P)
Pileipellis
simplex/
duplex
Sequestrate
sp Appearance of the agaricoid basidiomata Rozitoid/
cuphocy-
boid
Myxacioid Phlegmac Telamonioid Cortinarioid
Cortinarius 11 > 2000 11/130* N + S (X) S/(P) Duplex
(+ simpl.) X X (X) X X X
Phlegmacium 3 > 200 4/23 N (+ S) P/S Duplex
(+ simpl.) X X (X)
Thaxtero-
gaster 3 > 200 6/22 (N +) S S/P Duplex X X X
Calonarius – ~ 200 3/14 N P Simplex X X
Aureonarius – ~ 25 2/3 (N +) S S Duplex X
Cystinarius – ~ 10 2/2 N + S X S Duplex X (X)
Volvanarius – ~ 10 1/1 S (X) P Duplex X X
Hygronarius – ~ 10 2/2 (N +) S S Duplex X
Austrocorti-
narius < 5 1/1 S S Duplex X
Mystinarius – < 5 1/1 N + S S Duplex X X
Fungal Diversity
1 3
species: Cortinarius mucifluus Fr., Epicr. syst. mycol.
(Upsaliae): 274 (1838) [1836–1838].
Protoglossum Massee, Grevillea 19(no. 92): 97 (1891).
Current name of the type species: Cortinarius atratus (Rod-
way) Gasparini, Mycosphere 5(4): 542 (2014). Basionym of
the type species: Protoglossum luteum Massee, Grevillea
19(no. 92): 97 (1891).
Quadrispora Bougher & Castellano, Mycologia 85(2):
285 (1993). Current name of the type species: Cortinarius
oblongisporus (G.W. Beaton, Pegler & T.W.K. Young) Gas-
parini, IOSR Journal of Pharmacy 6(4): 3 (2014). Basionym
of the type species: Hymenogaster oblongisporus G.W. Bea-
ton, Pegler & T.W.K. Young, Kew Bull. 40(1): 188 (1985).
Rozites P. Karst., Bidr. Känn. Finl. Nat. Folk 32: XX
(1879). Current name of the type species: Cortinarius
caperatus (Pers.) Fr., Epicr. syst. mycol. (Upsaliae): 256
(1838) [1836–1838]. Basionym of the type species: Rozites
caperatus (Pers.) P. Karst., Bidr. Känn. Finl. Nat. Folk 32:
290 (1879).
Sericeocybe Rob. Henry, Bull. trimest. Soc. mycol. Fr.
109(1): 19 (1993). Current name of the type species: Corti-
narius caninus (Fr.) Fr., Epicr. syst. mycol. (Upsaliae): 285
(1838) [1836–1838]. Basionym of the type species: Agaricus
anomalus var. caninus Fr. 1821.
Telamonia (Fr.) Wünsche, Die Pilze: 87, 122 (1877). Bas-
ionym: Agaricus trib. Telamonia Fr., Syst. mycol. (Lundae)
1: 10, 210 (1821). Current name of the type species: Corti-
narius torvus (Fr.) Fr., Epicr. syst. mycol. (Upsaliae): 293
(1838) [1836–1838]. Basionym of the type species: Agaricus
torvus Fr., Observ. mycol. (Havniae) 2: 80 (1818).
Currently included subgenera: Cortinarius, Camphorati,
Dermocybe, Illumini, Infracti, Iodolentes, Leprocybe, Myxa-
cium, Orellani, Paramyxacium, and Telamonia (Fig.3).
Description: Basidiomata small- to large-sized, agaricoid
or sequestrate, development type stipitocarpic, very rarely
pileocarpic. Pileus 0.3–13cm, at first conical to hemispheri-
cal, then low conical, to low convex to plane, with or with-
out an umbo; surface smooth, innately fibrillose, tomentose
or ± scaly; in the vast majority of the species pale to dark
red-, ochraceous- or grey- brown, in other species ± yellow/
orange, ± white, ± red, greenish/olivaceous, purple, umber
to blackish; dry, viscid or glutinous, hygrophanous, with
hygrophanous spots or streaks or non-hygrophanous. Lamel-
lae in most species medium spaced, in others crowded or
distant; adnate, adnexed or emarginate; when young grey-
ish white, pale grey, pale to dark brown, or with a purplish
tint or purple, more rarely yellow, green/olivaceous, orange
or red. Stipe 1.5–13cm long, 0.1–2.5cm wide at the apex,
up to 5cm at the base; in the vast majority of the species
cylindrical to clavate, less often rooting or bulbous; silky-
fibrillose, white, pale to dark brown, with purplish tints or
purple, more rarely yellow, green/olivaceous, orange or red;
dry to viscid. Universal veil in the majority of the species
white, in others yellow/ochraceous, purple, green/oliva-
ceous, pink or red, in some species at first white and then
turning pink; sparse to abundant, forming incomplete and/
or complete girdles on the stipe, or a sock-like sheet on the
lower part of the stipe, more rarely forming a ring at the
upper part of the stipe; dry or viscid. Context in the vast
majority of the species brownish white, pale to dark red-,
ochraceous- or grey-brown, sometimes with a purplish tint,
in other species ± yellow/orange, ± red, greenish/olivaceous,
purple, umber or blackish. Odour in many species indistinct
or raphanoid, usually best observed in lamellae, in certain
groups pelargonium-like, cedar tree-like, fruity, perfume-
like, iodoform-like, earthy or unpleasant: in some taxa best
observed in the context of the stipe and honey-like, sweet
or raphanoid; in most species of C. sect. Obtusi and Acetosi
the odour is iodoform-like and best observed at the base
of the stipe when somewhat dried. KOH reaction in most
species negative in pileus, context and/or stipital veil, in
some groups red, yellow to orange-yellow, brown or black.
Basidiospores 4.5–20 × 3–10μm, in vast majority of the
species ± amygdaloid, ± ellipsoid or ± subglobose, less com-
monly obovoidly ellipsoid, fusoid, lacrymoid, citriform or
boletoid, finely to strongly verrucose. Cystidia absent in vast
majority of the species, cheilo- and/or pleurocystidia present
in some groups. Pileipellis ± duplex, hypoderm usually more
or less developed, lacking from C. subgen. Cortinarius.
Ecology and Distribution: In the Northern and Southern
Hemisphere with a wide range of hosts.
Notes: The species of this globally distributed, exception-
ally species-rich genus of Cortinariaceae are characterized
by mainly stipitocarpic development and a pileipellis duplex
with a more or less developed hypoderm. The basidiomata
range from very small to large, from dry to glutinous, and
are of varied colours although brown colours are the most
common. Secondary metabolites containing nitrogen are
currently only known from this genus of the family and are
present in the subgenera Cortinarius, Infracti, Orellani, and
section Subtorti (Stensrud etal. 2014).
Morphologically similar species, previously included in
this entity but phylogenetically distinct from the genus Cor-
tinarius, are found in the genera Aureonarius, Cystinarius,
Hygronarius, Thaxterogaster sect. Vibratiles and Phlegma-
cium subgen. Carbonella.
Cortinarius subgen. Cortinarius
Synonym: Inoloma (Fr.) Wünsche, Die Pilze: 87, 126
(1877). Basionym: Agaricus trib. Inoloma Fr., Syst. mycol.
(Lundae) 1: 216 (1821).
Currently included sections: Cortinarius.
Notes: The species of this small bihemispheric subge-
nus have a unique combination of characters and are easy
to identify at the subgeneric level. The basidiomata are
medium to rather large-sized, deep violet to almost blackish
Fungal Diversity
1 3
Fig. 3 Photos of the representatives of genus Cortinarius. A. C. sub-
gen. Cortinarius, C. harcynicus TN 04-525 (H), B. C. subgen. Der-
mocybe, C. neosanguineus TN 09-130 (H), C. subgen. Orellani, C.
rubellus TN 05-024 (H), D. C. subgen. Iodolentes, C. mammillatus
TN 06-249, E. C. subgen. Telamonia, C. badiolaevis TN 04-960 (H),
F. C. sect. Subtorti, C. subtortus TN 05-021 (H), G. C. subgen. Myxa-
cium, C. seidliae TN09-063 (H), and H. C. subgen. Paramyxacium,
C. caperatus TN 06-149 (H). Photos K. Liimatainen
Fungal Diversity
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violet, stipitocarpic, agaricoid (cortinarioid) with a dry
pileus and a dry stipe. The pileus is tomentose to scaly and
non-hygrophanous, and the KOH reaction on any surface of
the basidiomata is red. Pleurocystidia and cheilocystidia are
present and the pileipellis lacks a well-developed hypoderm.
For a recent morpho-genetic revision see Harrower etal.
(2015a, b).
Cortinarius subgen. Camphorati Liimat., Niskanen &
Ammirati, Index Fungorum 256: 2 (2015)
Current name of the type species: Cortinarius campho-
ratus (Fr.) Fr., Epicr. syst. mycol. (Upsaliae): 280 (1838)
[1836–1838]. Basionym of the type species: Agaricus cam-
phoratus Fr., Syst. mycol. (Lundae) 1: 218 (1821). Neotype:
S F-14265, in Brandrud etal., Cortinarius Flora Photograph-
ica I, pl. A12 (1989).
Possible synonym: Sericeocybe Rob. Henry, Bull. trimest.
Soc. mycol. Fr. 109(1): 19 (1993). Current name of the type
species: Cortinarius caninus (Fr.) Fr., Epicr. syst. mycol.
(Upsaliae): 285 (1838) [1836–1838]. Basionym of the type
species: Agaricus anomalus var. caninus Fr. 1821.
Currently included sections: Camphorati.
Notes: The core group of this subgenus, the small bihemi-
spheric C. sect. Camphorati, is easy to delimit based on
morphology: The basidiomata are medium to large-sized,
blue/purple, white to pale yellowish brown, stipitocarpic,
agaricoid (telamonioid) with a dry pileus and dry stipe. The
odour in many species is strong and unpleasant. Cheilocys-
tidia are present and the pileipellis is somewhat duplex, but
the hypoderm is not that well-developed. In the phylogenetic
analysis of Soop etal. (2019) this group was placed within a
larger entity including sections Anomali, Spilomei, Bolares,
Delibuti and Subtorti but without support. Further studies
will be needed to define the limits of the subgenus.
Cortinarius subgen. Dermocybe (Fr.) Trog, Mitt. naturf.
Ges. Bern 15–23: 43 (1844)
Basionym: Agaricus trib. Dermocybe Fr., Syst. mycol.
(Lundae) 1: 10, 227 [‘217’] (1821).
Current name of the type species: Cortinarius cin-
namomeus (L.) Gray [as ‘Cortinaria’], Nat. Arr. Brit. Pl.
(London) 1: 630 (1821). Basionym of the type species: Aga-
ricus cinnamomeus L., Sp. pl. 2: 1173 (1753). Neotype: S
F-44851, in Niskanen, Index Fungorum 221: 1 (2015).
Synonym: Dermocybe (Fr.) Wünsche, Die Pilze: 87,
125 (1877). Basionym: Agaricus trib. Dermocybe Fr., Syst.
mycol. (Lundae) 1: 10, 227 [‘217’] (1821).
Currently included sections: Dermocybe, Aureifolii, Cru-
entoides, Malicoriae, Pauperae and Sanguinei.
Notes: This bihemispheric subgenus includes agaricoid
(dermocyboid), stipitocarpic, small- to medium-sized spe-
cies with yellow, orange, red or olive colours. The stipe is
dry, and the pileus is dry, hygrophanous or not, and felty,
squamulose or glabrous. The pileipellis is somewhat duplex
with a poorly developed hypoderm.
Cortinarius subgen. Illumini Liimat., Niskanen & Kytöv.,
Index Fungorum 256: 2 (2015)
Current name and basionym of the type species: Cor-
tinarius illuminus Fr., Epicr. syst. mycol. (Upsaliae): 305
(1838) [1836–1838] (Lundae) 1: 218 (1821). Neotype: S
F-44877, in Niskanen etal., Index Fungorum 256: 1 (2015).
Currently included sections: Illumini.
Notes: This is a small bihemispheric subgenus that
includes agaricoid (telamonioid), stipitocarpic, medium-
sized species with a vivid red-brown to brown, dry, hygroph-
anous pileus, dry stipe, subglobose basidiospores and a
pileipellis duplex.
Cortinarius subgenus Infracti Niskanen & Liimat., sub-
gen. nov.
IndexFungorum IF551876
Current name and basionym of the type species: Corti-
narius infractus (Pers.) Fr., Epicr. syst. mycol. (Upsaliae):
261 (1838) [1836–1838]. Neotype: S F-41138, in Lii-
matainen etal., Persoonia 33: 120, (2014).
Etymology: Named after the type species of the subgenus.
Currently included sections: Infracti.
Description: Basidiomata medium-sized to large-sized,
agaricoid (phlegmacioid), development type stipitocarpic.
Pileus 3–10cm, at first hemispherical, then low convex to
almost plane, sometimes with a very low and broad umbo,
innately fibrillose; olivaceous grey, olivaceous brown or
umber brown, some species becoming yellow/ochraceous
brown with age; viscid or glutinous; not hygrophanous.
Lamellae crowded, adnate, adnexed to emarginate, dark
olivaceous brown to dark olivaceous grey at least when
young, sometimes with a purplish tint. Stipe 3–9cm long,
0.8–1.5cm wide at the apex, up to 2.5cm at base, cylindrical
to clavate; whitish grey to olivaceous grey, sometimes with a
purplish tint at the apex, dry. Universal veil yellow to yellow
brown, in some species white when young, rather sparse,
fibrillose. Context in pileus and stipe whitish to olivaceous
grey, sometimes purple at the apex of the stipe, marbled
hygrophanous. Odour in lamellae indistinct. Taste bitter.
NaOH reaction yellow to orange–yellow (Soop etal. 2018).
Basidiospores 7–9.5 × 5–7μm, subglobose to broadly ellip-
soid, moderately verrucose. Lamellar trama preparation with
abundant small red granules in Melzer´s. Cystidia absent.
Pileipellis duplex, hypoderm present but poorly developed.
Ecology and Distribution: In the Northern Hemisphere
with Fagaceae and Pinaceae.
Notes: A small subgenus of about 10 to 15 agaricoid
(phlegmacioid) species occurring in the Northern Hemi-
sphere. The species of this subgenus can be distinguished
by the combination of bitter taste, olivaceous tints, viscid
Fungal Diversity
1 3
to glutinous, innately fibrillose pileus, clavate to sometimes
almost cylindrical stipe and subglobose to broadly ellipsoid
spores. The development type is stipitocarpic.
Cortinarius subgenus Iodolentes Niskanen & Liimat.,
subgen. nov.
IndexFungorum IF552140
Current name and basionym of the type species: Cor-
tinarius aurae Niskanen & Liimat., in Hyde etal., Fungal
Diversity 100: 247 (2020). Holotype: K(M) 200315.
Etymology: The name refers to the iodoform-like odour
that many of the species of this subgenus have.
Currently included sections:Acetosi, Fragrantiores and
Obtusi.
Description: Basidiomata small- to medium-sized, aga-
ricoid (telamonioid) or sequestrate, development type stip-
itocarpic. Pileus 1–7cm, at first conical to hemispherical,
then low conical to low convex to plane, with an acute or
broader umbo, pileus margin in many smaller species more
or less pellucid-striate, surface often somewhat rimy; yellow
brown, red brown to dark brown; dry; hygrophanous. Lamel-
lae medium spaced to distant, adnate, adnexed or emar-
ginate, yellow brown to strong brown, often with a white
edge. Stipe 2.5–11cm long, 0.15–1.4cm wide at the apex,
cylindrical or rooting; at first white fibrillose, later very
pale brown to yellow brown. Universal veil white, sparse,
or forming complete and/or incomplete girdles on stipe.
Context in pileus ± brown, usually somewhat paler in stipe.
Odour in lamellae indistinct or in some species raphanoid
or cellar-like, at the base of stipe indistinct, raphanoid or
iodoform-like, the latter best observed when slightly dried.
Basidiospores 6.5–10.5 × 4.5–6.5μm, ovoid, amygdaloid
to ellipsoid, in C. fragrantior ovoid-subglobose, finely to
strongly verrucose. Cheilocystidia present in part of the spe-
cies, clavate to balloon-shaped. Pileipellis duplex, hypoderm
developed.
Ecology and Distribution: In Northern and Southern
Hemisphere with a wide range of hosts plants.
Notes: Cortinarius subgenus Iodolentes includes small- to
medium-sized telamonioid species with dry, ± brown pileus
and dry, initially white stipe. The context of the stipe is often
somewhat paler than in the pileus and does not become
darker towards the base of the stipe. Many species have an
iodoform-like odour at the base of the stipe and clavate to
balloon-shaped cheilocystidia.
The species of this subgenus were traditionally included
in the C. subgen. Telamonia due to their dry pileus and dry
stipe, but the first molecular studies showed that they should
be recognized as a separate taxon (Høiland and Holst-Jensen
2000; Peintner etal 2004; Garnica etal. 2005), which is also
supported by our phylogenetic analysis. Iodolentes belongs
to a well-supported branch (BS 94%) in genus Cortinar-
ius that also includes subgenera Dermocybe, Leprocybe,
Illumini and Orellani. Since the species of Iodolentes mor-
phologically differ from the species of other related subgen-
era we here describe the subgenus as new.
Cortinarius subgen. Leprocybe M.M. Moser, Z. Pilzk.
35(3 + 4): 232 (1969) em. Niskanen & Liimat
Current name and basionym of the type species: Cor-
tinarius cotoneus Fr., Epicr. syst. mycol. (Upsaliae): 289
(1838) [1836–1838]. Neotype: S F-44846, in Ammirati etal.,
Persoonia 46: 221 (2021).
Currently included sections: Leprocybe, Fuscotomen-
tosi, Melanoti, Persplendidi, Squamiveneti, Veneti, and
Veronicae.
Notes: The species of this subgenus occur in both the
Northern and Southern Hemispheres. The basidiomata are
small- to medium-sized (occasionally large-sized), agaricoid
(leprocyboid/dermocyboid) or sequestrate, and with a dry
pileus and dry stipe and with yellow, red, or greenish-olive
colours. At least some parts of the basidiomata are fluo-
rescent. For a recent morpho-genetic revision of Northern
Hemispheric Leprocybe see Ammirati etal. (2021) and
Bidaud etal. (2021).
Cortinarius subgen. Myxacium (Fr.) Trog, Mitt. naturf.
Ges. Bern 15–23: 42 (1844)
Basionym: Agaricus trib. Myxacium Fr., Syst. mycol.
(Lundae) 1: 247 (1821)
Current name of the type species: Cortinarius collinitus
(Sowerby) Gray [as 'Cortinaria collinita'], Nat. Arr. Brit.
Pl. (London) 1: 628 (1821). Basionym of the type species:
Agaricus collinitus Sowerby, Col. fig. Engl. Fung. Mushr.
(London) 1(no. 2): tab. 9 (1796). Lectotype: Sowerby, Col.
Fig. Engl. Fungi 1: pl. 9. 1795, in Gómez & Cadiñanos-
Aguirre, J des JEC 2: 135, (2018).
Synonyms: Myxacium (Fr.) P. Kumm., Führ. Pilzk.
(Zerbst): 22 (1871). Basionym: Agaricus trib. Myxacium
Fr., Syst. mycol. (Lundae) 1: 247 (1821).
Myxopholis Locq., Fl. Mycol., 3. Cortinariales-A.: 146
(1979) [1977]. Basionym and current name of the type
species: Cortinarius mucifluus Fr., Epicr. syst. mycol.
(Upsaliae): 274 (1838) [1836–1838].
Quadrispora Bougher & Castellano, Mycologia 85(2):
285 (1993). Current name of the type species: Cortinarius
oblongisporus (G.W. Beaton, Pegler & T.W.K. Young) Gas-
parini, IOSR Journal of Pharmacy 6(4): 3 (2014). Basionym
of the type species: Hymenogaster oblongisporus G.W. Bea-
ton, Pegler & T.W.K. Young, Kew Bull. 40(1): 188 (1985).
Currently included sections: Myxacium, Cuphomorphi,
Defibulati, Marmorati, and Quadrispora.
Notes: This is a bihemispherical subgenus with about
50 species. The basidiomata are medium-sized to small,
agaricoid (cuphocyboid, myxacioid) or sequestrate with a
viscid to glutionous pileus and glutinous to dry stipe with
Fungal Diversity
1 3
white, brown and/or purplish colours. Cylindrical stipes and
relatively large (up to 20μm long), mainly amygdaloid to
citriform basidiospores are also typical. For a recent mor-
pho-genetic revision of the subgenus see Soop etal. (2021).
The type species of the subgenus, C. collinitus, is
described from Britain. Recently, a lectotype for the species
was designated by Gómez and Cadiñanos-Aguirre (2018).
They also challenged the current interpretation of the name
and concluded the species to be more C. trivialis-like. We
agree with this conclusion and materials from Britain will
need to be sequenced for selection of a suitable epitype.
However, since both the current species called as C. collini-
tus as well as C. trivialis-like fungi belong to this subgenus,
we conclude that the subgeneric name Myxacium can be
confidently used for this clade although the fixing of the
name C. collinitus still requires a selection of an epitype.
Cortinarius subgen. Orellani (M.M. Moser) Gasparini,
Australas. Mycol. 23(2): 69 (2004)
Basionym: Cortinarius sect. Orellani M.M. Moser, Z.
