Mission impossible completed: unlocking the nomenclature of the largest and most complicated subgenus of Cortinarius, Telamonia

Abstract

So far approximately 144,000 species of fungi have been named but sequences of the majority of them do not exist in the public databases. Therefore, the quality and coverage of public barcode databases is a bottleneck that hinders the study of fungi. Cortinarius is the largest genus of Agaricales with thousands of species worldwide. The most diverse subgenus in Cortinarius is Telamonia and its species have been considered one of the most taxonomically challenging in the Agaricales. Its high diversity combined with convergent, similar appearing taxa have earned it a reputation of being an impossible group to study. In this study a total of 746 specimens, including 482 type specimens representing 184 species were sequenced. Also, a significant number of old types were successfully sequenced, 105 type specimens were over 50 years old and 18 type specimens over 100 years old. Altogether, 20 epi-or neotypes are proposed for recently commonly used older names. Our study doubles the number of reliable DNA-barcodes of species of C. subgenus Telamonia in the public sequence databases. This is also the first extensive phylogenetic study of the subgenus. A majority of the sections and species are shown in a phylogenetic context for the first time. Our study shows that nomenclatural problems, even in difficult groups like C. subgenus Telamonia, can be solved and consequently identification of species based on ITS barcodes becomes an easy task even for non-experts of the genus.

Full text

Vol.:(0123456789)
1 3
Fungal Diversity
https://doi.org/10.1007/s13225-020-00459-1
Mission impossible completed: unlocking thenomenclature
ofthelargest andmost complicated subgenus ofCortinarius,
Telamonia
KareLiimatainen1 · TuulaNiskanen1,2 · BálintDima3 · JosephF.Ammirati4 · PaulM.Kirk1 · IlkkaKytövuori2
Received: 9 May 2020 / Accepted: 6 August 2020
© The Author(s) 2020
Abstract
So far approximately 144,000 species of fungi have been named but sequences of the majority of them do not exist in the
public databases. Therefore, the quality and coverage of public barcode databases is a bottleneck that hinders the study of
fungi. Cortinarius is the largest genus of Agaricales with thousands of species world-wide. The most diverse subgenus in
Cortinarius is Telamonia and its species have been considered one of the most taxonomically challenging in the Agaricales.
Its high diversity combined with convergent, similar appearing taxa have earned it a reputation of being an impossible group
to study. In this study a total of 746 specimens, including 482 type specimens representing 184 species were sequenced.
Also, a significant number of old types were successfully sequenced, 105 type specimens were over 50 years old and 18
type specimens over 100 years old. Altogether, 20 epi- or neotypes are proposed for recently commonly used older names.
Our study doubles the number of reliable DNA-barcodes of species of C. subgenus Telamonia in the public sequence data-
bases. This is also the first extensive phylogenetic study of the subgenus. A majority of the sections and species are shown
in a phylogenetic context for the first time. Our study shows that nomenclatural problems, even in difficult groups like C.
subgenus Telamonia, can be solved and consequently identification of species based on ITS barcodes becomes an easy task
even for non-experts of the genus.
Keywords ITS· Type study· Barcode· Neotype· Phylogeny· Section
Introduction
So far approximately 144,000 species of fungi have been
named (Willis 2018) but sequences of the majority of them
do not exist in the GenBank or UNITE. Moreover, only a
small percentage of the names in the GenBank, about 4800
Electronic supplementary material The online version of this
article (https ://doi.org/10.1007/s1322 5-020-00459 -1) contains
supplementary material, which is available to authorized users.
* Kare Liimatainen
k.liimatainen@kew.org
Tuula Niskanen
tuula.niskanen@cortinarius.fi
Bálint Dima
cortinarius1@gmail.com
Joseph F. Ammirati
cort@u.washington.edu
Paul M. Kirk
p.kirk@kew.org
Ilkka Kytövuori
ilkka.kytovuori@gmail.com
1 Jodrell Laboratory, Royal Botanic Gardens, Kew,
SurreyTW93AB, UK
2 Botanical Museum, University ofHelsinki, P.O. Box7,
00014Helsinki, Finland
3 Department ofPlant Anatomy, Institute ofBiology,
Eötvös Loránd University, Pázmány Péter sétány 1/C,
1117Budapest, Hungary
4 Department ofBiology, University ofWashington,
Box351800, Seattle, WA98195-1800, USA