Pilzk. 35(3 + 4): 224 (1969)
Current name and basionym of the type species: Cor-
tinarius orellanus Fr., Epicr. syst. mycol. (Upsaliae): 288
(1838) [1836–1838]. Lectotype: Junghuhn, Observationes
Mycologicae in species fungorum tam novas tam male cog-
nitas, Linnaea V, t. 6, f. 9. 1830 (lectotypus hic designatus,
IF552141). Epitypus: Norway, Agder; Tvedestrand, Eidbo,
in forest with Tilia, Quercus and Corylus, 21 Sep 2014, coll.
I-L. Fonneland & D. Pettersen (O F-251482, epitypus hic
designatus IF552142), UNITE No. UDB036242 (ITS).
Currently included sections: Orellani.
Notes: This small bihemispherical subgenus is character-
ized by the lethal nephrotoxin bipyridine orellanine that has
caused severe poisonings and deaths in humans (Schumacher
& Høiland 1983; Danel etal. 2001) and is not found in any
other lineage in Cortinariaceae. The basidiomata of the
species of C. subgen. Orellani are medium-sized, stipitor-
carpic, agaricoid (cortinarioid) with yellow, orange-brown
and saturated reddish-brown colours, and with a dry pileus
and stipe. A tomentose to finely scaly pileus and cylindrical
to somewhat clavate stipe is also typical. The pileipellis is
duplex with a well-developed hypoderm.
Cortinarius subgen. Paramyxacium M.M. Moser & E.
Horak, Beih. Nova Hedwigia 52: V, 263 (1975) em. Nis-
kanen & Liimat.
Current name of the type species: Cortinarius paradoxus
M.M. Moser & E. Horak, Beih. Nova Hedwigia 52: 264
(1975). Holotype: IB 19650506.
Synonyms: Cuphocybe R. Heim, Revue Mycol., Paris
16: 8 (1951). Current name of the type species: Cortinarius
elaiochrous E. Horak, M.M. Moser, Peintner & Vilgalys,
Mycotaxon 83: 449 (2002). Basionym of the type species:
Cuphocybe olivacea R. Heim, Revue Mycol., Paris 16: 8
(1951).
Rozites P. Karst., Bidr. Känn. Finl. Nat. Folk 32: XX
(1879). Current name of the type species: Cortinarius
caperatus (Pers.) Fr., Epicr. syst. mycol. (Upsaliae): 256
(1838) [1836–1838]. Basionym of the type species: Rozites
caperatus (Pers.) P. Karst., Bidr. Känn. Finl. Nat. Folk 32:
290 (1879).
Currently included sections: Cuphocybe, Paramyxacium,
Rozites, Subcastanelli and clade/Achroi.
Notes: The centre of the diversity of this subgenus is in
the Southern Hemisphere with only a few species occur-
ring in the Northern Hemisphere. This subgenus contains
agaricoid (rozitoid, cuphocyboid) and sequestrate species
and the development type of agaricoid species is stipito-
carpic. Typical for the agaricoid species of this subgenus is
the membraneous veil that in most species forms a distinct
ring or collar on the stipe, or in a few species, thick gir-
dles or scales on the stipe. The pileus is viscid/glutinous
to dry, and many species also have squamules or scales on
the pileus, or the pileus is innately fibrillose, radially wrin-
kled and/or rimy. The basidiomata are usually medium- to
large-sized. The basidiospores are medium to large-sized
(8–16 × 5.5–9.5μm), usually ovoid, amygdaloid or citriform,
more rarely ellipsoid to very short and broadly ellipsoid.
Cortinarius subgen. Telamonia (Fr.) Trog, Mitt. naturf.
Ges. Bern 15–23: 43 (1844) em. Niskanen & Liimat.
Basionym: Agaricus trib. Telamonia Fr., Syst. mycol.
(Lundae) 1: 10, 210 (1821)
Current name of the type species: Cortinarius torvus (Fr.)
Fr., Epicr. syst. mycol. (Upsaliae): 293 (1838) [1836–1838].
Basionym of the type species: Agaricus torvus Fr., Observ.
mycol. (Havniae) 2: 80 (1818). Lectotype: Bulliard, Herb.
Fr. (Paris) 2: Tab. 96, pl. 600, 1782 [1781–82], in Lii-
matainen etal., Fungal Diversity 104: 323 (2020). Epitype:
S F-248482, in Liimatainen etal., Fungal Diversity 104:
323 (2020).
Synonym: Hydrocybe (Fr. ex Rabenh.) Wünsche, Die
Pilze: 87, 119 (1877). Basionym: Cortinarius a Hydrocybe
Fr. ex Rabenh., Deutschl. Krypt.-Fl. (Leipzig) 1: 488 (1844).
Type species Cortinarius duracinus Fr., Epicr. syst. mycol.
(Upsaliae): 304 (1838) [1836–1838].
Telamonia (Fr.) Wünsche, Die Pilze: 87, 122 (1877). Bas-
ionym: Agaricus trib. Telamonia Fr., Syst. mycol. (Lundae)
1: 10, 210 (1821).
Currently included sections: 80 sections, see Liimatainen
etal. (2020a).
Notes: This predominantly Northern Hemispheric line-
age is the most species-rich subgenus in Cortinariaceae
including hundreds of species. The basidiomata are small-
to medium-sized (to large), stipitorcarpic, agaricoid (tela-
monioid) with a dry pileus and stipe. The basidiomata are
Fungal Diversity
1 3
predominantly with brown, grey, white, and/or purplish
colours. The pileipellis is duplex, with a more or less
developed hypoderm. For a recent morpho-genetic revi-
sion of the subgenus see Liimatainen etal. (2020a).
Aureonarius Niskanen & Liimat. gen. nov.
IndexFungorum IF552143
Current name of the type species: Aureonarius kroegeri
(Niskanen, Liimat., E. Harrower, Berbee, Garnica &
Ammirati) Niskanen & Liimat. comb. nov. IF552144. Bas-
ionym of the type species: Cortinarius kroegeri Niskanen,
Liimat., E. Harrower, Berbee, Garnica & Ammirati, Index
Fungorum 294: 1 (2016). Holotype: UBC F15952.
Etymology: Derived from the latin word aureus meaning
golden, since species of this genus have yellow colours in
their basidiomata, and the generic name Cortinarius.
Currently included subgenera: Aureonarius and Cal-
listei (Fig.4).
Description: Basidiomata small- to medium-sized (rather
large-sized), agaricoid, development type stipitocarpic.
Pileus 1–11cm, at first hemispherical or conical, then low
convex or low conical to almost plane, some species with
an umbo; smooth, finely scaly, innately fibrillose or almost
tomentose yellow; orange, orange-red, orange-brown,
brownish red, yellow–brown, red-brown, umber or black-
ish brown; dry or viscid, hygrophanous or not. Lamellae
rather crowded, medium spaced to distant, adnate, adnexed
to emarginate, white, ± yellow, bright orange, yellow–brown
or ± red. Stipe 2–11cm long, 0.2–1.8cm wide at the apex,
up to 2.5cm wide at the base, clavate, cylindrical or taper-
ing downwards, yellowish white, yellow, yellow–brown to
orange-brown, in some species becoming more brownish
when pressed with the thumb or with age, dry to somewhat
viscid. Universal veil yellow, ochraceous, yellow–brown,
orange-red, orange-brown, brown–red or purple-brown,
sparse or more abundant and then forming complete and
incomplete girdles on the stipe. Context in pileus white,
pale yellow, yellow–brown, orange, orange-brown, red-
brown to umber, in stipe yellow, yellow–brown, orange or
red–orange. Odour of pileus surface or context indistinct or
like a recently extinguished candle (ozone) or apple-like,
odour in lamellae indistinct, raphanoid, cellar-like or raw
potato-like. KOH reaction ± red in stipital veil, pileus and/or
context, or negative. UV fluorescence somewhat yellow or
absent. Basidiospores 5–10.5 × 4.5–7μm, subglobose, ovoid,
broadly ellipsoid, ellipsoid or amygdaloid, finely, moderately
to coarsely verrucose. Chrysobasidia present in two species,
A. rubrocastaneus and A. rubrimarginatus. Cystidia absent.
Pileipellis duplex, hypoderm at least somewhat developed.
Ecology and Distribution: In the Northern and Southern
Hemispheres, with a centre of the diversity in the Southern
Hemisphere. In coniferous (Pinaceae) and deciduous forests
(Nothofagaceae, Fagaceae, Betulaceae).
Notes: The species of the bihemispheric genus Aureonar-
ius are characterised by vivid yellow, orange, or red colours,
at least in some parts of the basidiomata. The basidiomata
are small- to rather large-sized, agaricoid (cortinarioid/lep-
rocyboid), and the development type is stipitocarpic. No
sequestrate species are yet known to belong to this genus.
Some species have a weak yellow UV fluorescence, and
some species exhibit a ± red KOH-reaction in stipital veil,
pileus, or context. This taxon is well supported in our phy-
logenomic analyses, and we here describe it as a new genus.
Aureonarius subgenus Aureonarius
IndexFungorum IF552145
Etymology: Derived from the latin word aureus meaning
golden, since species of this genus have yellow colours in
their basidiomata, and the generic name Cortinarius.
Currently included sections: Aureonarius (= Cortinarius
sect. Limonii Nezdojm).
Description: Basidiomata small- to medium-sized, aga-
ricoid (cortinarioid/leprocyboid), development type stip-
itocarpic. Pileus 1–8cm, at first hemispherical or coni-
cal, then low convex or low conical to almost plane, often
with an umbo, yellow, orange, orange-red, orange-brown,
brownish red, yellow–brown, red-brown, umber, or black-
ish brown, dry or viscid, hygrophanous or not. Lamellae
rather crowded, medium spaced to somewhat distant, adnate,
adnexed to emarginate, ± yellow, bright orange, pale yel-
low–brown or ± red. Stipe 2–11cm long, 0.2–1.8cm wide
at the apex, cylindrical to fusoid, often tapering downwards,
yellowish white, yellow, yellow–brown to orange-brown, dry
to somewhat viscid. Universal veil yellow, ochraceous, yel-
low–brown, orange-brown, or brown–red, sparse or more
abundant and then forming complete and incomplete gir-
dles on the stipe. Context in pileus yellow–brown, orange,
orange-brown, red-brown to umber, in stipe yellow, yel-
low–brown, orange or red–orange. Odour in lamellae or
pileus surface indistinct. KOH reaction red to dark red in
stipital veil, pileus and/or context, or negative. UV fluo-
rescence weak or absent (Soop etal. 2018). Basidiospores
5–10.5 × 4.5–7μm, subglobose, broadly ellipsoid, ellipsoid
or amygdaloid, finely, moderately to coarsely verrucose.
Chrysobasidia present in two species, A. rubrocastaneus
and A. rubrimarginatus. Cystidia absent. Pileipellis duplex,
hypoderm developed, some species with a thin gelatinous
layer at the top of the epicutis.
Ecology and Distribution: The centre of the diversity of
this lineage is in New Zealand where the species occur in
Myrtaceae and Nothofagaceae forests. The three species
known from the Northern Hemisphere, associated with
Fagaceae and Pinaceae, are clustered in one monophyletic
lineage within one of the New Zealand lineages.
Notes: The species of this small, bihemispheric subge-
nus have small- to medium-sized, stipitocarpic, agaricoid
Fungal Diversity
1 3
(cortinarioid/leprocyboid) basidiomata with yellow, orange-
red and reddish-brown colours. The pileus is dry to viscid,
and the stipe is cylindrical to fusoid and dry. The lamellae
are ± yellow, bright orange, pale yellow–brown or ± red and
the basidiospores are subglobose, broadly ellipsoid, ellipsoid
or amygdaloid. A distinct odour in the lamellae or at the
pileus surface is lacking. The species of the sister subgenus
Callistei differ from the species of subgenus Aureonarius by
having white, pale yellow or greyish ochraceous lamellae at
least when young and somewhat yellow UV fluorescence.
In addition, some species of the subgenus Callistei have a
clavate stipe and a distinct smell at the pileus surface, con-
text or lamellae, and none of the species have amygdaloid
or ellipsoid spores.
Aureonarius subgenus Callistei (Liimat., Niskanen &
Ammirati) Niskanen & Liimat., comb. nov.
IndexFungorum IF552146
Basionym: Cortinarius subgen. Callistei Liimat., Nis-
kanen & Ammirati, in Niskanen, Liimatainen, Kytövuori &
Ammirati, Index Fungorum 256: 2 (2015).
Current name of the type species: Aureonarius callisteus
(Fr.) Niskanen & Liimat. comb. nov. IF552147. Basionym
of the type species: Agaricus callisteus Fr., Observ. mycol.
(Havniae) 2:51. 1818. Neotype: S CFP1219, in Brandrud
etal., Cortinarius flora photographica 5: pl. E30, (2012).
Currently included sections: Callistei and Collybiani.
Description: Basidiomata small- to medium-sized, aga-
ricoid (cortinarioid/leprocyboid), development type stipito-
carpic. Pileus 2–11cm, at first hemispherical to somewhat
conical, then low convex to almost plane, some species with
an umbo, smooth, finely scaly, innately fibrillose or almost
tomentose, yellow, yellow-orange, yellow–brown, orange-
brown, brownish red to mahogany-red, dry, hygrophanous or
not. Lamellae medium spaced to distant, adnate to emargin-
ate, at first almost white, pale yellow or yellowish brown,
later brownish yellow to brown. Stipe 3.5–11cm long,
0.5–1.5cm wide at the apex, up to 2.5cm wide at the base,
clavate, cylindrical to somewhat tapering; yellowish white,
pale yellow, yellow–brown, becoming more brownish when
pressed with the thumb and with age. Universal veil yel-
low, yellow–brown, orange-red or purple-brown, forming
complete and/or incomplete zones on the stipe, or sparse.
Context in pileus white to pale yellow, in stipe pale yellow,
yellow–brown to orange-brown, in many species becoming
darker with age. Odour of pileus surface or context like a
recently extinguished candle (ozone), apple-like or indis-
tinct, odour in lamellae indistinct, raphanoid, cellar-like or
raw potato-like. KOH reaction in pileus and/or stipital veil
brownish red to red. UV fluorescence somewhat yellow.
Basidiospores 5.5–9 × 5–7μm, subglobose, ovoid to broadly
ellipsoid, finely to moderately verrucose. Cystidia absent.
Pileipellis duplex, hypoderm at least somewhat developed.
Ecology and Distribution: In the Northern and Southern
Hemispheres in coniferous and deciduous forests.
Notes: The species of this small, bihemispheric subge-
nus have small- to medium-sized, stipitocarpic, agaricoid
(cortinarioid/leprocyboid) basidiomata with yellow, orange
and brownish-red colours. The pileus is dry, and the stipe is
clavate or cylindrical and dry. The lamellae are at first almost
white, pale yellow or yellowish brown and the basidiospores
are subglobose, ovoid to broadly ellipsoid. Many species
have a distinct odour either at the pileus surface or in the
lamellae. The species of the sister subgenus Aureonarius
have ± yellow, bright orange, pale yellow–brown or ± red
lamellae and an indistinct odour in the lamellae or at the
pileus surface. In addition, there is no UV fluorescence in
the basidiomata of the species of subgenus Aureonarius.
Aureonarius section Callistei Niskanen & Liimat., sect.
nov.
IndexFungorum IF552148
Current name of the type species: Aureonarius callisteus
(Fr.) Niskanen & Liimat.
Etymology: Named after A. callisteus, a species belonging
to this section.
Currently included species: C. neocallisteus, C. callis-
teus, C. infucatus, C. tofaceus.
Description: Basidiomata medium-sized (to large-
sized), agaricoid (cortinarioid/leprocyboid), development
type stipitocarpic. Pileus 3–11cm, at first hemispherical,
then low convex to almost plane, smooth to finely scaly to
almost tomentose, yellow, yellow-orange, yellow–brown to
orange-brown, dry, somewhat to not hygrophanous. Lamel-
lae medium spaced to distant, adnate to emarginate, at first
almost white, pale yellow or yellowish brown, later brown-
ish yellow to brown. Stipe 3.5–11cm long, 0.6–1.5cm
wide at the apex, up to 2.5cm wide at the base, clavate
or cylindrical; pale yellow, yellow brown, becoming more
brownish when pressed with the thumb and with age. Uni-
versal veil yellow to yellow–brown, forming complete and/
or incomplete zones on the stipe, sometimes sparse. Con-
text in pileus white to pale yellow, in stipe yellow–brown to
orange-brown, becoming darker with age. Odour of pileus
surface like a recently extinguished candle (ozone), apple-
like or indistinct, odour in lamellae raphanoid, cellar-like
or raw potato-like. KOH reaction in pileus and/or stipital
veil brownish red to red. UV fluorescence somewhat yel-
low. Basidiospores 6.5–9 × 5.5–7μm, subglobose to ovoid,
moderately verrucose. Cystidia absent. Pileipellis duplex,
hypoderm somewhat developed.
Ecology and Distribution: In the Northern Hemisphere
in coniferous and deciduous forests.
Notes: A Northern Hemispheric lineage in A. subgenus
Callistei. The representatives of the sister lineage, A. sect.
Collybiani from the Southern Hemispheric Nothofagaceae
Fungal Diversity
1 3
forests, often have a somewhat more reddish-coloured
pileus and darker universal veil. The group received full
support in the phylogenetic analysis of Soop etal. (2018,
2019).
Aureonarius section Collybiani Niskanen & Liimat., sect.
nov.
IndexFungorum IF552149
Current name of the type species: Aureonarius collybi-
anus (Soop) Niskanen & Liimat., comb. nov. IF552150.
Basionym of the type species: Cortinarius collybianus
Soop, Bull. Soc. mycol. Fr. 117(2): 121 (2001). Holotype:
PDD 70509.
Etymology: Named after the type species of the section.
Currently included species: C. collybianus, C. eucol-
lybianus, C. rubrodactylus.
Description: Basidiomata small- to medium-sized,
agaricoid (cortinarioid/leprocyboid), development type
stipitocarpic. Pileus 2–8.5cm, at first hemispherical to
somewhat conical, then convex to plano-convex, often
with an umbo, smooth to finely innately fibrillose, orange-
red, apricot-brown, brownish red to mahogany red, dry,
hygrophanous or not. Lamellae medium spaced to distant,
adnate to emarginate, at first white, pale yellow or grey-
ish ochraceous later more brownish. Stipe 4–9cm long,
0.5–1.2cm wide at the apex, somewhat clavate, cylindri-
cal to somewhat tapering, yellowish white, pale yellow,
yellow or brown-yellow, becoming more brownish at the
base with age. Universal veil orange-red, orange-brown, or
purple-brown, sparse. Context white or pale yellow. Odour
in context like wax-candles. KOH reaction dark red on
pileipellis and stipital veil or trivial. Not UV fluorescent.
Basidiospores 5.5–8.5 × 5–6μm, subglobose to broadly
ellipsoid, finely to moderately verrucose. Cystidia absent.
Pileipellis duplex, hypoderm developed.
Ecology and Distribution: In the Southern Hemisphere
in Nothofagaceae forests.
Notes: A Southern Hemispheric lineage in A. subge-
nus Callistei. The representatives of the sister lineage, A.
sect. Callistei from the Northern Hemisphere, often have
a somewhat paler, less reddish pileus and a yellow to yel-
low–brown universal veil. The group received full support
in the phylogenetic analysis of Soop etal. (2018, 2019).
Austrocortinarius Niskanen & Liimat., gen. nov.
IndexFungorum IF552197
Current name of the type species: Austrocortinarius
victoriaensis (Liimat.) Niskanen, comb. nov. IF552198
Basionym of the type species: Cortinarius victoriaensis
Liimat., Index Fungorum 506: 2 (2021); Holotype: K(M)
162337
Etymology: A genus of family Cortinariaceae that is cur-
rently only known from the Southern Hemisphere.
Currently included subgenera: The genus includes only
a few species and no infrageneric classification is proposed
at present.
Description: Basidiomata (medium- to) large-sized,
agaricoid (phlegmacioid), development type stipitocarpic.
Pileus 6–15cm, at first hemispherical, then convex to plano-
convex, margin often with hanging remnants of veil, white
to brownish white. Lamellae crowded, adnate to emarginate,
at first almost white to very pale brown, later pale brown
to brown. Stipe 7–15cm long, 1.2–3cm wide at the apex,
rooting, white. Universal veil white, peronate, often forming
a distinct ring at the upper part of the stipe. Context white.
Odour not recorded. KOH reaction not recorded. Basidi-
ospores 10–12 × 5–6.5μm, amygdaloid to citriform, finely
verrucose. Cystidia absent. Pileipellis simplex, hypoderm
not developed.
Ecology and Distribution: In the Southern Hemisphere in
Australia and New Zealand. In Myrtaceae forests.
Notes: Austrocortinarius is a small, Southern Hemi-
spheric genus currently only known from Australia and New
Zealand. The representatives of the genus are easy to recog-
nize by the combination of pileipellis simplex, large, ± white
basidiomata and a peronate universal veil often forming a
distinct ring at the upper part of the rooting stipe. In addi-
tion, the basidiomata are agaricoid (phlegmacioid), stipito-
carpic and the spores are rather large and amygdaloid. The
sequence data deposited in the public repositories, as well
as morphology, indicates that C. australiensis would also
belong to this genus but the type specimen of the species
has not been studied to confirm the placement. The species
of genus Austrocortinarius are most reminiscent of those in
P. subgenus Phlegmacium, sect. Arguti and clades Obsoleti
and Caligati but those lineages of Phlegmacium are only
known from the Northern Hemisphere. Rooting, phlegma-
cioid species are also found from genus Thaxterogaster, but
none of those species has the same, unique combination of
characters than the representatives of the genus Austrocorti-
narius. Based on the morphological and molecular data we
here consider this distinct lineage as its own genus.
Calonarius Niskanen & Liimat., gen. nov.
IndexFungorum IF552199
Current name of the type species: Calonarius typicus
(Liimat.) Niskanen, comb. nov. IF552200. Basionym of the
type species: Cortinarius typicus Liimat., in Niskanen &
Liimatainen, Index Fungorum 487: 2 (2021). Holotype: H
7068019.
Etymology: Derived from the section name Calochroi and
the generic name Cortinarius.
Currently included subgenera: Calonarius, Calochroi and
Fulvi (Fig.4).
Description: Basidiomata medium- to large-sized, usu-
ally agaricoid (phlegmacioid) but a few species sequestrate,
Fungal Diversity
1 3
development type pileocarpic. Pileus 3–15cm, at first hemi-
spherical, then convex to plano-convex, surface in part of
the species with small appressed scales or patches of veil
and/or innately fibrillose, often colourful with white, yellow,
orange, green, olivaceous, brown, blackish and/or purple
colours, glutinous. Lamellae crowded, adnate to adnexed to
emarginate, white, pale grey, yellow, greenish, olivaceous,
pale brown or purple. Stipe 3–12cm long, 0.7–3cm wide
at the apex, with more or less, usually distinctly, margin-
ated bulb at the base (up to 4.5cm wide), bulb in some
species flattened; white, pale grey, yellow, olivaceous green
or purple. Universal veil white, ochraceous yellow, oliva-
ceous/greenish yellow, orange, brown, brown-olive or pur-
ple, found at the bulb margin. Context white, greyish white,
yellow or greenish yellow, in some species with a purplish,
greenish or olivaceous tint. Odour in lamellae indistinct,
earth-like, malty or yeast-like, curry-like, sweet, in one spe-
cies of anise. KOH reaction pink, red, yellowish or orange-
brown, olivaceous green, black or in some species negative.
Basidiospores 8.5–16 × 5.5–9.5μm, amygdaloid to citriform,
distinctly and coarsely verrucose. Cystidia absent. Pileipellis
simplex with a well-developed gelatinous layer, hypoderm
not developed.
Ecology and Distribution: In the Northern Hemisphere.
Forming ectomycorrhizal associations mainly with the trees
of Fagaceae (Castanea, Castanopsis, Chrysolepis, Fagus,
Notholithocarpus, Quercus) and Pinaceae (Abies, Larix,
Picea, Pinus, Pseudotsuga, Tsuga), some species also with
Betulaceae (Alnus, Corylus, Carpinus), Cistaceae (Cistus,
Helianthemum) and Malvaceae (Tilia) (Garnica etal. 2011).
Most species are rare and have narrow ecological prefer-
ences (Frøslev etal. 2007), and the majority are calcicolous
or calciphilous.
Notes: This species-rich genus is currently only known
from the Northern Hemisphere. The species are predomi-
nantly calcicolous or calciphilous, many are rare and have
narrow ecological preferences and are thus included in
national red lists in several countries and/or used as indi-
cator species. One species, C. meinhardii, is also included
in the global red list of fungi. Typical for the members of
this genus are medium- to large-sized, pileocarpic, aga-
ricoid (phlegmacioid) or sometimes sequestrate, often
brightly coloured basidiomata with a more or less, usually
distinctly marginated bulb at the base of the stipe. Amyg-
daloid to citriform coarsely verrucose basidiospores and
simplex pileipellis are also typical. Some species have a
positive KOH-reaction (± red, yellowish or orange-brown,
olivaceous green, black). The species are most reminiscent
of those in the genera Phlegmacium and Thaxterogaster,
but the combination of simplex pileipellis, marginated bulb
and amygdaloid to citriform, coarsely verrucose basidi-
ospores distinguish the members of Calonarius from the
other phlegmacioid species. This group has been recognized
as a separate, well-supported lineage since early molecular
studies (Peintner etal. 2004; Garnica etal. 2005) and is also
supported by morphological characteristics, and here were
propose a name for it in generic level. For the most recent
morpho-genetic study of the group see Frøslev etal. (2007)
and Garnica etal. (2009).
Calonarius subgenus Calonarius
IndexFungorum IF552151
Etymology: Derived from the section name Calochroi and
the generic name Cortinarius.
Currently included sections: Calonarius, Humolentes,
and Rufoolivacei.
Description: Basidiomata medium- to large-sized, usually
agaricoid (phlegmacioid) but at least one species seques-
trate, development type pileocarpic. Pileus 4–14cm, at first
hemispherical, then convex to plano-convex, surface in some
species with patches or scales of veil or spots, with grey,
greenish, olivaceous, yellow, orange-brown, copper brown,
red brown, umber brown and purplish colours, rarely cream-
coloured, glutinous. Lamellae crowded, adnate to adnexed
to emarginate, greyish white, pale ochraceous grey, pale
yellow, greenish grey, greyish green, yellowish green or
olivaceous, rarely with a purplish tint. Stipe 4–12cm long,
0.8–2.5cm wide at the apex, with a marginated or more
rarely rounded bulb at the base (up to 4.5cm wide), bulb
in a few species flattened; white, in some species purplish,
greyish green or yellowish green or with a purple, yellow or
olivaceous tint. Universal veil white, yellow, greyish or yel-
lowish green, brown, brown-olive, purple or purplish red, in
some species somewhat glutinous, found at the bulb margin.
Context white to grey, rarely greenish yellow, in some spe-
cies with a purplish, greenish or olivaceous tint at the stipe.
Odour in lamellae indistinct, earth-like, malty or yeast-like,
curry-like, sweet, in two species of anise. KOH reaction
negative or in some species yellow–brown, orange-brown,
olivaceous green, brown–red or purplish, rarely blood-red.
Basidiospores 10–16 × 6–9.5μm, amygdaloid to citriform,
distinctly and coarsely verrucose. Cystidia absent. Pileipellis
simplex with a well-developed gelatinous layer, hypoderm
not developed.
Ecology and Distribution: In the Northern Hemisphere.
Forming ectomycorrhizal associations mainly with the spe-
cies of Fagaceae, Pinaceae, Betulaceae, and Malvaceae.
Most species are rare and have narrow ecological prefer-
ences, and the majority are calcicolous or calciphilous.
Notes: The species of this subgenus mainly have lamel-
lae without purple tones and the context is white to grey,
rarely greenish-yellow, and in some species with a purplish,
greenish or olivaceous tint at the stipe. The KOH-reaction,
if present, is not red for most species. The basidiomata are
medium- to large-sized, pileocarpic, agaricoid (phlegma-
cioid) and the pileipellis is simplex.
Fungal Diversity
1 3
Calonarius section Humolentes Niskanen & Liimat., sect.
nov.
IndexFungorum IF552330
Current name of the type species: Calonarius humolens
(Brandrud) Niskanen & Liimat., comb. nov. IF552331.
Basionym of the type species: Cortinarius humolens
Brandrud, in Brandrud, Lindström, Marklund, Melot &
Muskos, Cortinarius, Flora Photographica (Matfors) 4: 20.
1998. Holotype: O CFP1281.
Etymology: Named after the type species of the section.
Currently included species: C. anaunianus, C. carovio-
laceus, C. elotus, C. elotoides, C. glaucoelotus, C. hilde-
gardiae, C. humolens, C. lavandulochlorus, C. mariekris-
tinae, C. osloensis, C. praetermissus, C. pseudoglaucopus,
C. rapaceoides, C. saporatus, and C. xanthodryophilus.
Description: Basidiomata medium- to large-sized,
agaricoid (phlegmacioid), development type pileocarpic.
Pileus 4–14cm, at first hemispherical, then convex to
plano-convex, in some species with small drop-like spots
or appressed scales, or more rarely innately fibrillose;
with ochraceous/yellow, grey, greenish/olivaceous, and/
or ochraceous brown colours, rarely orange/reddish brown,
glutinous. Lamellae crowded (to medium spaced), adnate
to adnexed to emarginate, at first pale ochraceous grey,
more or less yellow, olivaceous or with a yellow, oliva-
ceous or greenish tint, later more ochraceous brown. Stipe
3–8cm long, 0.8–2.5cm wide at the apex, with a margin-
ated or rounded bulb at the base (up to 4.5cm wide), bulb
in some species flattened; white, pale yellow, or with a
yellow, olivaceous or purple tint. Universal veil white, yel-
low, greyish or yellowish green, purple or brown, in some
species somewhat glutinous, found at the bulb margin, in
some species volva-like when young. Context white, grey,
in some species with an olivaceous, yellow or purplish
tint. Odour in most species in flesh/lamellae earth-like or
raphanoid, in some species malty. KOH reaction nega-
tive. Basidiospores 9–14 × 5–8μm, amygdaloid to citri-
form, distinctly and coarsely verrucose. Cystidia absent.
Pileipellis simplex with a well-developed gelatinous layer,
hypoderm not developed.
Ecology and Distribution: In the Northern Hemisphere.
In deciduous and coniferous forests on calcareous or base-
rich ground. Forming ectomycorrhizal associations mainly
with the species of Fagaceae, Pinaceae, Betulaceae and
Tilia.
Notes: The species of this section are found from the
Norther Hemisphere and grow on calcareous or base-rich
ground ground with deciduous or coniferous trees. Basidi-
omata have ochraceous/yellow, grey, greenish/olivaceous
and ochraceous brown colours and most species have an
earth-like or raphanoid odour in flesh/lamellae. In addition,
the species lack KOH reaction and they do not have purplish
colours in lamellae. The basidiomata are medium- to large-
sized, pileocarpic, agaricoid (phlegmacioid) and the pileipel-
lis is simplex.
The clade name/Humolentes was first introduced for this
group by Brandrud etal. (2019) and includes clades/Pseu-
doglaucopodes and Caroviolacei recognized by Garnica
etal. (2009). For the most recent phylogenetic study of the
group see Brandrud etal. (2019) and Fellin etal. (2021).
Calonarius subgenus Calochroi Niskanen & Liimat., sub-
gen. nov.
IndexFungorum IF552332
Current name of the type species: Calonarius flavipallens
(Kytöv., Liimat. & Niskanen) Niskanen & Liimat., comb.
nov. IF552333. Basionym of the type species: Cortinarius
flavipallens Kytöv., Liimat. & Niskanen, in Liimatainen,
Niskanen, Dima, Kytövuori, Ammirati & Frøslev, Persoonia
33: 125 (2014). Holotype: H 6032745.
Etymology: Named after C. calochrous, a species belong-
ing to this subgenus.
Currently included sections: Calochroi, Nymphicolores,
Platypodes, and Sodagniti.
Description: Basidiomata medium- to large-sized, usually
agaricoid (phlegmacioid) but at least one species seques-
trate development type pileocarpic. Pileus 3–10cm, at first
hemispherical, then convex to plano-convex, surface in most
species with small appressed scales or patches of veil, ochra-
ceous white, yellow, ochraceous to brown, in some species
with bluish, purplish, greenish or olivaceous tints, in some
species completely purple, glutinous. Lamellae crowded,
adnate to adnexed to emarginate, greyish white with a pur-
plish tint to pale greyish purple to distinctly purple, in a few
species yellow to brown. Stipe 3–12cm long, 0.7–2.5cm
wide at the apex, with a distinctly marginated bulb at the
base (up to 4cm wide), bulb in some species flattened, grey-
ish white, in many species with a purplish tint at the top of
the stipe, in some species completely purple at least when
young. Universal veil whitish to ochraceous yellow, in some
species purple or olivaceous, found at the bulb margin. Con-
text whitish to greyish white, in part of the species pale pur-
ple to purple at least at the top of the stipe. Odour in many
species in lamellae somewhat earth-like in older basidi-
omata. KOH reaction in many species pink, reddish brown
or blood red in some parts of the basidiomata, indistinct
in part of the species. Basidiospores 8.5–13 × 5.5–8.5μm,
amygdaloid to citriform, distinctly and coarsely verrucose.
Cystidia absent. Pileipellis simplex with a well-developed
gelatinous layer, hypoderm not developed.
Ecology and Distribution: In the Northern Hemisphere.
Forming ectomycorrhizal associations mainly with the spe-
cies of Fagaceae, Pinaceae, Betulaceae, Cistaceae and
Fungal Diversity
1 3
Malvaceae. Most species are rare and have narrow eco-
logical preferences, and the majority are calcicolous or
calciphilous.
Notes: This is the most species-rich lineage within the
genus Calonarius. Most species are characterized by a
combination of lamellae with a purplish tint or completely
purplish and lack of anthraquinonoid pigments (Frøslev
etal. 2007). The basidiomata are medium- to large-sized,
pileocarpic, agaricoid (phlegmacioid) and the pileipellis is
simplex. The clade Calochroi was also recovered as a well-
supported lineage in previous studies by Frøslev etal. (2007)
and Garnica etal. (2009).
Calonarius section Nymphicolores Niskanen & Liimat.,
sect. nov.
IndexFungorum IF552334
Current name of the type species: Calonarius molochi-
nus (Bidaud & Ramm) Niskanen & Liimat., comb. nov.
IF552335. Basionym of the type species: Cortinarius
molochinus Bidaud & Ramm, in Bidaud, Moënne-Loccoz,
Reumaux, Carteret & Eyssartier, Atlas des Cortinaires
(Meyzieu) 11: 613. 2001. Holotype: PC 3676.
Etymology: Named after C. nymphicolor, a species
belonging to the section.
Currently included species: C. nymphicolor and C.
molochinus.
Description: Basidiomata medium- to large-sized, agari-
coid (phlegmacioid), development type pileocarpic. Pileus
3.5–10cm, at first hemispherical, then convex to plano-
convex, with small patches of whitish veil at centre, entirely
pinkish purple or at least margin purple when young, centre
cream, pale ochraceous yellow to ± brownish; glutinous.
Lamellae crowded, adnate to adnexed to emarginate, grey-
ish to purple. Stipe 3–7.5cm long, 0.7–2cm wide at the
apex, with a flattened bulb; white, with a purplish tint at
the apex. Universal veil white or yellow, becoming yellow
to ochraceous brown with age. Context white. Odour indis-
tinct or ± earthy. KOH reaction pink on pileus and bulbi-
pellis, somehat rose or negative in context. Basidiospores
8.5–11 × 5–6.5μm, amygdaloid to ellipsoid, coarsely ver-
rucose. Cystidia absent. Pileipellis simplex with a well-
developed gelatinous layer, hypoderm not developed.
Ecology and Distribution: In the Northern Hemisphere.
In deciduous forests on calcareous ground. Forming ectomy-
corrhizal associations mainly with the species of Fagaceae.
Notes: The species of the section are found from the
Northerh Hemispheric deciduous forests on calcareous
ground. Typical are purplish colours in pileus and stipe and
often also in lamellae as well as initially white or yellow
universal veil. The KOH reaction is pink on pileus and bul-
bipellis and the spores are amygdaloid to ellipsoid, coarsely
verrucose. The basidiomata are medium- to large-sized,
pileocarpic, agaricoid (phlegmacioid) and the pileipellis is
simplex. The clade received full support in our phylogenetic
analysis.
Calonarius subgenus Fulvi Niskanen & Liimat., subgen.
nov.
IndexFungorum IF552337
Current name of the type species: Calonarius elegantio-
occidentalis (Garnica & Ammirati) Niskanen & Liimat.,
comb. nov. IF552338. Basionym of the type species: Corti-
narius elegantio-occidentalis Garnica & Ammirati, in Gar-
nica, Spahn, Oertel, Ammirati & Oberwinkler, BMC Evol.
Biol. 11(213): 13 + Additional file3: 23 (2011). Holotype:
WTU, Ammirati 13226.
Etymology: This subgenus includes part of the species
previously included in section Fulvi.
Currently included sections: Fulvi, Atrovirentes, Aureop-
ulverulenti, Dibaphi, and Splendentes.
Description: Basidiomata medium- to large-sized, agari-
coid (phlegmacioid), development type pileocarpic. Pileus
3–15cm, at first hemispherical, then convex to plano-con-
vex, surface in many species with small appressed scales
and/or innately fibrillose, ± yellow, olivaceous green, grey-
ish green, yellow brown, orange-brown, rose brown to red
brown, in some species with purplish tints, in one species
completely purplish, center saffron orange, brown, chest-
nut brown, purplish brown to blackish; glutinous. Lamellae
crowded, adnate to adnexed to emarginate, greyish white,
grey, yellow, greenish yellow, olivaceous yellow, ochraceous
brown to olivaceous green, in some species with a purplish
tint. Stipe 4–12cm long, 1–3cm wide at the apex, with
more or less, usually distinctly marginated bulb at the base
(up to 4.5cm wide), bulb in some species flattened; white,
pale yellow, yellow, greenish yellow to olivaceous green,
in some species with a purplish tint or completely purplish.
Universal veil ± yellow, orange or purple, found at the bulb
margin. Context yellow, greenish yellow, pale yellow or
white, in some species with a purplish tint. Odour in lamel-
lae indistinct or malt-like. KOH reaction pink, vinaceous,
blood-red, olivaceous green, olivaceous brown, red brown
or black. Basidiospores 9–15 × 5.5–9μm, amygdaloid to cit-
riform, distinctly and coarsely verrucose. Cystidia absent.
Pileipellis simplex with a well-developed gelatinous layer,
hypoderm not developed.
Ecology and Distribution: In the Northern Hemisphere.
Forming ectomycorrhizal associations mainly with the spe-
cies of Fagaceae, Pinaceae, Betulaceae, and Malvaceae.
Most species are rare and have narrow ecological prefer-
ences, and the majority are calcicolous or calciphilous.
Notes: Most species of this subgenus are characterized by
yellow colours in the lamellae and/or stipe. If the lamellae
are purple, then the pileus is not yellow. Part of the spe-
cies have anthraquinonoid pigments (Frøslev etal. 2007).
The basidiomata are medium- to large-sized, pileocarpic,
Fungal Diversity
1 3
agaricoid (phlegmacioid) and the pileipellis is simplex. The
group also received good support (BS 96%) in the analysis
by Garnica etal. (2009).
Cystinarius Niskanen & Liimat., gen. nov.
IndexFungorum IF552491
Current name of the type species: Cystinarius rubigino-
sus (Ammirati, Bojantchev, Niskanen & Liimat.) Liimat. &
Niskanen, comb. nov. IF559243. Basionym of the type spe-
cies: Cortinarius rubiginosus Ammirati, Bojantchev, Nis-
kanen & Liimat., Index Fungorum 506: 1 (2021). Holotype:
H 7072000.
Etymology: Derived from the word cystidia, a property of
this genus, and the generic name Cortinarius.
Currently included subgenera: Cystinarius and Crassi
(Fig.4).
Description: Basidiomata medium- to large-sized, aga-
ricoid (phlegmacioid/cortinarioid), development type stip-
itocarpic. Pileus 1.5–11cm, at first hemispherical, then
convex to plano-convex, surface fibrillose to tomentose,
orange yellow, yellow brown, ochraceous brown, greyish
brown, red brown to blackish brown, in some species with
spots, dry or somewhat viscid. Lamellae crowded to medium
spaced, adnate to adnexed to emarginate, white to greyish
yellow. Stipe 3–11cm long, 0.7–2.5cm wide at the apex,
cylindrical, clavate to fusiform, white, pale grey to greyish
yellow, in some species staining yellow or pinkish, dry to
somewhat viscid. Universal veil white, pale grey, ochraceous
yellow to brown, in some species darkening, often sparse,
forming thin bands on the stipe. Context in stipe white to
yellowish brown, in some species darker in the pileus, in
subgenus Rubicunduli staining more or less yellow when
exposed. Odour in lamellae indistinct. KOH reaction nega-
tive. Basidiospores 6–9 × 3.5–5μm, ellipsoid to amygdaloid
to subfusoid, almost smooth to very finely and indistinctly
verrucose. Lamellae with cylindrical, clavate or capitate
cheilo- and pleurocystidia. Pileipellis somewhat duplex-like,
the hypoderm is poorly developed but the hyphae beneath
the epicutis are hypoderm-like (elements that are short and
wide).
Ecology and Distribution: In the Northern and Southern
Hemisphere with coniferous and deciduous trees.
Notes: This small bihemispheric genus is easy to rec-
ognize by the unique combination of small basidiospores
(6–9 × 3.5–5μm) and presence of cheilo- and pleurocystidia.
The basidiomata are medium- to large-sized, stipitocarpic
and agaricoid (phlegmacioid/cortinarioid). The pileipellis is
somewhat duplex. The species form a well-supported line-
age (BS 99%) distinct from the other genera of Cortinari-
aceae, a relationship already recovered by the phylogenetic
analysis of Stensrud etal. (2014) and Soop etal. (2019), and
we here describe the genus as new.
Cystinarius subgen. Cystinarius
IndexFungorum IF552339
Currently included sections: Cystinarius.
Description: Basidiomata medium- to large-sized, agari-
coid (cortinarioid), development type stipitocarpic. Pileus
1.5–8cm, at first hemispherical, then convex to plano-con-
vex, surface fibrillose, orange yellow, yellow brown, greyish
brown, red brown to blackish brown, in some species with
spots, dry or viscid. Lamellae crowded to medium spaced,
adnate to adnexed to emarginate, white to greyish yellow.
Stipe 3–8cm long, 0.7–2cm wide at the apex, cylindrical to
clavate, white, pale grey to greyish yellow, staining yellow
or pinkish, dry to somewhat viscid. Universal veil white,
pale grey to ochraceous yellow, in some species darkening,
often sparse, forming thin bands on the stipe. Context in the
whole basidiomata white to yellowish brown, in some spe-
cies darker in the pileus, staining more or less yellow when
exposed. Odour in lamellae indistinct. KOH reaction nega-
tive. Basidiospores 6–9 × 3.5–5μm, ellipsoid to amygdaloid
to subfusoid, very finely and indistinctly verrucose. Lamel-
lae with cylindrical, clavate or capitate cheilo- and pleuro-
cystidia. Pileipellis somewhat duplex-like, the hypoderm is
poorly developed but the hyphae beneath the epicutis are
hypoderm-like (elements that are short and wide).
Ecology and Distribution: In the Northern and Southern
Hemisphere with coniferous and deciduous trees.
Notes: This small bihemispheric subgenus includes
medium- to large-sized, stipitocarpic, agaricoid (cortinari-
oid) species whose context stains more or less yellow when
exposed. Small basidiospores and the presence of cheilo-
and pleurocystidia is also typical. The pileus is dry to viscid
and the stipe is dry and the pileipellis is somewhat duplex
with a poorly developed hypoderm. The species of the sister
group, C. subgen. Crassi, lack bright yellow colours in their
basidiomata.
Cystinarius subgen. Crassi Niskanen & Liimat., subgen.
nov.
IndexFungorum IF552493
Current name of the type species: Cystinarius eutactus
(Soop) Niskanen & Liimat., comb. nov. IF552494. Basionym
of the type species: Cortinarius eutactus Soop, N.Z. Jl Bot.
43(2): 552. 2005. Holotype: PDD 78807.
Etymology: Named after C. crassus, a species belonging
to this subgenus.
Currently included sections: Crassi.
Description: Basidiomata (medium- to) large-sized,
agaricoid (phlegmacioid), development type stipitocarpic.
Pileus 4–11cm, at first hemispherical, then convex to
plano-convex, surface fibrillose to tomentose, ochraceous
brown, greyish brown to dark red brown, dry or some-
what viscid. Lamellae crowded, adnate to adnexed to
emarginate, white to yellowish grey. Stipe 4–11cm long,
Fungal Diversity
1 3
Fig. 4 Photos of the representatives of Cortinariaceae. A Calonarius
subgen. Calochroi, C. metarius TN 06-268 (H), B C. subgen. Calon-
arius, C. odorifer TN 05-138 (H), C C. subgen. Fulvi, C. sp. TN
11-128 (H), D Aureonarius limonius, TN 07-282 (H), E Cystinarius
rubiginosus TN 12-223 (H), F Hygronarius renidens TN 05-197 (H),
G Mystinarius lustrabilis TN 05-218 (H), H. Volvanarius olivaceo-
vaginatus K235015. Photos AF K. Liimatainen, H R. Healy
Fungal Diversity
1 3
1–2.5cm wide at the apex, fusiform, cylindrical to clavate,
white, in one species becoming brownish red from the
apex, dry. Universal veil white, ochraceous to red brown,
sparse. Context in stipe white, in pileus very pale brown
to brown. Odour in lamellae indistinct. KOH reaction
negative. Basidiospores 6.5–9 × 3.5–4.5μm, ellipsoid to
amygdaloid, almost smooth to very finely and indistinctly
verrucose. Lamellae with cylindrical or clavate cheilo- and
pleurocystidia. Pileipellis somewhat duplex-like, the hypo-
derm is poorly developed but the hyphae beneath the epi-
cutis are hypoderm-like (elements that are short and wide).
Ecology and Distribution: In the Northern and Southern
Hemisphere with coniferous and deciduous trees.
Notes: The members of this small bihemispheric subge-
nus have medium- to large-sized, stipitocarpic, agaricoid
(phlegmacioid) basidiomata with dry to somewhat vis-
cid, ± brown pileus and a white, dry stipe. Small, narrow
basidiospores and the presence of cheilo- and pleurocys-
tidia is also typical. The pileipellis is somewhat duplex
with a poorly developed hypoderm. The species of the
sister subgenus Cystinarius differ by having bright colours
at least in some parts of their basidiomata and a context
that stains more or less yellow when exposed.
Hygronarius Niskanen & Liimat., gen. nov.
IndexFungorum IF552519
Current name of the type species: Hygronarius renidens
(Fr.) Niskanen & Liimat., comb. nov. IF552520. Basionym
of the type species: Cortinarius renidens Fr., Epicr. syst.
mycol. (Upsaliae): 308 (1838) [1836–1838]. Lectotype:
Batsch, Elench. Fung., tab. 6: 23. 1783 (lectotypus hic des-
ignatus, IF552494). Epitypus: Finland, Varsinais-Suomi;
Lohja, herb-rich spruce forest, on calcareous ground, 20
Aug 2000, coll. I. Kytövuori 00–021, H 6107047 (epitypus
hic designatus IF552495), GenBank No. OL958653 (ITS).
Etymology: Derived from the word hygrophanous, since
the species of this genus have a hygrophanous pileus, and
the generic name Cortinarius.
Currently included subgenera: Hygronarius and Visincisi
(Fig.4).
Description: Basidiomata small- to medium-sized, aga-
ricoid (telamonioid), development type stipitocarpic. Pileus
1–6cm, at first somewhat hemispherical or conical, then
convex to plano-convex, with or without an umbo, yel-
low–brown to red-brown, dry to viscid, hygrophanous.
Lamellae medium spaced to almost crowded, adnate to
emarginate, pale brown to rusty brown. Stipe 2.5–9cm
long, 0.3–0.8cm wide at the apex, cylindrical to some-
what clavate; greyish white, pale brown, brownish yellow
to brown, in some species covered by silky-white fibrils
when young. Universal veil white to yellow–brown, sparse
to distinct. Context ± brown. Odour in lamellae indistinct
or slightly raphanoid. Basidiospores 6–10 × 4.5–6 μm,
subglobose, broadly ellipsoid to ellipsoid, finely to coarsely
verrucose. Cystidia absent. Pileipellis duplex, hypoderm
developed.
Ecology and Distribution: In the Northern and Southern
Hemisphere with deciduous and coniferous trees.
Notes: This small bihemispheric genus includes small- to
medium-sized, stipitocarpic, agaricoid (telamonioid) spe-
cies with yellow–brown to red-brown colours. The stipe is
dry and the pileus is dry or viscid and hygrophanous. The
basidiospores are subglobose or ellipsoid and the pileipellis
is duplex with a more or less developed hypoderm. The spe-
cies are morphologically reminiscent of those in Cortinarius
subgenus Iodolentes and Telamonia but are genetically dis-
tinct from them (Garnica etal. 2005; Stensrud etal. 2014)
and for a well-supported clade (BS 92%) in our phylogenetic
analysis. Thus, we here recognize them as their own genus.
Hygronarius subgen. Hygronarius
IndexFungorum IF552521
Currently included sections: Hygronarius (= C. sect.
Renidentes Moënne-Locc. & Reumaux).
Description: Basidiomata small- to medium-sized, aga-
ricoid (telamonioid), development type stipitocarpic. Pileus
1.5–6cm, at first somewhat hemispherical, then convex to
plano-convex, sometimes with an umbo, red-brown, dry to
viscid, hygrophanous. Lamellae medium spaced to almost
crowded, adnate to emarginate, pale brown to rusty brown.
Stipe 2.5–7cm long, 0.3–0.6cm wide at the apex, cylin-
drical (to somewhat clavate); greyish white, pale brown to
brownish yellow, in some species covered by silky-white
fibrils when young. Universal veil absent or very sparse.
Context ± brown. Odour in lamellae indistinct or slightly
raphanoid. Basidiospores 6–7 × 4.5–6μm, subglobose to
broadly ellipsoid, finely verrucose. Cystidia absent. Pileipel-
lis duplex, hypoderm developed.
Ecology and Distribution: In the Northern and Southern
Hemisphere with deciduous and coniferous trees.
Notes: Typical for this small bihemispheric subgenus
are small- to medium-sized, stipitocarpic, agaricoid (tela-
monioid) basidiomata with red-brown colours, absent or
sparse universal veil and subglobose to broadly ellipsoid
basidiospores. They can most easily be distinguished from
the species of H. subgen. Viscincisi by the size of the basidi-
ospores: the basidiospores of Viscincisi species are larger,
7–10 × 4.5–6μm.
Hygronarius subgen. Viscincisi Niskanen & Liimat., sub-
gen. nov.
IndexFungorum IF552522
Current name of the type species: Hygronarius viscincisus
(Soop) Niskanen & Liimat., comb. nov. IF552523 Basionym
of the type species: Cortinarius viscincisus Soop, Australas.
Mycol. 31: 6 (2013). Holotype: PDD 97544.
Fungal Diversity
1 3
Fig. 5 Photos of the representatives of genera Phlegmacium and
Thaxterogaster. A Phlegmacium subgen. Phlegmacium, P. saginum
TN 05-232 (H), B P. subgen. Phlegmacium, P. largum TN 08-060
(H), C. P. subgen. Bulbopodium, P. olivaceodionysae TN 06-311 (H),
D P. subgen. Cyanicium, P. violaceorubens TN 07-062 (H), E T. sect.
Lustrati, T. leucophanes TN 05-161 (H), F T. subgen. Variegati, T.
variegatus TN 05-182 (H), G T. sect. Vibratiles, T. sp TN 05-210 (H),
H T. subgen. Scauri, T. subpurpurascens TN 08-059 (H). Photos K.
Liimatainen
Fungal Diversity
1 3
Etymology: Named after the type species of this genus.
Currently included sections: Austroduracini, Viscincisi.
Description: Basidiomata small- to medium-sized, aga-
ricoid (telamonioid), development type stipitocarpic. Pileus
1–6cm, at first somewhat hemispherical or conical, then
convex to plano-convex, with or without an umbo, yel-
low–brown to red-brown, dry to viscid, hygrophanous.
Lamellae medium spaced to almost crowded, adnate to
emarginate, pale brown to rusty brown. Stipe 2.5–9cm
long, 0.3–0.8cm wide at the apex, cylindrical to some-
what clavate; greyish white, pale brown, brownish yellow
to brown, in some species covered by silky-white fibrils
when young. Universal veil white to yellow–brown, sparse
to distinct. Context ± brown. Odour in lamellae indistinct or
slightly raphanoid. Basidiospores 7–10 × 4.5–6μm, ellip-
soid, moderately to coarsely verrucose. Cystidia absent.
Pileipellis duplex, hypoderm developed.
Ecology and Distribution: In the Southern Hemisphere in
forests of Nothofagaceae.
Notes: The members of this small subgenus only occur in
the Southern Hemisperic Nothofagaceae forests. The basidi-
omata are small- to medium-sized, stipitocarpic, agaricoid
(telamonioid) with yellow–brown to red-brown colours,
sparse to distinct universal veil and ellipsoid basidiospores.
They can most easily be distinguished from the species of
H. subgen. Hygronarius by the basidiospores: the basidi-
ospores of Hygronarius species are smaller, 6–7 × 4.5–6μm
and subglobose to broadly ellipsoid.
Mystinarius Niskanen & Liimat., gen. nov.
IndexFungorum IF552524
Current name of the type species: Mystinarius lustra-
bilis (Moënne-Locc.) Niskanen & Liimat., comb. nov.
IF552525. Basionym of the type species: Cortinarius lustra-
bilis Moënne-Locc., in Bidaud, Moënne-Loccoz, Reumaux
& Henry, Atlas des Cortinaires (Meyzieu) 10: 492 (2000).
Holotype: PC 1226.
Etymology: Derived from the latin word “mysticus” and
the generic name Cortinarius.
Currently included subgenera: Mystinarius (Fig.4).
Description: Basidiomata medium-sized, agaricoid
(myxacioid/phlegmacioid), development type stipitocarpic.
Pileus 3–6cm, at first hemispherical, then low convex, rimy
fibrillose, brownish yellow to reddish brown, somewhat
viscid to almost dry. Lamellae crowded to medium spaced,
emarginate, yellowish brown to brown. Stipe 5–10cm long,
0.7–1.2cm wide at the apex, cylindrical or tapering down-
wards to somewhat clavate, silky fibrillose, white to yellow,
dry. Universal veil white, sparse, presumably somewhat vis-
cid. Context yellow. Odour in lamellae indistinct or sweet-
ish. Taste in pileus context somewhat bitter. Basidiospores
8.5–9.5 × 5–6μm, ovoid to amygdaloid, finely to moderately
verrucose. Pileipellis duplex, hypoderm developed.
Ecology and Distribution: In Northern and Southern
Hemisphere in coniferous and Nothofagaceae forests.
Notes: The species of this small, bihemispheric genus
have medium-sized, stipitocarpic, agaricoid (myxacioid/
phlegmacioid) basidiomata with a yellow to reddish brown,
somewhat viscid to almost dry pileus and a white to yel-
low, dry stipe. The basidiospores are medium-sized and
the pileipellis is duplex. The species of this genus resemble
morphologically most of those in Thaxterogaster subgenus
Multiformes, T. sect. Pinophili or T. sect. Vibratiles. How-
ever, they are not closely related to Thaxterogaster or other
genera of Cortinariaceae and we here propose a new genus,
Mystinarius, for them.
Phlegmacium (Fr.) Wünsche, Die Pilze: 87, 128 (1877)
em. Niskanen & Liimat.
Basionym: Agaricus trib. Phlegmacium Fr., Syst. mycol.
(Lundae) 1: 10, 227 [‘217’] (1821).
Sanctioning citation: Fr., Syst. mycol. 1: 10 (1821).
Current name of the type species. Phlegmacium saginum
(Fr.) Niskanen & Liimat., comb. nov. IF552795. Basionym of
the type species: Agaricus saginus Fr., Syst. mycol. (Lundae)
1: 226 (1821). Neotype: IB 19770098, in Melot, Docum.
Mycol. XVI (63–64): 130, (1986).
Synonyms: Bulbopodium Earle, Bull. New York Bot.
Gard. 5: 441 (1909). Current name of the type species:
Phlegmacium caerulescens (Schaeff.) Wünsche, Die Pilze:
131 (1877). Basionym of the type species: Agaricus caer-
ulescens Schaeff., Fung. bavar. palat. nasc. (Ratisbonae) 4:
17 (1774).
Cyanicium Locq., Fl. Mycol., 3. Cortinariales-A.: 146
(1979) [1977]. Current name of the type species: Phleg-
macium cyanites (Fr.) M.M. Moser, Die Gatt. Phlegm.: 337
(1960). Basionym of the type species: Cortinarius cyanites
Fr., Epicr. syst. mycol. (Upsaliae): 279 (1838) [1836–1838].
Meliderma Velen., České Houby 2: 399 (1920). Current
name of the type species: Phlegmacium mussivum (Fr.) Nis-
kanen & Liimat., comb. nov. IF552796. Basionym of the
type species: Agaricus mussivus Fr., Epicr. syst. mycol.
(Upsaliae): 178 (1838) [1836–1838].
Currently included subgenera: Phlegmacium, Bulbopo-
dium, Carbonella and Cyanicium (Fig.5).
Description: Basidiomata medium- to large-sized, rarely
small, agaricoid (phlegmacioid, rarely telamonioid), some
species sequestrate, development type stipitocarpic or pile-
ocarpic. Pileus (1–)3–12(–20) cm, at first hemispherical to
convex, low convex to almost plane when old, rarely with a
broad umbo; surface in some species innately fibrillose or
radially wrinkled, rarely scaly or with patches of veil; from
white, pale ochraceous and yellow to dark brown and umber
with yellow, orange, red, greyish, greenish or purplish tints,
in some species completely purple; dry, viscid or glutinous.
Lamellae in the vast majority of species crowded, in some
Fungal Diversity
1 3
species medium spaced, emarginate, at first pale greyish
white to purple, later pale brown to purplish brown, darker
in the species of subgenus Carbonella. Stipe 3–13(–20) cm
long, (0.4–)0.8–2(–3) cm wide at the apex, up to 5cm at
base, clavate to bulbous with a rounded or a marginate bulb,
or cylindrical to rooting, in the vast majority of the species
more or less white, sometimes with purplish tints, in some
species grey, dry. Universal veil white, grey, yellow, greenish
yellow, more or less brown or purple, sparse to abundant,
forming incomplete and complete girdles on the stipe. Con-
text in many species in pileus and stipe white, sometimes
with purplish colours, in some species grey, yellow-greenish,
olive-grey, pale greyish purple to vinaceous brown, in the
species of subgenus Cyanicium becoming vinaceous red
on exposure. Odour in lamellae indistinct or in some spe-
cies raphanoid, earthy, grassy, rubbery, sweetish, fruity or
farinaceous. KOH reaction in pileus context negative, yel-
low, orange, reddish lilac, olivaceous or pale brown. Basidi-
ospores 7–12.5(–17) × 4–8.5(–10) μm, amygdaloid, ellipsoid
or citriform, in some species subglobose, finely to strongly
verrucose. Pileipellis duplex, simplex in P. subgen. Cyani-
cium and some lineages of P. subgen. Phlegmacium, epicutis
in many species ± gelatinous.
Ecology and Distribution: In the Northern Hemisphere
with the species of Fagales, Pinaceae and Tilia. In the
Southern Hemisphere at least in Nothofagaceae forests.
The centre of the diversity is in the Northern Hemisphere:
two of the four subgenera, Bulbopodium and Cyanicium,
are only know from the Northern Hemisphere and the vast
majority of the species of the P. subgenus Bulbopodium are
also boreal.
Notes: This genus includes many of the species tradi-
tionally placed in the Cortinarius subgenus Phlegmacium.
Typical for the species are a dry stipe and viscid to glutinous
pileus, or if dry, then the KOH reaction in the context of
the pileus is usually yellow. Most species have a pileipel-
lis duplex with a more or less developed hypoderm but the
species of the subgenus Cyanicium and some lineages of P.
subgen. Phlegmacium have a simplex pileipellis. The spe-
cies of the genus Calonarius that were previously included
in this group can be distinguished from the species of the
genus Phlegmacium by the combination of pileocarpic
basidiomata, a marginated bulb and simplex pileipellis. For
the phlegmacioid species in the genus Thaxterogaster, a dis-
tinguishing combination of characters that would work for
all groups is harder to give, but as a rule the phlegmacioid
species encountered in the Southern Hemisphere mainly
belong to the genus Thaxterogaster and the lineages in the
Northern Hemisphere that can be confused with the spe-
cies of the genus Phlegmacium are subgenera Multiformes,
Riederorum, Scauri and Varieagati and sections Pinophili
and Vespertini. Some phlegmacioid lineages also exist in the
genus Cortinarius, namely Infracti and Subtorti, but they
have stipitocarpic basidiomata and round spores. The species
of the small Southern Hemispheric genus Volvanarius have a
phlegmacioid appearance as well, but they are small in size
and the majority of the species have a volva. Some species of
the genus Cystinarius may also be confused with the species
of the genus Phlegmacium, but Cystinarius species have dis-
tinct cheilo- and pleurocystidia and a dry pileus. Lastly, the
basidiomata of the genus Austrocortinarius resemble those
in P. subgenus Phlegmacium, sect. Arguta and clades/Obso-
leti and Caligati but those lineages of Phlegmacium are only
known from the Northern Hemisphere whereas the genus
Austrocortinarius occurs in the South Pacific.
Phlegmacium subgen. Phlegmacium
Currently included sections: Phlegmacium, Arguta,
Claricoloria, Elastica, Obsoleta, Phlegmacioida, Percomia,
Rhizophora, Seraria, and Varia as well as clade/Caligata.
Notes: The centre of the diversity of this species-rich
lineage is in the Northern Hemisphere but some members
of the group are also encountered in the Southern Hemi-
sphere. Basidiomata are medium- to large-sized, predomi-
nantly stipitocarpic, agaricoid (phlegmacioid). The stipe is
dry, and the pileus is viscid to glutinous, or if dry then the
KOH reaction in the context of the pileus is usually yellow.
The pileipellis is either duplex or simplex. The members of
the other species-rich subgenus, Bulbopodium, have mainly
pileocarpic basidiomata.
Phlegmacium subgen. Bulbopodium (Earle) Niskanen &
Liimat., comb. & stat. nov.
IndexFungorum IF552797
Basionym: Bulbopodium Earle, Bull. New York Bot.
Gard. 5: 441 (1909).
Current name of the type species: Phlegmacium caer-
ulescens (Schaeff.) Wünsche, Die Pilze: 131 (1877). Bas-
ionym of the type species: Agaricus caerulescens Schaeff.,
Fung. bavar. palat. nasc. (Ratisbonae) 4: 17 (1774). Lecto-
type: Schaeffer, Fung. Bav. 1: Tab. 34, figs I, II, III, 1762, in
Brandrud etal., Cortinarius Flora Photographica II, pl. B11
(1992). Epitype: S F-44815, in Liimatainen etal., Persoonia
33:118 (2014).
Currently included sections: Amoenolentia, Arcifolia,
Aureocistophila, Bulbopodium (= Caerulescentes ss. Soop
etal. 2019), Caerulea (= /Eucaerulei ss. Soop etal. 2019),
Caerulescentia (= /Camptori ss. Soop etal. 2019), Dionysae,
Glaucocephala, Glaucopodes, Subhymenogaster, and Taura.
Notes: Representatives of this species-rich subgenus
are thus far only known from the Northern Hemisphere.
Basidiomata are medium- to large-sized, predominantly
pileocarpic, agaricoid (phlegmacioid) or rarely sequestrate.
The pileus is viscid to glutinous, and the stipe is dry. The
Fungal Diversity
1 3
members of the other species-rich subgenus, Phlegmacium,
have mainly stipitocarpic basidiomata and some species also
have a yellow to pale brown KOH reaction in the pileus
context.
Phlegmacium subgen. Carbonella (Soop) Niskanen &
Liimat., comb. & stat. nov.
IndexFungorum IF553054
Basionym: Cortinarius sect. Carbonelli Soop, in Soop,
Dima, Cooper, Park & Oertel, Persoonia 42: 286 (2019).
Current name of the type species: Phlegmacium carbonel-
lum (Soop) Niskanen & Liimat., comb. nov. IF552947.
Basionym of the type species: Cortinarius carbonellus
Soop, Bull. Soc. mycol. Fr. 117(2): 120 (2001). Holotype:
PDD 70502.
Currently included sections: Carbonella.
Notes: The species of this subgenus occur in the Nothof-
agaceae forests of New Zealand. The species are charac-
terized by small-sized, agaricoid (telamonioid), stipito-
carpic basidiomata with dark grey-brown to bluish-grey,
or purple-brown to umber colours and dry, hygrophanous
pileus and dry stipe. The universal veil is pale grey-brown,
purple to pale brownish-red and sparse. The alkaline reac-
tion is orange to reddish-lilac in the context and red on
lamellae (Soop 2017; Soop etal. 2019). Morphologically
the species of this subgenus are most reminiscent of those
in Cortinarius subgenus Telamonia (dry pileus and stipe,
development type of the basiodiomata stipitocarpic) but
phylogenetically the subgenus is most closely related to
the genus Phlegmacium (pileus viscid to glutinous, or if
dry then KOH-reaction usually yellow, stipe dry, devel-
opment type of the basiodiomata stipito- or pileocarpic).
Since they represent a well-supported lineage within the
genus Phlegmacium and are morphologically distinct from
their closest relatives we here recognize them in a subge-
neric level.
Phlegmacium subgen. Cyanicium (Locq.) Niskanen &
Liimat., comb. & stat. nov.
IndexFungorum IF553369
Basionym: Cyanicium Locq., Fl. Mycol., 3. Cortinari-
ales-A.: 146 (1979) [1977].
Current name of the type species: Phlegmacium
cyanites (Fr.) M.M. Moser, Die Gatt. Phlegm.: 337 (1960).
Basionym of the type species: Cortinarius cyanites Fr.,
Epicr. syst. mycol. (Upsaliae): 279 (1838) [1836–1838].
Neotype: S, Taylor 2005069, in Liimatainen etal., Persoo-
nia 33:118 (2014).
Currently included sections: Cyanicium.
Notes: This small Northern Hemispheric subgenus
includes species with a unique combination of charac-
ters: basidiomata are (medium- to) large-sized, agari-
coid (phlegmacioid) and stipitocarpic with greyish-blue,
greyish-brown to darker purplish-brown colors. The pileus
is at first viscid but soon dry. A context that becomes vina-
ceous-red on exposure and lamellar trama hyphae with
abundant small to large to worm-like blood red guttules
in Melzer’s reagent is typical. The KOH reaction is nega-
tive in the context and the pileipellis structure is simplex.
Thaxterogaster Singer, Mycologia 43(2): 216 (1951) em.
Niskanen & Liimat.
Current name and basionym of the type species: Thax-
terogaster magellanicus Singer [as ‘magellanicum’], Myco-
logia 43(2):219. 1951. Neotype: Argentina, Tierra del Fuego,
Ushuaia, los Humedales, under Nothofagaceae, 16 Feb
2015, L. Domínguez LSD2380b, CORDC00006504, Gen-
Bank no. MN855076 (ITS), in Nouhra etal., Mycologia 113:
1040 (2021).
Synonyms: Gigasperma E. Horak, N.Z. Jl Bot. 9(3): 491
(1971). Current name of the type species: Thaxterogaster
crypticus (E. Horak) Niskanen & Liimat.
Hygramaricium Locq., Fl. Mycol., 3. Cortinariales-
A.: 146 (1979) [1977]. Current name of the type species:
Thaxterogaster causticus (Fr.) Niskanen & Liimat., comb.
nov. IF552948. Basionym of the type species: Cortinarius
causticus Fr., Epicr. syst. mycol. (Upsaliae): 270 (1838)
[1836–1838]. Neotype: 6031321 (H), in Niskanen etal.,
Index Fungorum 477: 3 (2021).
Rapacea E. Horak, Kew Bull. 54(3): 789 (1999). Current
name of the type species: Thaxterogaster mariae (E. Horak)
Niskanen & Liimat., comb. nov. IF552949. Basionym of the
type species: Rapacea mariae E. Horak, Kew Bull. 54(3):
789 (1999). Holotype: PDD 69747.
Currently included subgenera: Thaxterogaster, Cretaces,
Multiformes, Riederorum, Scauri, and Variegati (Fig.5).
Description: Basidiomata small- to large-sized, agaricoid
(phlegmacioid, myxacioid) or sequestrate, development type
stipitocarpic to pileocarpic. Pileus 1–12cm, at first hemi-
spherical, then convex to plano-convex; surface smooth,
innately fibrillose or somewhat scaly; ± white, ± yellow,
pale to dark brown with greyish, ochraceous or reddish tints,
umber to blackish, ± purple or with a purplish tint, some
species with olivaceous colours; dry, viscid or glutinous,
with hygrophanous spots or streaks or non-hygrophanous.
Lamellae crowded to medium spaced, adnate, adnexed
or emarginate; when young white, pale grey, pale brown,
green, ± purple or with a purplish tint. Stipe 4–13cm long,
0.3–2.5cm wide at the apex, up to 5cm at the base; cylindri-
cal, clavate, rooting or bulbous, bulb rounded to ± margin-
ate; white, pale brown, ± purple, some species with greenish
colours, in species of sect. Purpurascentes usually turning
deeper purple when bruised, dry. Universal veil white, pur-
plish white, purple, in some species turning pink; sparse
to more abundant, in pileocarpic species at the bulb mar-
gin and pileus margin, in stipitocarpic species forming a
Fungal Diversity
1 3
thin sock-like sheet or incomplete and complete girdles on
the stipe, usually dry but in some species viscid. Context
in pileus white, pale brown, purple or dark blue/blackish
green, in stipe white, ± yellow, pale brown, ± purple or with
purplish or green/olivaceous colours. Odour in lamellae
indistinct, in the context of the stipe indistinct, honey-like
or sweet, garlic-like in T. crypticus. KOH reaction negative
or red in pileus, context and/or stipital veil. Basidiospores
6–18 × 3.5–9μm, subglobose, amygdaloid, fusoid to ellip-
soid, finely to strongly verrucose (subglobose, smooth and
very large in T. crypticus 25–35μm in diam.). Cystidia
absent. Pileipellis duplex, hypoderm present.
Ecology and Distribution: In the Northern Hemisphere
with species of Fagaceae, Betulaceae, Tilia and Pinaceae.
In the Southern Hemisphere in Nothofagaceae forests.
Notes: The species of this bihemispherical genus have
traditionally been classified in phlegmacioid and myxa-
cioid taxa in genus Cortinarius or in sequestrate genera/
taxa. The size of the basidiomata ranges from small to large
and vary in coloration from white, ochraceous, greenish,
brown to purple. Typical for all agaricoid species, however,
is a pileipellis duplex and a negative or, more rarely, red (in
pileus, context and/or stipital veil) KOH reaction. Several
lineages of this genus have a honey-like or sweet smell in the
context, not typical in other genera of the family Cortinari-
aceae and otherwise known only in Cortinarius subgenus
Myxacium. The development type of basidiomata ranges
from stipitocarpic to pileocarpic.
Thaxterogaster subgen. Thaxterogaster
Currently included sections: Alboaggregati,
Thaxterogaster.
Description: Basidiomata medium- to large-sized, aga-
ricoid (phlegmacioid) or sequestrate, development type
stipitocarpic. Pileus 3.5–7.5(–12) cm, at first hemispheri-
cal, then convex to plano-convex, with an umbo, finely
fibrillose, white, yellow–brown or with purplish tint, dry to
viscid. Lamellae crowded, adnate to adnexed to emargin-
ate, white to greyish white. Stipe 4–10cm long, 1–2cm
wide at the apex, cylindrical to rooting; white. Universal
veil white, rather abundant in agaricoid species, peronate,
sometimes forming a small collar. Context white. Odour in
lamellae indistinct. Basidiospores 10.5–17.5 × 5.5–9.5μm,
fusoid-amygdaloid to ellipsoid, moderately verrucose. Cys-
tidia absent.
Ecology and Distribution: In New Zealand and South
America, in Nothofagaceae forests.
Notes: This small Southern Hemispheric subgenus
includes medium- to large-sized, stipitocarpic, agaricoid
(phlegmacioid) and sequestrate species. The pileus is viscid
and the stipe is cylindrical to rooting. The species resem-
ble those in South American T. subgenus Cretaces but the
species of subgenus Cretaces have smaller basidiospores
(6–10 × 3–5.5μm) and a sparse universal veil.
Thaxterogaster subgen. Cretaces (Soop & Dima) Nis-
kanen & Liimat., comb. et stat. nov.
IndexFungorum IF552950
Basionym: Cortinarius sect. Cretaces Soop & Dima,
in Soop, Dima, Cooper, Park & Oertel, Persoonia 42:279.
2019. Current name of the type species: Thaxterogaster
cretax (Soop) Niskanen & Liimat., comb. nov. IF552951.
Basionym of the type species: Cortinarius cretax Soop, Bull.
Soc. Mycol. Fr. 118(3):185. 2003. (2002). Holotype: PDD
73148.
Currently included sections: Cretaces.
Notes: The representatives of this small subgenus are only
encountered in the Southern Hemisphere. The basidiomata
are medium- to large-sized, stipitocarpic, agaricoid (phleg-
macoid) with a viscid pileus and a dry stipe. The colour of
the pileus ranges from white to yellow–brown, lamellae are
white to pale grey and crowded, the stipe is usually rooting
and the universal veil is sparse. The odour in the lamellae is
indistinct or marzipan-like. Basidiospores are fusoid-amyg-
daloid, 6–10 × 3–5.5μm and weakly verrucose. The species
are most reminiscent of those in T. subgen. Thaxterogaster
but the agaricoid species in subgenus Thaxterogaster have
larger basidiospores and more abundant universal veil.
Thaxterogaster subgen. Multiformes Niskanen & Liimat.,
subgen. nov.
IndexFungorum IF552952
Current name of the type species: Thaxterogaster caesio-
phylloides (Kytöv., Liimat., Niskanen, Brandrud & Frøslev)
Niskanen & Liimat., comb. nov. IF552953. Basionym of the
type species: Cortinarius caesiophylloides Kytöv., Liimat.,
Niskanen, Brandrud & Frøslev, in Liimatainen, Niskanen,
Dima, Kytövuori, Ammirati & Frøslev, Persoonia 33: 130.
2014. Holotype: H 6029792.
Etymology: Named after C. multiformis, a species belong-
ing to this subgenus.
Currently included sections: Multiformes.
Description: Basidiomata medium- to large-sized, agari-
coid (phlegmacioid), development type pileocarpic. Pileus
4–12cm, at first hemispherical, then convex to plano-con-
vex, in some species innately fibrillose, white, cream-col-
oured, pale yellow, ochraceous yellow, orange-yellow, grey-
brown or red brown, rarely bluish brown, viscid to glutinous,
with hygrophanous spots or streaks. Lamellae crowded to
almost crowded, emarginate, at first greyish white, later very
pale greyish brown, in some species with a bluish tint. Stipe
4–13cm long, 0.7–2cm wide at the apex, up to 3.5cm at the
base, clavate to bulbous, bulb rounded to slightly marginate,
rarely almost cylindrical, at first white, later pale brown, in
some species with a bluish tint at the apex, dry. Universal
Fungal Diversity
1 3
veil white, sparse, at the bulb margin, rarely somewhat vis-
cid. Context in pileus white or pale brown to brown near
the pileus surface, in stipe white, in some species with a
bluish tint at the apex of the stipe. Odour in the flesh of the
bulb and/or stipe honey-like. KOH reaction negative. Basidi-
ospores 7.5–11 × 4.5–6.5μm, ovoid-amygdaloid, amygda-
loid, fusoid to ellipsoid, finely to rather strongly verrucose.
Cystidia absent. Pileipellis duplex, epicutis with a glutinous
layer on the top, hypoderm present.
Ecology and Distribution: In the Northern Hemisphere
with coniferous (Pinaceae) and deciduous trees (Fagaceae,
Betulaceae).
Notes: The species of this Northern Hemispheric sub-
genus are medium- to large-sized and phlegmacioid with
a viscid to glutinous pileus and a dry stipe. They are char-
acterised by having a pileocarpic development type of the
basidiomata, pileipellis duplex, greyish-white lamellae when
young and a honey-like smell in the context. In addition, the
colour of the pileus ranges from cream-coloured to yellow-
ochraceous to red-brown and the stipe is white. Some spe-
cies have bluish tints in their basidiomata. The phylogenetic
analysis of Soop etal. (2019) indicated that morphologically
similar species of T. sect. Cremeolini could also belong to
this subgenus as well as species of sect.Malvacei Moser, but
the group did not receive strong support.
Thaxterogaster subgen. Riederorum Niskanen & Liimat.,
subgen. nov.
IndexFungorum IF553567
Current name of the type species: Thaxterogaster mal-
achioides (P.D. Orton) Niskanen & Liimat., comb. nov.
IF553568. Basionym of the type species: Cortinarius mala-
chioides P.D. Orton, Naturalist, Leeds (Suppl.): 148. 1958.
Holotype: K(M) 94426.
Etymology: Named after C. riederi, a species belonging
to this subgenus.
Currently included sections: Riederorum.
Description: Basidiomata medium- to large-sized, agari-
coid (phlegmacioid), development type pileocarpic. Pileus
2.5–12cm, at first hemispherical, then convex to plano-
convex, innately fibrillose; cream-coloured, greyish white,
pale grey, yellow ochraceous, ochraceous brown, fulvous
brown or grey-brown, sometimes with an olivaceous tint;
viscid to glutinous, in some species with hygrophanous spots
or streaks. Lamellae crowded, emarginate, at first violet-
blue, later greyish brown. Stipe 5–12cm long, 0.7–2.5cm
wide at the apex, up to 5cm at the base; clavate to bulbous,
bulb rounded to ± marginate; at first white with a bluish
tint to completely bluish violet, later greyish white to pale
ochraceous brown, becoming ± brown if damaged, bruised
or with age, dry. Universal veil bluish white, very sparse,
remnants, if visible, near the bulb margin and on the pileus
margin. Context in pileus and bulb bluish white to white, in
stipe bluish white to bluish violet, violet colour fading with
age. Odour in lamellae indistinct. KOH reaction negative.
Basidiospores 10–14.5 × 6–9μm, ellipsoid to amygdaloid,
moderately to strongly verrucose. Cystidia absent. Pileipellis
duplex, epicutis with a glutinous layer on the top, hypoderm
present, well to somewhat developed.
Ecology and Distribution: In the Northern Hemisphere
with coniferous (Pinaceae) and deciduous trees (Fagaceae,
Betulaceae, Tilia).
Notes: The species of this small Northern Hemispheric
subgenus are characterized by having medium- to large-
sized, agaricoid (phlegmacioid), pileocarpic basidiomata
with a viscid to glutinous, innately fibrillose pileus, dry stipe
and bluish-violet colours in the lamellae and stipe. Typical
are also large (> 10μm long) ellipsoid to amygdaloid basidi-
ospores, a negative KOH-reaction and pileipellis duplex. For
a recent morpho-genetic revision of the group see Brandrud
etal. (2018).
Thaxterogaster subgen. Scauri Niskanen & Liimat., sub-
gen. nov.
IndexFungorum IF553569
Current name of the type species: Thaxterogaster herpe-
ticus (Fr.) Niskanen & Liimat., comb. nov. IF553570. Bas-
ionym of the type species: Cortinarius herpeticus Fr., Epicr.
syst. mycol. (Upsaliae): 268. 1838 (1836–1838). Neotype:
S F-44759, in Liimatainen etal., Persoonia 33:119 (2014).
Etymology: Named after C. scaurus, a species belonging
to this subgenus.
Currently included sections: Scauri and Purpurascentes.
Description: Basidiomata medium- to large-sized, agari-
coid (phlegmacioid), in T. sect. Purpurascentes some spe-
cies sequestrate, development type pileocarpic (Scauri) or
somewhat stipitocarpic to stipitocarpic (Purpurascentes).
Pileus 1–10cm, at first hemispherical, then convex to
plano-convex; surface innately fibrillose or not; greyish
white, pale (greyish) ochraceous, pale ochraceous brown,
olivaceous brown, red brown, dark brown, blackish brown,
purplish grey or pale purple; viscid to glutinous, many spe-
cies with hygrophanous spots or veins. Lamellae crowded
to medium spaced, emarginate, greyish brown, green to oli-
vaceous, soon brown, in part of the species with a purplish
tint, completely purple or bluish grey. Stipe 3–12cm long,
0.3–2cm wide at the apex, up to 3.5cm at the base; cylindri-
cal, clavate or more or less bulbous, bulb usually ± margin-
ate; at first pale greyish purple/blue, pale purple to purplish
green, later yellowish grey, yellow–brown or purple, in spe-
cies of sect. Purpurascentes usually turning deeper purple
when bruised, dry. Universal veil green, purple, ochraceous
yellow or white, sparse to more abundant, in pileocarpic
species at the bulb and pileus margin, in stipitocarpic spe-
cies forming a sock/like sheet or incomplete and/or complete
girdles on the stipe. Context in pileus white, brownish white,
Fungal Diversity
1 3
pale brown, purple or dark blue/blackish green, in stipe pale
purple, greenish purple, pale olivaceous to yellow-green.
Odour in the context of the stipe honey-like or sweet in many
species. KOH reaction negative, in a few species blood red
on pileus and/or stipital veil (Soop 2017). Basidiospores
7–12 × 4.5–7μm, broadly ellipsoid, ellipsoid to amygdaloid,
moderately to coarsely verrucose. Cystidia absent. Pileipellis
duplex, epicutis with a glutinous layer on the top, hypoderm
present.
Ecology and Distribution: In the Northern Hemisphere
with coniferous (Pinaceae) and deciduous trees (Fagaceae,
Betulaceae, Tilia), and in the Southern Hemisphere in
Nothofagaceae forests.
Notes: Typical for the species of this bihemispheric sub-
genus are medium- to large-sized, agaricoid (phlegmacioid)
basidiomata with purplish and/or greenish tints/colours and
a pileipellis duplex. The pileus is viscid to glutinous, and
the stipe is dry, and many species have a honey-like or sweet
smell in the context. The iodine (lugol) reaction is posi-
tive in the context and lamellae (Garnica etal. 2005; Soop
etal. 2019). The development type of the basidiomata is
pileocarpic or stipitocarpic, or for at least 14 Australian spe-
cies sequestrate..
Thaxterogaster subgen. Variegati Niskanen & Liimat.,
subgen. nov.
IndexFungorum IF553571
Current name of the type species: Thaxterogaster var-
iegatus (Bres.) Niskanen & Liimat., comb. nov. IF553573.
Basionym of the type species: Cortinarius variegatus Bres.,
Fung. trident. 1(4–5):56. 1884. Lectotype: Bresadola, Fung.
trident. 1(4–5): tab. LXII, 1884 (lectotypus hic designatus,
IF553574). Epitype: Finland, Kuusamo; Oulanka biological
station, dry pine heath forest (Pinus sylvestris) on sandy soil,
20 Sept. 2005, coll. K. Liimatainen & T. Niskanen, 05–182,
H 6031519 (epitypus hic designatus IF553602), GenBank
No. OL985940 (ITS).
Currently included sections: Variegati.
Description: Basidiomata medium- to large-sized, aga-
ricoid (phlegmacioid), development type ± stipitocarpic.
Pileus 3.5–10cm, at first hemispherical then plano-convex;
surface rimy at least when young: red-brown, darker from
the centre; viscid to glutinous, with some hygrophanous
spots. Lamellae crowded, adnexed to emarginate, greyish
white to pale grey. Stipe 5–15cm long, 1–1.5cm wide at the
apex, up to 2cm at the base; cylindrical, clavate to bulbous,
bulb marginate or not; silky-fibrillose, white, dry. Universal
veil at first white, later pink to purplish pink, forming a thin
sheet or some girdles on the 1/3 lowest part of the stipe.
Context white. Odour in lamellae indistinct. KOH reaction
negative. Basidiospores 6–8 × 3–4μm, amygdaloid-fusoid,
smooth to finely verrucose. Cystidia absent. Pileipellis
duplex, hypoderm present.
Ecology and Distribution: In the Northern Hemisphere
with coniferous (Pinaceae) and more rarely with deciduous
trees (Fagaceae).
Notes: The most characteristic features of this mono-
typic boreal subgenus are the initially white universal veil
that becomes pinkish with age and small, almost smooth
amygdaloid-fusoid spores. In addition, the basidiomata are
medium- to large-sized, the development type is ± stipito-
carpic and the pileipellis has a well-developed hypoderm.
The pileus is red-brown, and the stipe is white. Our phylo-
genetic analysis also suggests that T. sect. Turmales could
be included in this subgenus, but the relationship is not well-
supported. Section Turmales includes morphologically simi-
lar species with small (< 9μm long), amygdaloid-fusoid,
finely verrucose spores, and in at least one species of the
section, C. turmalis, the mycelium becomes rose-coloured
after exposure to air.
Volvanarius Niskanen & Liimat., gen. nov.
IndexFungorum IF553603
Current name of the type species: Volvanarius chloros-
plendidus (Furci, Niskanen, San-Fabian, Liimat. & Salgado
Salomón) Niskanen & Liimat., comb nov. IF553604. Basio-
nym of the type species: Cortinarius chlorosplendidus Furci,
Niskanen, San-Fabian, Liimat. & Salgado Salomón, in Lii-
matainen, Niskanen, San-Fabian, Mujic, Peintner, Dresch,
Furci, Nouhra, Matheny & Smith, Mycologia 112(2): 335.
2020. Holotype: K(M) 235086.
Etymology: Derived from the word volva, that many spe-
cies of this genus have, and the generic name Cortinarius.
Currently included subgenera: Thaumasti, Volvanarius
(Fig.4).
Description: Basidiomata small- to rather small-sized,
agaricoid (phlegmacioid) or rarely sequestrate, develop-
ment type pileocarpic. Pileus 1.5–6cm, at first hemispheri-
cal, then low convex to almost plane, yellow, ochraceous,
ochraceous brown, orange-brown, olive brown to greenish,
dry or viscid. Lamellae medium crowded to almost crowded,
adnate, at first very pale brownish grey, later pale greyish
brown. Stipe 3–8.5cm long, 0.4–1.2cm wide at the apex,
cylindrical, with a bulbous base (up to 2.5cm), with silky
fibrillose surface, white, pale yellow, or pale greenish. Uni-
versal veil white or ochraceous, in some species with orange
spots, often forming a volva at the base of the stipe. Context
in most species white with ochraceous, greenish or brown-
ish tints, in some species context in pileus brown. Odour
in lamellae indistinct. Chemical reactions: context of the
bulb turns red with ammonia (Moser and Horak 1975; Soop
etal. 2019). Basidiospores 7–11.5 × 4–6.5μm, citriform to
amygdaloid, rarely ellipsoid, finely to strongly verrucose.
Cystidia (cheilocystidia) balloon-shaped, present in some
species. Pileipellis duplex, hypoderm developed.
Fungal Diversity
1 3
Ecology and Distribution: In the Southern Hemisphere
with species of Nothofagaceae.
Notes: This small genus is only known from the South-
ern Hemispheric Nothofagaceae forests. Members of this
group can easily be identified in the field by the small and
Phlegmacium-like basidiomata with a bulbous stipe, and the
universal veil that in most species forms a distinct volva at
the base of the stipe. Typical are also citriform to amygda-
loid, rarely ellipsoid basidiospores and pileipellis duplex.
A few species have balloon-shaped cheilocystidia. For a
recent morphogenetic revision of C. sect. Thaumasti see
Liimatainen etal. (2020b).
Discussion
Which criteria make agood genus?
The primary criterion for recognizing a taxonomic rank such
as genus is a natural, monophyletic group of species that is
supported, for a given phylogenetic analysis. However, the
rank and what other criteria should be used to delimit it are
more or less subjective. Ultimately, the aim is to find a com-
munity consensus for practical solution to describing diver-
sity. At least in species-rich fungal groups, monophyletic
clades with good support often exist at different nested levels
for a given phylogeny and there are, therefore, multiple ways
in which generic limits could be drawn.
A genus is usually also defined by its morphological,
chemical, or ecological characteristics that distinguish it
from its relatives. Our proposed classification is largely in-
line with circumscription of other genera in the same order
based on a combination of phylogenetic, morphological,
chemical, and ecological traits. Although a universal set
of objective criteria is not realistic for classifying all life
with its multitudes of variation, the objective of achieving
both coherence and practicality in recognizing evolutionary
uniqueness of the generic rank within a higher taxon could
be applied at approximately the same level of inclusive-
ness. This ideal exists not only because a basic assumption
is that a certain taxonomic rank reflects a level of cohesion
around a similar set of traits and phylogenetic patterns, but
also because in practice it makes comparisons between dif-
ferent genera across different groups of organisms (e.g., in
ecological, evolutionary, and conservation studies) more
meaningful.
One additional aspect to take into consideration when
delimiting genera is the amount of diversity to be included.
Within reason, monotypic genera should be avoided, since
the general aim of classification above the species level is
to group closely related units together towards increasingly
larger units, so that each taxonomic level, with cumulative
inclusiveness, would deliver information that is something
more than the previous unit. In practise, however, monotypic
entities are hard to completely avoid since some clades are
just less diverse than others. On the opposite end, if made
possible by the other criteria above, we also try to avoid
overly diverse entities in which we run out of infrageneric
taxonomic ranks, i.e., subgenera and sections, to classify the
distinct monophyletic groups identified within the genus.
When delimiting genera, we aim to find a balance between
the number of genera and the amount of diversity they
include.
The trend in fungal taxonomy, after the introduction of
molecular tools, has in many cases been towards smaller and
natural genera (e.g., Buyck etal. 2008, Sánchez-García etal.
2014, Matheny etal. 2020). The work is still ongoing and
it would be important that all genera would go through the
same re-evaluation so that, in the end, we would be applying
a similar set of criteria for recognizing genera across higher
fungal taxa. This process has and will, without a doubt,
lead to nomenclatural changes, but the end result should
be improvement—a natural, more meaningful, and stable
classification that will provide a good framework for under-
standing and classifying fungal diversity in an evolutionary
context.
Generic delimitation withinCortinariaceae
Current delimitation ofthegenus Cortinarius
andassociated problems
The size of genus Cortinarius with thousands of species and
tremendous morphological variation among them have con-
tributed to a poor understanding of their true diversity and
evolutionary relationships. Even at local scales, Cortinarius
has often been too diverse to manage accurately as a whole
and taxonomists have tended to specialize on certain groups.
For example, in Funga Nordica (Niskanen etal. 2008) Cor-
tinarius is the only genus in which different authors have
written different subkeys of the genus.
The idea of splitting the genus Cortinarius into several
genera is not a new one. Based on morphological data,
different genera—i.e., Dermocybe, Phlegmacium, and
Rozites—have been recognized in the past (e.g., Moser
1960; Moser and Horak 1975). The main issue with all pre-
vious classifications, however, has been that they were either
unnatural or keeping them would have led to the splitting of
the genus into far too many, upractical entities. Alternative
solutions on how to divide the genus into natural units have
not been possible until now, because the phylogenetic stud-
ies done so far have not been able to resolve the deeper nodes
of the phylogeny, beyond the section level (e.g., Peintner
etal 2004; Garnica etal. 2005; Soop etal. 2019).
If we look at how the current situation lines with other
families in the suborder Agaricineae, the ectomycorrhizal
Fungal Diversity
1 3
family Inocybaceae provides a good point of reference for a
comparison. It is a species-rich family, around 2600 species
are recognized based on ITS sequence data using an SH
threshold of 1.5% in UNITE (2021). Before its most recent
molecular revisions (Matheny and Bougher 2006; Alvarado
etal. 2010; Matheny etal. 2020), it used to be a monotypic
family including one genus, Inocybe. Now, the family is
delimited into seven genera based on morpho-genetic data.
If the current system to delimit genus Cortinarius is nei-
ther practical nor does it align well with comparable gen-
era in the suborder, the reasonable conclusion would be to
split the genus. It is hard to see a reason, other than keeping
nomenclatoric stability, to maintain Cortinarius as a single
genus.
New classification proposed andjustification forthenew
generic delimitations
Here we propose the classification of family Cortinari-
aceae into ten genera—Aureonarius, Austrocortinarius,
Calonarius, Cortinarius, Cystinarius, Hygronarius, Mysti-
narius, Phlegmacium, Thaxterogaster, and Volvanarius
based on the phylogenomic analysis of 75 single-copy
nuclear orthologs from 19 species, complemented with a
wider 5-locus analysis of 245 recognized species. There are
names already in existance for the three largest genera, as
well as three to eleven generic level synonyms. Where sev-
eral names of a genus are possible, the most ancient syno-
nym must be chosen (Art. 11.4, 52.1), which explains why a
name like Thaxterogaster now applies to a large taxon with
only some gastroid members. Seven genera are described as
new to science. The position of Stephanopus within subor-
der Agaricineae, for which no sequence data exist, remains
to be studied. The genera have been delimited (i) to be the
largest monophyletic units with statistical support, (ii) to be
supported by morphological traits, (iii) to facilitate classifi-
cation of Cortinariaceae diversity into infrageneric ranks,
(iv) to avoid oversplitting, and (v) to be in line with other
genera of gilled macrofungi.
To strike a balance between the number of genera and the
amount of morphological and species diversity they include,
a few exceptions to the principles above have been made. For
example, we propose that the small telamonioid subgenus
Carbonella should be included in an otherwise rather uni-
form genus Phlegmacium, because treating Carbonella as a
separate genus would have required the division of the genus
Phlegmacium into four separate genera. Also, the genus Cor-
tinarius s. str. has been kept as one unit, rather than further
split it into several smaller genera, even though morphologi-
cally it still is quite a variable genus (and six out of eleven
entities currently recognized as subgenera already have a
generic level name), because many relationships still remain
unresolved and many species are not currently included in
any well-supported group.
Based just on monophyly, the phylogeny would have
allowed us to propose also other solutions for the generic
classification within family Cortinariaceae. We could have
delimited just two genera, Thaxterogaster and Cortinar-
ius, but that would not have led to any major taxonomic
improvements that accurately capture the morphological
variation contained within a single genus. Also, having five
genera—Thaxterogaster, Hygronarius, Austrocortinarius,
Cortinarius, and Phlegmacium—could have been an option.
However, in this scenario, the species-rich genera Phlegma-
cium and Calochroi, as well as the smaller Aureonarius, Cys-
tinarius, and Mystinarius (all entities with morphological
characteristics that make them easily distinguishable from
one another) would have been grouped together.
All genera recognized have a combination of morpho-
logical traits that distinguish them from their closely related
taxa. The smaller genera are more uniform morphologically
and thus more easily defined and recognized. Additionally,
one of the four species-rich genera, Calonarius, is also mor-
phologically very uniform. However, the three largest genera
contain clades that differ morphologically from the others
within that genus. This broadens the infrageneric variation
and affects the diagnostic value of the generic descriptions,
although the vast majority of the species in those genera can
still be distinguished based on morphology from members
of other genera.
An estimation of the species diversity in the ten puta-
tive genera of Cortinariaceae is given in Table3. Genus
Cortinarius s. str. still remains the largest genus in the fam-
ily, with 2000 species estimated worldwide, classified into
eleven subgenera and more than a hundred sections. In the
Northern Hemisphere, the next most species-rich genera are
Phlegmacium (four subgenera, 15 sections) and Calonarius
(three subgenera), while in the Southern Hemisphere (where
Calonarius does not occur and where the diversity of Phleg-
macium is low) the second largest genus is Thaxterogaster
(six subgenera, 18 sections). All these three genera are esti-
mated to have more than or ~ 200 species. The remaining six
genera are smaller, containing 25 species.
Thus, how does the suggested delimitation of genera
compare to other groups of gilled fungi? When comparing
our proposal to the new Inocybaceae classification the new
proposal and its justifcation are very similar. In both cases,
after careful examination of global morpho-genetic data, the
previously monotypic family has been divided into several
genera to better recognize and communicate the amount of
morphological, ecological, and biological diversity observed
within each of them. We propose four larger and six smaller
genera, whereas the current framework for Inocybaceae
includes four larger (> 50 species) and three smaller gen-
era of which one is monotypic. In both cases the genus on
Fungal Diversity
1 3
which the family name is based remains the largest one
with hundreds of species. The refined classification will be
more practical to use for comparative studies and is more
appropriate for conservation studies and for identification
of diversity hotspots than the previous, more inclusive one
(Matheny etal. 2020).
When comparing the sizes of the four most species-rich
genera—Cortinarius, Phlegmacium, Thaxterogaster, and
Calonarius—to other genera of gilled fungi, the new clas-
sification still retains the size signature of the old classifica-
tion, only now with more segregated units that are better
refined. Because comparison at a global scale is challenging
since, for many genera, data are still lacking and/or can be
strongly biased, and because local keys or checklists from
the most intensively studied areas of the world can provide
a better basis for a more accurate comparison, we here use
species numbers from Funga Nordica (Niskanen etal. 2008)
to give an idea of the sizes of the new genera. Based on the
proposed classification, Cortinarius would still be at least
the second largest genus (206 species) after Entoloma (232
species) in Northern Europe and, most likely even remain
the largest when all species from the region are recorded.
The size of Calonarius (~ 80 species) would be equal with
Psathyrella, while the size of Phlegmacium (~ 60 species)
with Tricholoma and the size of Thaxterogaster (~ 30 spe-
cies) with Amanita although for the latter the comparison
does not strictly apply, since Thaxterogaster has its centre of
the diversity in the Southern Hemisphere and, globally, the
size of this genus is equal to that of Phlegmacium.
To maintain nomenclatural stability as much as possible,
we have kept the currently accepted sectional framework
that has been created based on molecular data. In addition,
the new generic names have been designed to have the same
ending -narius as in Cortinarius, to keep the species epithets
as they currently are, whenever possible.
In ecological and conservation studies, the go-to oper-
ational taxonomic unit is the species rank. Under current
usage, the next rank used is the genus level, since no infor-
mation on infrageneric ranks are associated with a name
in fungal DNA barcoding databases. Therefore, the new
classification proposed here will benefit ecological and
other research by providing more biologically relevant cat-
egories. For example, the recognition of genus Calonarius,
with many rare representatives that have narrow ecological
preferences, will help highlight its uniqueness. Moreover,
this classification will also advance communication of con-
servation priorities, as many of the species in Calonarius
are included on national red lists across Europe (e.g. Stoltze
and Pihl 1998; SLU Artdatabanken 2020) with C. meinhar-
dii also making it into the global red list (Brandrud 2019).
Having Calonarius as a separate genus will help draw focus
into this group and provide a tool to better recognize it by
ecologists and conservation biologists.
Previous phylogenetic studies inCortinariaceae
Genera recognized in this study have also been recovered
in the two previous multi-gene studies of Cortinariaceae
that included data from the RPB1 region, in addition to
the traditionally used ITS and LSU regions (Garnica etal.
2016; Soop etal 2019). Garnica etal. (2016) recognized the
following clades: Phlegmacioid clade 1 (= Phlegmacium),
Phlegmacioid clade 2 (= Thaxterogaster), Phlegmacioid
clade 3 (= Calonarius), Renidentes (= Hygronarius), and
Coleopodes (= Volvanarius). Also, Aureonarius, Corti-
narius, Cystinarius, and Mystinarius were included in their
analysis and formed their own respective clades, although
they were not named in the tree. The only genus not repre-
sented in this earlier study is Austrocortinarius, but it was
included in the phylogeny of Soop etal. (2019), which also
recovered the same clades inferred by Garnica etal. (2016).
Although many lineages in Garnica etal. (2016) received
good support, they were unsupported in the analysis of Soop
etal. (2019) and, therefore, were not recognized as formally
named taxa.
The phylogenies based on ITS and LSU alone can recover
the proposed genera to some extent, but these two gene
regions do not suffice to resolve all of the infrafamilial rela-
tionships correctly. Particularly, they fail in recovering the
monophyly of genera Cortinarius s. str. and Aureonarius
(Garnica etal. 2005; Stensrud etal. 2014). Rather, the ITS
and LSU regions are most suitable for shallow level classi-
fication, i.e., species and sections. To get a better idea on the
higher level classification of Cortinariaceae, at least RPB1
would be needed, in addition to ITS and LSU. For optimal
resolution, genome-wide data should be used.
How does thenew proposed classification differ
fromtheexisting one?
Moving from one to ten genera is a big change, but our pro-
posal is not a large leap in circumscribing the known diver-
sity. First, most of the diversity belongs to four large gen-
era, Cortinarius (species-rich in the Northern and Southern
Hemispheres), Phlegmacium (species-rich in the Northern
Hemisphere, but far fewer species in the Southern Hemi-
sphere), Thaxterogaster (species-rich in the Southern Hemi-
sphere, but far fewer species in the Northern Hemisphere)
and Calonarius (restricted to the Northern Hemisphere).
Secondly, most of the species-level diversity remains within
genus Cortinarius s. str., which includes most of the spe-
cies with telamonioid, cortinarioid (including dermocyboid
and leprocyboid) or myxacioid habits, and all species with
Fungal Diversity
1 3
rozitoid or cuphocyboid habits (Table3). Third, except for
Aureonarius and Hygronarius, the small genera Austrocorti-
narius, Cystinarius, Mystinarius, and Volvanarius have long
been enigmatic and difficult to confidently place within the
previous classifications based on morphological traits only.
The main difference to the previous classification is the
transfer of most phlegmacioid species to three separate
genera: Phlegmacium, Calonarius, and Thaxterogaster.
Of these, Calonarius is the easiest one to distinguish based
solely on morphology. It has been recognized as a separate
lineage from very early phylogenetic studies onwards (Pei-
ntner etal. 2004; Garnica etal. 2005) and several molecular
studies have focused on it (e.g., Frøslev etal. 2007; Gar-
nica etal. 2009, 2011). In the Northern Hemisphere, genus
Phlegmacium contains most of the species traditionally
included in C. subgenus Phlegmacium, except for sections
Multiformes, Scauri/Purpurascentes, Riederorum, Lus-
trati, Pinophili, Turmales, and C. variegatus which instead
belong to genus Thaxterogaster. In the Southern Hemisphere
Nothofagaceae forests, however, the default genus for phleg-
macioid species is Thaxterogaster. So far, no Calonarius
species have been found in the Southern Hemisphere and
far fewer species of Phlegmacium occur in the Southern
than in the Northern Hemisphere. The peculiar species of
Southern Hemisphere C. sect. Thaumasti, which look like
miniature, volvate Phlegmacium, are treated in their own
genus, Volvanarius; whereas the two distinctive, big, white
species that occur in Australia and New Zealand are placed
in genus Austrocortinarius. In addition, sections Crassi and
Rubicunduli, traditionally classified with either phlegma-
cioid or cortinarioid species, are treated in their own genus,
Cystinarius. Consequently, few stipitocarpic lineages with
a phlegmacioid appearance—sections Subtorti, Infracti,
Dulciolentes, Cuphocybe, and Vinaceolamellati—are kept
in genus Cortinarius.
The myxacioid species mostly remain in genus Cor-
tinarius and only sections Vibratiles and Austrocyanites
are moved into Thaxterogaster, while C. lustrabilis and C.
badiohepaticus are placed in the genus Mystinarius. Also,
cortinarioid species mainly remain in genus Cortinarius and
the only changes are the placement of sections Callistei and
Limoni in their own genus Aureonarius, and the placement
of section Rubicunduli in genus Cystinarius. In addition,
most telamonioid species belong to genus Cortinarius and
only some tens of species are placed in other lineages: genus
Hygronarius and P. subgenus Carbonella. All rozitoid and
cuphocypoid species belong to genus Cortinarius.
Sequestrate species have already been shown to belong
to different lineages of Cortinariaceae (Peintner etal.
2001; Nouhra etal. 2021). Although most of them belong
to either Cortinarius or Thaxterogaster, they are found in
all four of the largest Cortinariaceae genera (Cortinarius,
Phlegmacium, Calonarius, and Thaxterogaster), as well as
in the small genus Volvanarius.
Infrageneric classification
Our main goal for this study, was to produce a revised
generic framework for family Cortinariaceae based on a
robust phylogeny derived from genomic data. Futhermore,
a base for subgeneric classification is also proposed by rec-
ognizing clades with strong to full support, while indicat-
ing the possible limits of the already existing subgenera. In
some genera, i.e., Aureonarius and Phlegmacium, for which
single-copy gene data from a wide range of species already
exist, all species were placed in moderately to fully sup-
ported groups (Fig.2). Elsewhere, e.g., the two species-rich
genera Cortinarius and Thaxterogaster, further multi-gene
studies will be needed to clarify the infrageneric relation-
ships and, at present, only the morphologically and geneti-
cally most distinct groups are here recognized. A total of 30
subgenera are recognized of which 10 are here described as
new to science.
For the most species-rich genus, Cortinarius, 11 subgen-
era and 130 sections are currently recognized, although most
of these sections(80) belong to the most species-rich sub-
genus of Cortinariaceae, C. subgen. Telamonia. Morpho-
logical variation in the genus is broad but correlates rather
well with the phylogeny. The vast majority of cortinarioid
(dermocyboid, leprocyboid) and telamonioid species are
placed in the strongly supported (BS 97%) crown group of
Cortinarius. The group includes the cortinarioid subgenera
Dermocybe, Leprocybe, and Orellani, and the telamonioid
subgenera Iodolentes, Illumini, and Telamonia, plus several
sections and species whose relationships were not well-
resolved (Fig.2). The small phlegmacioid subgenus Infracti
and the New Zealand endemic C. pholiotellus form a fully
supported (BS 100%) sister group to the crown group. Lead-
ing to this crown clade, we find a grade comprised mainly
myxacioid, rozitoid, cuphocypoid, and sequestrate species,
together with C. sect. Anomali, which was previously placed
in C. subgen. Telamonia. Tentative limits of the previously
described subgenera are marked in the tree (Fig.2). Sub-
genus Cortinarius, which includes the type species of the
genus C. violaceus, is tentatively placed in an unsupported
clade sister to all other Cortinarius species. It is a mor-
phologically unique group characterized by dark purple to
blackish-purple species with dry velvety-squamulose pileus
and cheilocystidia (Harrower etal. 2015a, b).
In the genus Phlegmacium, four subgenera and 22 sec-
tions are recognized. The subgenera are all moderately to
fully supported in the phylogenetic analysis and are also
supported by morphological traits. The two large groups,
P. subgen. Phlegmacium and P. subgen. Bulbopodium, are
Fungal Diversity
1 3
characterized by different basidiomata development types:
the basidiomata of Phlegmacium are stipitocarpic and those
of Bulbopodium pileocarpic. The small subgenus Cyanicium
includes species with bluish to violet (brownish)grey basid-
iomata and reddening context. They are phlegmacioid in
appearance but have previously also been placed in Telamo-
nia and Sericeocybe (Soop etal. 2019). Also, Carbonella is
placed with moderate support in genus Phlegmacium in our
analysis. It differs from all other lineages of Phlegmacium
by having telamonioid basidiomata and is thus recognized
here as its own subgenus.
Thaxterogaster includes phlegmacioid, myxacioid and a
few telamonioid species. At present, five new subgenera,
in addition to the autonym, are proposed to describe the
morphologically most distinct units and to serve as anchors
for further studies. Twenty-eight sections are recognized.
More sampling of species from the Southern Hemisphere
is needed to better understand the infrageneric relationships
and evolution of the genus.
In Calonarius, three subgenera and 11 sections are recog-
nized. In the smaller genera Aureonarius, Cystinarius, and
Hygronarius, two subgenera are recognized for each genus,
and this division is also supported by morphological traits.
In Austrocortinarius, Mystinarius, and Volvanarius, which
only contain a few species, no further infrageneric classifica-
tion is proposed.
Fungariomics
Two approaches were used to create genomic data for our
phylogenomic study: shallow WGS and targeted capture
sequencing. For the shallow WGS, the goal was to produce
sufficient sequence data representative of the whole genome
to assemble into contiguous sequences and then fish our tar-
gets from the resulting assemblies. For the targeted capture
sequencing, the baits are first designed for the chosen targets,
then genomic libraries of chosen samples are enriched, via in
solution hybridization with our baits, so that only the targets
will be sequenced. We wanted to compare the performance
of these two methods in fungi, for which the genome size
is relatively small and thus producing low cost WGS data
is possible.
Results obtained from targeted capture sequencing were
remarkably better than those achieved via shallow WGS.
With targeted capture sequencing, over 85% of the 75 targets
used for the phylogenomics analysis were recovered, for nine
out of 11 specimens. Whereas with WGS, the same percent-
age was only achieved for three out of nine specimens. The
same specimen of Cystinarius crassus was processed with
both methods and 33% of the targets were recovered with
WGS, compared to 99% with targeted capture sequencing.
The advantage of the targeted capture sequencing approach
is that only the targeted regions of the systematic group stud-
ied are sequenced whereas in WGS the whole genome is
sequenced, including possible contaminants present in the
basidiomata collected from the wild or gained during the
preservation process (Dentinger etal. 2016). Thus, prior
knowledge of the genome size of the studied species might
help optimize the number of specimens to be pooled for
WGS; although, even then, some of the capacity might be
lost to sequencing accompanying organisms. Somewhat
better results could potentially have also been achieved by
improved assembly quality through assembly refinement and
it could also be possible that differences in the preservation
state and molecular processing of sampled Cortinariaceae
species might as well have had some impact in the results
(Brewer etal. 2019; Forrest etal. 2019). However, a thor-
ough comparison and evaluation of the effects of these dif-
ferent factors to the success rate is beyond the scope of this
study and overall, they would not entirely explain the differ-
ences observed in performance.
The age of the fungarium specimens sampled ranged
from one to 21year for the WGS and from four to 13year
for the targeted capture sequencing. Targeted capture
sequencing studies in plants have used herbarium speci-
mens ~ 50–200years old (Brewer etal. 2019; Shee etal.
2020), and even thousands of years old aDNA in archae-
ogenomic studies (Kistler etal. 2020). Therefore, targeted
capture sequencing is also a very promising approach for
fungariomics and provides a way to unlock the full poten-
tial of specimens stored in fungaria worldwide for phyloge-
netic analysis. In our study we generated genomic libraries
with medium sized inserts, which we chose to sequence in
a MiSeq using the Nano chemistry (250 × 250bp), due to
the small number of specimens processed in total. However,
especially for the older specimens, where the DNA is likely
to be more degraded, the Illumina platforms (i.e., HiSeq,
NextSeq, NovaSeq), taking shorter sized fragments as tem-
plate, would be more suitable.
Our results join the existing evidence (Dodsworth etal.
2019) showing that targeted capture sequencing provides a
cost-efficient approach (Hale etal. 2020) to produce data for
phylogenomic analyses for species-rich groups, like Corti-
nariaceae, in which one can use the same set of baits for
a large range of species (e.g., see Liu etal. 2019, mosses;
Johnson etal. 2019, angiosperms; or Widhelm etal. 2021,
peltigeralean lichens). The initial bait design and capture
reactions add costs compared to the WGS and therefore
the latter approach can be more appropriate for smaller
taxonomic groups, unless an enrichment panel is already
available. It is difficult to provide a precise threshold on
the number of species for which targeted capture sequenc-
ing becomes cheaper than WGS in fungi, since costs of
genomic data are in constant flux and costs also depend
Fungal Diversity
1 3
on the genome size and intended sequencing coverage, as
well as the number of baits to be included in the enrichment
panel. Nonetheless, to give some idea of the difference in
volume between these two methods, for Cortinariaceae we
estimated that about ~ 200 specimens could be pooled into a
single Illumina MiSeq 2 × 300bp paired end run, when using
targeted capture sequencing, compared to five to six speci-
mens in total for the WGS approach. On the other hand, in
some cases it might be justified to choose the WGS approach
for other reasons, e.g., its potential added value of providing
data on other genomic features or other loci, not included
for a given enrichment panel, for analyses at different taxo-
nomic levels.
Conclusions
This study is the first family revision in Agaricales based on
genomic data and hopefully many others will soon follow.
We have come a long way from the time of Fries when all
gilled fungi were in one genus, Agaricus (Fries 1821). Since
then, mycologists have, in most cases, created smaller and
smaller genera due to the increased understanding of the
diversity and enhanced ability to collect data of the organ-
isms for taxonomic studies. The same phenomenon has
happened in plants and animals. The genus Cortinarius has
been an especially difficult group for taxonomists, because it
includes an enormous amount of morphological and species
diversity. While there have been previous efforts to divide
the genus into more manageable, practical, and natural units,
none have achieved a natural classification for the whole
group. Our proposed classification for Cortinariaceae is
more equilevant to contemporary concepts in other genera
of gilled fungi and we hope that our framework will be more
user-friendly, facilitating the identification, conservation and
ecological studies on these fascinating organisms.
Additional new combinations ofthespecies
andsections belonging tothedierent
genera offamily Cortinariaceae
Aureonarius
Aureonarius armiae (Soop) Niskanen & Liimat., comb. nov.
IF553605
Basionym: Cortinarius armiae Soop, Bresadoliana 1(2): 19.
2013.
Aureonarius aurantiobrunneus (Ammirati, Halling & Gar-
nica) Niskanen & Liimat., comb. nov.
IF553606
Basionym: Cortinarius aurantiobrunneus Ammirati, Halling
& Garnica, in Ammirati, Garnica, Halling, Mata, Mueller &
Carranza, Can. J. Bot. 85(9): 801. 2007.
Aureonarius austrolimonius (M.M. Moser & E. Horak) Lii-
mat. & Niskanen, comb. nov.
IF553607
Basionym: Cortinarius austrolimonius M.M. Moser & E.
Horak, Beih. Nova Hedwigia 52: 454. 1975.
Aureonarius callisteus (Fr.) Niskanen & Liimat., comb. nov.
IF553608
Basionym: Agaricus callisteus Fr., Observ. mycol. (Havniae)
2: 51. 1818.
Aureonarius caryotis (Soop) Niskanen & Liimat., comb.
nov.
IF553621
Basionym: Cortinarius caryotis Soop, Bull. Soc. mycol. Fr.
117(2): 97. 2001.
Aureonarius caryotoides (Soop) Niskanen & Liimat., comb.
nov.
IF553622
Basionym: Cortinarius caryotoides Soop, in Soop, Wallace
& Dima, N.Z. Jl Bot. 56(2): 166. 2018.
Aureonarius controversus (Gasparini) Niskanen & Liimat.,
comb. nov.
IF553623
Basionym: Cortinarius controversus Gasparini, in Gasparini
& Soop, Australas. Mycol. 27(3): 190. 2008.
Fungal Diversity
1 3
Aureonarius eucollybianus (Soop) Niskanen & Liimat.,
comb. nov.
IF553624
Basionym: Cortinarius eucollybianus Soop, in Soop, Wal-
lace & Dima, N.Z. Jl Bot. 56(2): 171. 2018.
Aureonarius infucatus (Fr.) Niskanen & Liimat., comb. nov.
IF553625
Basionym: Cortinarius infucatus Fr., Öfvers. K. Svensk.
Vetensk.-Akad. Förhandl. 18(1): 26. 1861.
Aureonarius limonius (Fr.) Niskanen & Liimat., comb. nov.
IF553626
Basionym: Agaricus limonius Fr., Observ. mycol. (Havniae)
2: 56. 1818.
Aureonarius neocallisteus (Kranab., Ammirati, Liimat. &
Niskanen) Niskanen & Liimat., comb. nov.
IF553640
Basionym: Cortinarius neocallisteus Kranab., Ammirati,
Liimat. & Niskanen, in Niskanen, Liimatainen, Kytövuori,
Lindström, Dentinger & Ammirati, Mycologia 108(5): 1024.
2016.
Aureonarius rubrimarginatus (Soop) Niskanen & Liimat.,
comb. nov.
IF553641
Basionym: Cortinarius rubrimarginatus Soop, in Soop,
Wallace & Dima, N.Z. Jl Bot. 56(2): 169. 2018.
Aureonarius rubrocastaneus (Soop) Niskanen & Liimat.,
comb. nov.
IF553643
Basionym: Gymnopilus rubrocastaneus Soop, Bull. Soc.
mycol. Fr. 117(2): 128. 2001.
Aureonarius rubrodactylus (Soop) Niskanen & Liimat.,
comb. nov.
IF553644
Basionym: Cortinarius rubrodactylus Soop, Australas.
Mycol. 31: 3. 2013.
Aureonarius tofaceus (Fr.) Niskanen & Liimat., comb. nov.
IF553645
Basionym: Cortinarius tofaceus Fr., Epicr. syst. mycol.
(Upsaliae): 281. 1838. [1836–1838].
Aureonarius viscilaetus (Soop) Niskanen & Liimat., comb.
nov.
IF553777
Basionym: Cortinarius viscilaetus Soop, Bull. Soc. mycol.
Fr. 117(2): 114. 2001.
Austrocortinarius
Austrocortinarius australiensis (Cleland & Cheel) Liimat.
& Niskanen, comb. nov.
IF553778
Basionym: Rozites australiensis Cleland & Cheel, Trans. &
Proc. Roy. Soc. S. Australia 42: 90. 1918.
Calonarius
Calonarius sect. Atrovirentes (Bidaud, Moënne-Locc. &
Reumaux) Niskanen & Liimat., comb. & stat. nov.
IF553779
Basionym: Cortinarius ser. Atrovirens Bidaud, Moënne-
Locc. & Reumaux, in Bidaud, Carteret, Eyssartier, Moënne-
Loccoz & Reumaux, Atlas des Cortinaires (Meyzieu) 14:
958. 2004.
Calonarius sect. Aureopulverulenti (Brandrud & Melot)
Niskanen & Liimat., comb. & stat. nov.
IF553780
Basionym: Cortinarius subsect. Aureopulverulenti Brandrud
& Melot, Nordic Jl Bot. 10(5): 535. 1990.
Calonarius sect. Dibaphi (Brandrud & Melot) Niskanen &
Liimat., comb. & stat. nov.
IF553781
Fungal Diversity
1 3
Basionym: Cortinarius subsect. Dibaphi Brandrud & Melot,
Nordic Jl Bot. 10(5): 537. 1990.
Calonarius sect. Flavovirentes (Bidaud, Moënne-Locc. &
Reumaux) Niskanen & Liimat., comb. & stat. nov.
IF553782
Basionym: Cortinarius ser. Flavovirens Bidaud, Moënne-
Locc. & Reumaux, in Bidaud, Carteret, Eyssartier, Moënne-
Loccoz & Reumaux, Atlas des Cortinaires (Meyzieu) 14:
958. 2004.
Calonarius sect. Osmophori (Bidaud & Reumaux) Niskanen
& Liimat., comb. & stat. nov.
IF553783
Basionym: Cortinarius stirps Osmophorus Bidaud & Reu-
maux, in Bidaud, Moënne-Loccoz, Reumaux, Carteret &
Eyssartier, Atlas des Cortinaires (Meyzieu) 16: 1097. 2006.
Calonarius sect. Platypodes (Bidaud, Moënne-Locc. & Reu-
maux) Niskanen & Liimat., comb. & stat. nov.
IF553784
Basionym: Cortinarius ser. Platypodes Bidaud, Moënne-
Locc. & Reumaux, in Bidaud, Moënne-Loccoz, Reumaux,
Carteret & Eyssartier, Atlas des Cortinaires (Meyzieu) 11:
611. 2001.
Calonarius sect. Rufoolivacei (Brandrud & Melot) Niskanen
& Liimat., comb. & stat. nov.
IF553785
Basionym: Cortinarius subsect. Rufoolivacei Brandrud &
Melot, Nordic Jl Bot. 10(5): 538. 1990.
Calonarius sect. Sodagniti (Bidaud, Moënne-Locc. & Reu-
maux) Niskanen & Liimat., comb. & stat. nov.
IF553786
Basionym: Cortinarius ser. Sodagniti Bidaud, Moënne-
Locc. & Reumaux, Docums Mycol. 24(no. 95): 43. 1994.
Calonarius sect. Splendentes (Kühner & Romagn. ex Bran-
drud & Melot) Niskanen & Liimat., comb. & stat. nov.
IF553787
Basionym: Cortinarius subsect. Splendentes Kühner &
Romagn. ex Brandrud & Melot, Nordic J. Bot. 10(5): 538.
1990.
Calonarius adonis (Bojantchev & Ammirati) Niskanen &
Liimat., comb. nov.
IF553788
Basionym: Cortinarius adonis Bojantchev & Ammirati, in
Bojantchev, Index Fungorum 247: 1. 2015.
Calonarius albertii (Dima, Frøslev & T.S. Jeppesen) Nis-
kanen & Liimat., comb. nov.
IF553789
Basionym: Cortinarius albertii Dima, Frøslev & T.S.
Jeppesen, in Frøslev, Jeppesen & Læssøe, Mycol. Res.
110(9): 1050. 2006.
Calonarius albidolilacinus (Ammirati, Bojantchev, Beug,
Liimat., Niskanen & Garnica) Niskanen & Liimat., comb.
nov.
IF553790
Basionym: Cortinarius albidolilacinus Ammirati,
Bojantchev, Beug, Liimat., Niskanen & Garnica, in Lii-
matainen, Index Fungorum 241: 1. 2015.
Calonarius alcalinophilus (Rob. Henry) Niskanen & Lii-
mat., comb. nov.
IF553791
Basionym: Cortinarius alcalinophilus Rob. Henry, Bull.
trimest. Soc. mycol. Fr. 67(3): 301. 1952. (1951).
Calonarius alnobetulae (Kühner) Niskanen & Liimat.,
comb. nov.
IF553792
Basionym: Cortinarius alnobetulae Kühner, Docums Mycol.
20(77): 92. 1989.
Calonarius amabilis (Bojantchev, Ammirati & Pastorino)
Niskanen & Liimat., comb. nov.
IF553793
Fungal Diversity
1 3
Basionym: Cortinarius amabilis Bojantchev, Ammirati &
Pastorino, in Bojantchev, Index Fungorum 247: 1. 2015.
Calonarius amnicola (A.H. Sm.) Niskanen & Liimat., comb.
nov.
IF553794
Basionym: Cortinarius amnicola A.H. Sm., Bull. Torrey
bot. Club 69(1): 48. 1942.
Calonarius anaunianus (Fellin & R.J. Ferrari) Niskanen &
Liimat., comb. nov.
IF553795
Basionym: Cortinarius anaunianus Fellin & R.J. Ferrari,
in Fellin, Ercole, Ferrari & Vizzini, Phytotaxa 520(3): 230.
2021.
Calonarius anetholens (Ammirati, Garnica, Bojantchev,
Beug, Liimat. & Niskanen) Niskanen & Liimat., comb. nov.
IF553800
Basionym: Cortinarius anetholens Ammirati, Garnica,
Bojantchev, Beug, Liimat. & Niskanen, in Liimatainen,
Index Fungorum 241: 1. 2015.
Calonarius arcuatorum (Rob. Henry) Niskanen & Liimat.,
comb. nov.
IF553801
Basionym: Cortinarius arcuatorum Rob. Henry, Bull. tri-
mest. Soc. mycol. Fr. 55(1): 80. 1939.
Calonarius arenicola (A.H. Sm.) Niskanen & Liimat., comb.
nov.
IF553802
Basionym: Cortinarius arenicola A.H. Sm., Bull. Torrey
bot. Club 69(1): 49. 1942.
Calonarius atrovirens (Kalchbr.) Niskanen & Liimat., comb.
nov.
IF553803
Basionym: Cortinarius atrovirens Kalchbr., Icon. Sel.
Hymenomyc. Hung. (Budapest) 2: tab. 19. 1874.
Calonarius aureocalceolatus (M.M. Moser & Peintner) Nis-
kanen & Liimat., comb. nov.
IF553804
Basionym: Cortinarius aureocalceolatus M.M. Moser &
Peintner, Journal des JEC, Journées Européenes du Corti-
naire 5(no. 4): 30. 2002.
Calonarius aureofulvus (M.M. Moser) Niskanen & Liimat.,
comb. nov.
IF553851
Basionym: Cortinarius aureofulvus M.M. Moser, Sydowia
6(1–4): 154. 1952.
Calonarius aureopulverulentus (M.M. Moser) Niskanen &
Liimat., comb. nov.
IF553852
Basionym: Cortinarius aureopulverulentus M.M. Moser,
Sydowia 6(1–4): 152. 1952.
Calonarius aurora (M.M. Moser & Ammirati) Niskanen &
Liimat., comb. nov.
IF553853
Basionym: Cortinarius aurora M.M. Moser & Ammirati, in
Moser, McKnight & Ammirati, Mycotaxon 55: 305. 1995.
Calonarius barbaricus (Brandrud) Niskanen & Liimat.,
comb. & stat. nov.
IF553854
Basionym: Cortinarius callochrous var. barbaricus Bran-
drud, Cortinarius, Flora Photographica (Matfors) 3: 27.
1994.
Calonarius barbarorum (Bidaud, Moënne-Locc. & Reu-
maux) Niskanen & Liimat., comb. nov.
IF553855
Basionym: Cortinarius barbarorum Bidaud, Moënne-Locc.
& Reumaux, in Bidaud, Moënne-Loccoz, Reumaux, Carteret
& Eyssartier, Atlas des Cortinaires (Meyzieu) 11: 612. 2001.
Calonarius bigelowii (Thiers & A.H. Sm.) Niskanen & Lii-
mat., comb. nov.
Fungal Diversity
1 3
IF554100
Basionym: Cortinarius bigelowii Thiers & A.H. Sm., Myco-
logia 61: 531. 1969.
Calonarius cacodes (M.M. Moser & Ammirati) Niskanen
& Liimat., comb. nov.
IF554101
Basionym: Cortinarius cacodes M.M. Moser & Ammirati,
Mycotaxon 74(1): 6. 2000.
Calonarius caesiocinctus (Kühner) Niskanen & Liimat.,
comb. nov.
IF554102
Basionym: Cortinarius caesiocinctus Kühner, Docums
Mycol. 20(77): 92. 1989.
Calonarius caesiolatens (Rob. Henry ex Bidaud & Reu-
maux) Niskanen & Liimat., comb. nov.
IF554103
Basionym: Cortinarius caesiolatens Rob. Henry ex Bidaud
& Reumaux, in Bidaud, Carteret, Eyssartier, Moënne-Loc-
coz & Reumaux, Atlas des Cortinaires (Meyzieu) 14: 960.
2004.
Calonarius callochrous (Pers.) Niskanen & Liimat., comb.
nov.
IF554104
Basionym: Agaricus callochrous Pers., Syn. meth. fung.
(Göttingen) 2: 282. 1801.
Calonarius calojanthinus (M.M. Moser & Ammirati) Nis-
kanen & Liimat., comb. nov.
IF559246
Basionym: Cortinarius calojanthinus M.M. Moser &
Ammirati, Mycotaxon 72: 291. 1999.
Calonarius caroviolaceus (P.D. Orton) Niskanen & Liimat.,
comb. nov.
IF554105
Basionym: Cortinarius caroviolaceus P.D. Orton, Trans. Br.
mycol. Soc. 43(2): 208. 1960.
Calonarius catharinae (Consiglio) Niskanen & Liimat.,
comb. nov.
IF554106
Basionym: Cortinarius catharinae Consiglio, Riv. Micol.
39(3): 199. 1997. [1996].
Calonarius cedretorum (Maire) Niskanen & Liimat., comb.
nov.
IF554107
Basionym: Cortinarius cedretorum Maire, Bull. Soc. mycol.
Fr. 30(2): 210. 1914.
Calonarius chailluzii (Frøslev & T.S. Jeppesen) Niskanen
& Liimat., comb. nov.
IF554108
Basionym: Cortinarius chailluzii Frøslev & T.S. Jeppesen,
in Frøslev, Jeppesen & Læssøe, Mycol. Res. 110(9): 1051.
2006.
Calonarius cisticola (Frøslev & T.S. Jeppesen) Niskanen &
Liimat., comb. nov.
IF554109
Basionym: Cortinarius cisticola Frøslev & T.S. Jeppesen,
in Frøslev, Jeppesen & Læssøe, Mycol. Res. 110(9): 1051.
2006.
Calonarius citrinipedes (A.H. Sm.) Niskanen & Liimat.,
comb. nov.
IF554400
Basionym: Cortinarius citrinipedes A.H. Sm., Bull. Torrey
bot. Club 69(1): 54. 1942.
Calonarius citrinus (J.E. Lange ex P.D. Orton) Niskanen &
Liimat., comb. nov.
IF554401
Basionym: Cortinarius citrinus J.E. Lange ex P.D. Orton,
Bull. trimest. Soc. mycol. Fr. 55(2): 176. 1960.
Fungal Diversity
1 3
Calonarius claroflavus (Rob. Henry) Niskanen & Liimat.,
comb. nov.
IF554402
Basionym: Cortinarius claroflavus Rob. Henry, Bull. tri-
mest. Soc. mycol. Fr. 67(3): 297. 1952. (1951).
Calonarius cobaltinus (Kytöv., Liimat. & Niskanen) Nis-
kanen & Liimat., comb. nov.
IF554403
Basionym: Cortinarius cobaltinus Kytöv., Liimat. & Nis-
kanen, in Liimatainen, Index Fungorum 22: 1. 2013.
Calonarius coniferarum (M.M. Moser) Niskanen & Liimat.,
comb. & stat. nov.
IF554404
Basionym: Phlegmacium multiforme var. coniferarum M.M.
Moser, Gatt. Phlegm.: 349. 1960.
Calonarius corrosus (Fr.) Niskanen & Liimat., comb. nov.
IF554405
Basionym: Cortinarius corrosus Fr., Epicr. syst. mycol.
(Upsaliae): 266. 1838. [1836–1838].
Calonarius cupreorufus (Brandrud) Niskanen & Liimat.,
comb. nov.
IF554406
Basionym: Cortinarius cupreorufus Brandrud, in Brandrud,
Lindström, Marklund, Melot & Muskos, Cortinarius, Flora
Photographica (Matfors) 3: 27. 1994.
Calonarius dalecarlicus (Brandrud) Niskanen & Liimat.,
comb. nov.
IF554407
Basionym: Cortinarius dalecarlicus Brandrud, in Brandrud,
Lindström, Marklund, Melot & Muskos, Cortinarius, Flora
Photographica vol. 2 (Sweden): 33. 1992.
Calonarius dibaphus (Fr.) Niskanen & Liimat., comb. nov.
IF554408
Basionym: Cortinarius dibaphus Fr., Epicr. syst. mycol.
(Upsaliae): 266. 1838. [1836–1838].
Calonarius elegantiomontanus (Garnica & Ammirati) Nis-
kanen & Liimat., comb. nov.
IF554409
Basionym: Cortinarius elegantiomontanus Garnica &
Ammirati, in Garnica, Spahn, Oertel, Ammirati & Ober-
winkler, BMC Evol. Biol. 11(213 [reprint]): 13 + Additional
file3: 20. 2011.
Calonarius elegantior (Fr.) Niskanen & Liimat., comb. &
stat. nov.
IF554410
Basionym: Agaricus multiformis β elegantior Fr., Observ.
mycol. (Havniae) 2: 64. 1818.
Calonarius elegantissimus (Rob. Henry) Niskanen & Lii-
mat., comb. nov.
IF554411
Basionym: Cortinarius elegantissimus Rob. Henry, Docums
Mycol. 20(77): 69. 1989.
Calonarius elotoides (M.M. Moser & McKnight) Niskanen
& Liimat., comb. nov.
IF554412
Basionym: Cortinarius elotoides M.M. Moser & McKnight,
in Moser, McKnight & Ammirati, Mycotaxon 55: 311. 1995.
Calonarius elotus (Fr.) Niskanen & Liimat., comb. nov.
IF554413
Basionym: Cortinarius elotus Fr., Epicr. syst. mycol.
(Upsaliae): 264. 1838. (1836–1838).
Calonarius evosmus (Joachim ex Bidaud & Reumaux) Nis-
kanen & Liimat., comb. nov.
IF554414
Basionym: Cortinarius evosmus Joachim ex Bidaud & Reu-
maux, in Bidaud, Moënne-Loccoz, Reumaux, Carteret &
Eyssartier, Atlas des Cortinaires (Meyzieu) 16: 1097. 2006.
Fungal Diversity
1 3
Calonarius flavaurora (M.M. Moser & McKnight) Niskanen
& Liimat., comb. nov.
IF554415
Basionym: Cortinarius flavaurora M.M. Moser & McK-
night, in Moser, McKnight & Ammirati, Mycotaxon 55:
321. 1995.
Calonarius flavoaurantians (Boccardo, Cleric. & Vizzini)
Niskanen & Liimat., comb. nov.
IF554416
Basionym: Cortinarius flavoaurantians Boccardo, Cleric. &
Vizzini, in Vizzini, Clericuzio, Boccardo & Ercole, Myco-
logia 104(6): 1504. 2012.
Calonarius flavobulbus (Ammirati & M.M. Moser) Nis-
kanen & Liimat., comb. nov.
IF554417
Basionym: Cortinarius flavobulbus Ammirati & M.M.
Moser, in Moser & Ammirati, Sydowia 49(1): 34. 1997.
Calonarius flavovirens (Rob. Henry) Niskanen & Liimat.,
comb. nov.
IF554418
Basionym: Cortinarius flavovirens Rob. Henry, Bull. tri-
mest. Soc. mycol. Fr. 55(2): 182. 1939.
Calonarius frondosophilus (Bidaud) Niskanen & Liimat.,
comb. nov.
IF554419
Basionym: Cortinarius frondosophilus Bidaud, in Bidaud,
Moënne-Loccoz, Reumaux, Carteret & Eyssartier, Atlas des
Cortinaires (Meyzieu) 11: 612. 2001.
Calonarius fulvoarcuatorum (Garnica & Ammirati) Nis-
kanen & Liimat., comb. nov.
IF554420
Basionym: Cortinarius fulvoarcuatorum Garnica & Ammi-
rati, in Garnica, Spahn, Oertel, Ammirati & Oberwinkler,
BMC Evol. Biol. 11(213 [reprint]): 13 + Additional file3:
5. 2011.
Calonarius fulvocitrinus (Brandrud) Niskanen & Liimat.,
comb. nov.
IF554421
Basionym: Cortinarius fulvocitrinus Brandrud, in Brandrud,
Lindström, Marklund, Melot & Muskos, Cortinarius, Flora
Photographica (Matfors) 4: 20. 1998.
Calonarius fulvoincarnatus (Joachim ex Bidaud, Moënne-
Locc. & Reumaux) Niskanen & Liimat., comb. nov.
IF554422
Basionym: Cortinarius fulvoincarnatus Joachim ex Bidaud,
Moënne-Locc. & Reumaux, in Bidaud, Moënne-Loccoz,
Reumaux, Carteret & Eyssartier, Atlas des Cortinaires
(Meyzieu) 11: 613. 2001.
Calonarius glaucescens (Jul. Schäff.) Niskanen & Liimat.,
comb. nov.
IF554423
Basionym: Phlegmacium glaucescens Jul. Schäff., Die Gat-
tung Phlegmacium (Schleimköpfe). Die Pilze Mitteleuropas
4: 359. 1960.
Calonarius glaucoelotus (Brandrud, Dima, Krisai, Ballarà
& Peintner) Niskanen & Liimat., comb. nov.
IF554427
Basionym: Cortinarius glaucoelotus Brandrud, Dima, Kri-
sai, Ballarà & Peintner, in Crous et. al., Persoonia 45: 405.
2020.
Calonarius haasii (M.M. Moser) Niskanen & Liimat., comb.
& stat. nov.
IF554428
Basionym: Phlegmacium arquatum var. haasii M.M. Moser,
Gatt. Phlegm.: 353. 1960.
Calonarius hildegardiae (Schmidt-Stohn, Brandrud &
Dima) Niskanen & Liimat., comb. nov.
IF554429
Basionym: Cortinarius hildegardiae Schmidt-Stohn,
Brandrud & Dima, in Brandrud, Schmidt-Stohn & Dima,
Sydowia 71: 119. 2019.
Fungal Diversity
1 3
Calonarius insignibulbus (Bidaud & Moënne-Locc.) Nis-
kanen & Liimat., comb. nov.
IF554430
Basionym: Cortinarius insignibulbus Bidaud & Moënne-
Locc., in Bidaud, Moënne-Loccoz, Reumaux, Carteret &
Eyssartier, Atlas des Cortinaires (Meyzieu) 11: 613. 2001.
Calonarius intricatus (Bojantchev, Ammirati & N. Siegel)
Niskanen & Liimat., comb. nov.
IF554431
Basionym: Cortinarius intricatus Bojantchev, Ammirati &
N. Siegel, in Bojantchev, Index Fungorum 247: 1. 2015.
Calonarius ionochlorus (Maire) Niskanen & Liimat., comb.
nov.
IF554432
Basionym: Cortinarius ionochlorus Maire, Publ. Inst. Bot.
Barcelona 3(no. 4): 113. 1937.
Calonarius jardinensis (Garnica, Ammirati & Halling) Nis-
kanen & Liimat., comb. nov.
IF554433
Basionym: Cortinarius jardinensis Garnica, Ammirati &
Halling, in Garnica, Spahn, Oertel, Ammirati & Oberwin-
kler, BMC Evol. Biol. 11(213 [reprint]): 13 + Additional
file3: 7. 2011.
Calonarius juxtadibaphus (Rob. Henry) Niskanen & Lii-
mat., comb. nov.
IF554434
Basionym: Cortinarius juxtadibaphus Rob. Henry, Bull.
trimest. Soc. mycol. Fr. 99(1): 11. 1983.
Calonarius kristinae (Brandrud) Niskanen & Liimat., comb.
nov.
IF554435
Basionym: Cortinarius kristinae Brandrud, in Frøslev, Bran-
drud & Dima, Mycol. Progr. 16(2): 151. 2017.
Calonarius laberiae (Münzmay, B. Oertel & Saar) Niskanen
& Liimat., comb. nov.
IF554436
Basionym: Cortinarius laberiae Münzmay, B. Oertel &
Saar, Journal des JEC, Journées Européenes du Cortinaire
12(11): 36. 2009.
Calonarius langeorum (Frøslev & T.S. Jeppesen) Niskanen
& Liimat., comb. nov.
IF554437
Basionym: Cortinarius langeorum Frøslev & T.S. Jeppesen,
in Frøslev, Jeppesen & Læssøe, Mycol. Res. 110(9): 1052.
2006.
Calonarius lavandulochlorus (Eyssart.) Niskanen & Liimat.,
comb. nov.
IF554438
Basionym: Cortinarius lavandulochlorus Eyssart., Journal
des JEC, Journées Européenes du Cortinaire 14(no. 13): 53.
2011.
Calonarius lentus (Boccardo, Cleric., Dovana & Vizzini)
Niskanen & Liimat., comb. nov.
IF554439
Basionym: Cortinarius lentus Boccardo, Cleric., Dovana &
Vizzini, in Dovana, Boccardo, Clericuzio & Vizzini, Phyto-
taxa 447(1): 35. 2020.
Calonarius lilacinovelatus (Reumaux & Ramm) Niskanen
& Liimat., comb. nov.
IF554440
Basionym: Cortinarius lilacinovelatus Reumaux &
Ramm, in Bidaud, Moënne-Loccoz, Reumaux, Carteret &
Eyssartier, Atlas des Cortinaires (Meyzieu) 11: 613. 2001.
Calonarius lilaciotinctus (Garnica & Ammirati) Niskanen
& Liimat., comb. nov.
IF554441
Basionym: Cortinarius lilaciotinctus Garnica & Ammi-
rati, in Garnica, Spahn, Oertel, Ammirati & Oberwinkler,
BMC Evol. Biol. 11(213 [reprint]): 13 + Additional file3:
10. 2011.
Fungal Diversity
1 3
Calonarius luteicolor (Ammirati, Bojantchev, Niskanen &
Liimat.) Niskanen & Liimat., comb. nov.
IF554442
Basionym: Cortinarius luteicolor Ammirati, Bojantchev,
Niskanen & Liimat., in Liimatainen, Niskanen, Dima, Kytö-
vuori, Ammirati & Frøslev, Persoonia 33: 126. 2014.
Calonarius luteolus (M.M. Moser) Niskanen & Liimat.,
comb. nov.
IF554443
Basionym: Cortinarius luteolus M.M. Moser, in Moser &
Horak, Beih. Nova Hedwigia 52: 322. 1975.
Calonarius mariekristinae (Brandrud & Dima) Niskanen &
Liimat., comb. nov.
IF554444
Basionym: Cortinarius mariekristinae Brandrud & Dima, in
Brandrud, Schmidt-Stohn & Dima, Sydowia 71: 122. 2019.
Calonarius meinhardii (Bon) Niskanen & Liimat., comb.
nov.
IF554740
Basionym: Cortinarius meinhardii Bon, Docums Mycol.
16(63–64): 66. 1986.
Calonarius metarius (Kauffman) Niskanen & Liimat., comb.
nov.
IF554741
Basionym: Cortinarius metarius Kauffman, Pap. Mich.
Acad. Sci. 1: 137. 1921.
Calonarius mikedavisii (Bojantchev) Niskanen & Liimat.,
comb. nov.
IF554742
Basionym: Cortinarius mikedavisii Bojantchev, Mycotaxon
118: 267. 2011. (2012).
Calonarius moseri (E. Horak) Niskanen & Liimat., comb.
nov.
IF554743
Basionym: Phlegmacium moseri E. Horak, Schweiz. Z.
Pilzk. 40: 93. 1962.
Calonarius murellensis (Cors. Gut., Ballarà, Cadiñanos,
Palazón & Mahiques) Niskanen & Liimat., comb. nov.
IF554744
Basionym: Cortinarius murellensis Cors. Gut., Ballarà,
Cadiñanos, Palazón & Mahiques, Butll. Soc. Micol. Valen-
ciana 10: 160. 2005.
Calonarius napus (Fr.) Niskanen & Liimat., comb. nov.
IF554745
Basionym: Cortinarius napus Fr., Epicr. syst. mycol.
(Upsaliae): 263. 1838. [1836–1838].
Calonarius natalis (D. Antonini & M. Antonini) Niskanen
& Liimat., comb. nov.
IF554746
Basionym: Cortinarius natalis D. Antonini & M. Antonini,
Fungi Non Delineati, Raro vel Haud Perspecte et Explorate
Descripti aut Definite Picti 22: 19. 2002.
Calonarius nymphicolor (Reumaux) Niskanen & Liimat.,
comb. nov.
IF554747
Basionym: Cortinarius nymphicolor Reumaux, in Bidaud,
Moënne-Loccoz, Reumaux & Henry, Atlas des Cortinaires,
Pars V (Annecy): 151. 1993.
Calonarius ochraceopallescens (Moënne-Locc. & Reu-
maux) Niskanen & Liimat., comb. nov.
IF554805
Basionym: Cortinarius ochraceopallescens Moënne-Locc.
& Reumaux, in Bidaud, Moënne-Loccoz, Reumaux, Carteret
& Eyssartier, Atlas des Cortinaires (Meyzieu) 11: 613. 2001.
Calonarius odoratus (Joguet ex M.M. Moser) Niskanen &
Liimat., comb. nov.
IF554806
Basionym: Phlegmacium odoratum Joguet ex M.M. Moser,
Gatt. Phlegm.: 360. 1960.
Fungal Diversity
1 3
Calonarius odorifer (Britzelm.) Niskanen & Liimat., comb.
nov.
IF554807
Basionym: Cortinarius odorifer Britzelm., Ber. naturhist.
Augsburg 28: 123. 1885.
Calonarius olearioides (Rob. Henry) Niskanen & Liimat.,
comb. nov.
IF554808
Basionym: Cortinarius olearioides Rob. Henry, Docums
Mycol. 17(no. 68): 36. 1987.
Calonarius oliveopetasatus (M.M. Moser) Niskanen & Lii-
mat., comb. nov.
IF555319
Basionym: Cortinarius oliveopetasatus M.M. Moser, in
Moser & Ammirati, Mycotaxon 74(1): 29. 2000.
Calonarius olympianus (A.H. Sm.) Niskanen & Liimat.,
comb. nov.
IF555320
Basionym: Cortinarius olympianus A.H. Sm., Contr. Univ.
Mich. Herb. 2: 13. 1939.
Calonarius osloensis (Brandrud, T.S. Jeppesen & Frøslev)
Niskanen & Liimat., comb. nov.
IF555321
Basionym: Cortinarius osloensis Brandrud, T.S. Jeppesen &
Frøslev, in Frøslev, Brandrud & Jeppesen, Mycotaxon 97:
369. 2006.
Calonarius osmophorus (P.D. Orton) Niskanen & Liimat.,
comb. nov.
IF555322
Basionym: Cortinarius osmophorus P.D. Orton, Trans. Br.
mycol. Soc. 43(2): 210. 1960.
Calonarius parasuaveolens (Bon & Trescol) Niskanen &
Liimat., comb. & stat. nov.
IF555417
Basionym: Cortinarius sodagnitus var. parasuaveolens Bon
& Trescol, Docums Mycol. 19(73): 36. 1988.
Calonarius piceae (Frøslev, T.S. Jeppesen & Brandrud) Nis-
kanen & Liimat., comb. nov.
IF555418
Basionym: Cortinarius piceae Frøslev, T.S. Jeppesen &
Brandrud, in Frøslev, Brandrud & Jeppesen, Mycotaxon
97: 372. 2006.
Calonarius platypus (M.M. Moser) Niskanen & Liimat.,
comb. nov.
IF556647
Basionym: Phlegmacium platypus M.M. Moser, Die Gat-
tung Phlegmacium (Schleimköpfe). Die Pilze Mitteleuropas
4: 353. 1960.
Calonarius praetermissus (Bergeron ex Reumaux) Niskanen
& Liimat., comb. nov.
IF556639
Basionym: Cortinarius praetermissus Bergeron ex Reu-
maux, in Bidaud, Moënne-Loccoz, Reumaux & Henry, Atlas
des Cortinaires, Pars V (Annecy): 151. 1993.
Calonarius prasinus (Schaeff.) Niskanen & Liimat., comb.
nov.
IF558128
Basionym: Agaricus prasinus Schaeff., Fung. bavar. palat.
nasc. (Ratisbonae) 4: 51. 1774.
Calonarius pseudocisticola (Boccardo, Dovana, Dima, L.
Albert, Borovička, Mikšík, Saar & Vizzini) Niskanen &
Liimat., comb. nov.
IF557225
Basionym: Cortinarius pseudocisticola Boccardo, Dovana,
Dima, L. Albert, Borovička, Mikšík, Saar & Vizzini, in
Dovana, Boccardo, Borovička, Vizzini, Saar, Albert, Mikšík,
Clericuzio & Dima, Phytotaxa 518(1): 17. 2021.
Fungal Diversity
1 3
Calonarius pseudocupreorufus (Niskanen, Liimat. & Ammi-
rati) Niskanen & Liimat., comb. nov.
IF558777
Basionym: Cortinarius pseudocupreorufus Niskanen, Lii-
mat. & Ammirati, in Liimatainen, Niskanen, Dima, Kytö-
vuori, Ammirati & Frøslev, Persoonia 33: 127. 2014.
Calonarius pseudoglaucopus (Jul. Schäff. ex M.M. Moser)
Niskanen & Liimat., comb. nov.
IF557992
Basionym: Phlegmacium pseudoglaucopus Jul. Schäff. ex
M.M. Moser, Die Gattung Phlegmacium (Schleimköpfe).
Die Pilze Mitteleuropas 4: 354. 1960.
Calonarius pseudoparvus (Bidaud) Niskanen & Liimat.,
comb. nov.
IF557993
Basionym: Cortinarius pseudoparvus Bidaud, in Bidaud,
Moënne-Loccoz, Reumaux, Carteret & Eyssartier, Atlas des
Cortinaires (Meyzieu) 11: 614. 2001.
Calonarius quercus-ilicis (Chevassut & Rob. Henry) Nis-
kanen & Liimat., comb. & stat. nov.
IF557864
Basionym: Cortinarius elegantior var. quercus-ilicis Che-
vassut & Rob. Henry, Docums Mycol. 5(no. 20): 34. 1975.
Calonarius rapaceoides (Bidaud, G. Riousset & Riousset)
Niskanen & Liimat., comb. nov.
IF558098
Basionym: Cortinarius rapaceoides Bidaud, G. Riousset &
Riousset, Micologia 2000 (Trento): 68. 2000.
Calonarius rapaceotomentosus (Delaporte & Eyssart.) Nis-
kanen & Liimat., comb. nov.
IF558101
Basionym: Cortinarius rapaceotomentosus Delaporte &
Eyssart., in Delaporte, Eyssartier & Moënne-Loccoz, Bull.
Soc. mycol. Fr. 118(1): 12. 2002.
Calonarius rufo-olivaceus (Pers.) Niskanen & Liimat.,
comb. nov.
IF558102
Basionym: Agaricus rufo-olivaceus Pers., Syn. meth. fung.
(Göttingen) 2: 285. 1801.
Calonarius sancti-felicis (Frøslev & T.S. Jeppesen) Nis-
kanen & Liimat., comb. nov.
IF558109
Basionym: Cortinarius sancti-felicis Frøslev & T.S.
Jeppesen, in Frøslev, Jeppesen & Læssøe, Mycol. Res.
110(9): 1053. 2006.
Calonarius sannio (M.M. Moser) Niskanen & Liimat.,
comb. nov.
IF558111
Basionym: Cortinarius sannio M.M. Moser, in Moser &
Ammirati, Mycotaxon 72: 315. 1999.
Calonarius saporatus (Britzelm.) Niskanen & Liimat.,
comb. nov.
IF558125
Basionym: Cortinarius saporatus Britzelm., Zur Hymeno-
mycetenkunde 3: 5. 1897.
Calonarius saxamontanus (Fogel) Niskanen & Liimat.,
comb. nov.
IF556154
Basionym: Cortinarius saxamontanus Fogel, Mycologia
86(6): 798. 1995. [1994].
Calonarius selandicus (Frøslev & T.S. Jeppesen) Niskanen