Fungal Diversity
1 3
species, are based on sequences from type materials or other
reliable sources (Schoch etal. 2014).
Currently species identification of fungi in academic stud-
ies is almost solely based on nrDNA ITS barcodes (Lindahl
etal. 2013). Thus, those collections with taxonomically cor-
rect names that are not in any public sequence repositories
are basically omitted in academic research. Therefore, the
quality and coverage of public barcode databases is a bot-
tleneck that hinders the study of fungi (Schoch etal. 2014).
Depositing the ITS sequences in public repositories like
GenBank does not automatically make them useful for iden-
tification. Two excellent platforms for delivering sequence-
based identification information for the end-users include
RefSeq under GenBank (Schoch etal. 2014) and UNITE
(Kõljalg etal. 2013). However, in both cases an extra step by
an expert, in addition to the normal sequence submission, is
required, but unfortunately often is left undone, making part
of the already existing information unusable.
Cortinarius (Pers.) Gray is the largest genus of Agaricales
with thousands of species world-wide (Kirk etal. 2008). They
are important ectomycorrhizal fungi and often discovered in
ecological studies. Only three large studies of type specimens
based on ITS sequence data in Cortinarius have been made
so far. Two of them are from C. subgen. Phlegmacium (Fr.)
Trog: Liimatainen etal. (2014) includes over 230 sequences
of type specimens representing over 150 species and Frøslev
etal. (2007) has over 50 sequences of 79 species. The third
one is from C. subgen. Telamonia (Fr.) Trog and includes
over 60 sequences of 33 species (Liimatainen etal. 2017).
The most species-rich subgenus in Cortinarius is Telamo-
nia. Members of this subgenus mainly occur in the Northern
Hemisphere (Garnica etal. 2005, Soop etal. 2019) and are
especially dominant in coniferous forests. Six relatively large
phylogenetic studies of Cortinarius have been published so
far (Garnica etal. 2005, 2016; Harrower etal. 2011; Peint-
ner etal. 2004; Soop etal. 2019; Stensrud etal. 2014). The
number of species of Telamonia included in these studies
varies from 8 to 70, and the phylogenies have been mainly
based on the analysis of ITS and LSU regions, with some also
including sequences of rpb1 or rpb2 regions. These studies
have shown that the majority of the traditionally morpho-
logically delimited species of Telamonia (Bidaud etal. 2017;
Brandrud etal. 2012; Moser 1983; Niskanen etal. 2012)
form a monophyletic group. However, sections or subgenera
that have been shown not to belong to Telamonia include
Anomali Konrad & Maubl., Balaustini Moënne-Locc. &
Reumaux, Camphorati Liimat., Niskanen & Ammirati, Ful-
vescentes Melot, Illumini Liimat., Niskanen & Kytöv., Obtusi
Melot, Renidentes Moënne-Locc. & Reumaux, and Rigentes
Melot. Morphological characters that define the species of
Telamonia are basidiomata with dry pileus and dry stipe and
mainly brownish colours, with the exception of some whitish,
bluish-purple or orange-red species. No larger phylogenetic
analysis on the infrasubgeneric relationships have been done
so far, but studies on specific sections including sequences
from type specimens have been published: Armillati M.M.
Moser (Niskanen etal. 2011), Bovini M.M. Moser (Niskanen
etal. 2013), Brunnei Melot (Niskanen etal. 2009), Colymba-
dini Melot/Cinnabarini Melot/Uracei Melot (Ammirati etal.
2017; Dima etal. 2014), Disjungendi Kytöv., Liimat., Nis-
kanen & Ammirati (Liimatainen etal. 2015), Hydrocybe (Fr.
ex Rabenh.) Gillot & Lucand (Suárez-Santiago etal. 2009)
and Saturnini Moënne-Locc. & Reumaux/Bicolores (M.M.
Moser) Melot (Liimatainen etal. 2017).
The nuclear ribosomal internal transcribed spacer (ITS),
which has been proposed as the universal barcode marker for
fungi (Schoch etal. 2012), is also the main locus used in the
species level taxonomy of Cortinarius. The treshold value for
barcoding Cortinarius species has been proposed to be 99%
(Garnica etal. 2016). However, there already is evidence that
a few morphologically distinct Telamonia species only have
1 base difference (99.8% similarity) in the ITS region, e.g.
C. laniger Fr./C. solis-occaus Melot (Niskanen etal. 2012)
and C. paragaudis Fr./C. pinigaudis Niskanen, Kytöv. & Lii-
mat. (Niskanen etal. 2011) and in the case of C. confirma-
tus Rob. Henry the intraspecific variation is > 1%, although
the species has a wide morphological and ecological range
and based on ITS sequences there are 3 supported subclades
which might be separate taxa (Liimatainen etal. 2017).
In this study our aim was to provide a revision of Corti-
narius, subgen. Telamonia as well as an extensive ITS data-
base for the identification of the species. Almost all type
specimens of the species described in the subgenus were
studied and an epi- or neotype is proposed for all recent
frequently used older names when possible. In addition, a
phylogenetic tree is produced as a framework for the infrasu-
bgeneric classification of the species; including many that
are included in a phylogenetic analysis for the first time.
Materials andmethods
Taxon sampling
The type specimens of Telamonia species published over
many years by Ammirati, D. Antonini, M. Antonini,
Bergeron, Bidaud, Bohus, Bouteville, Bresadola, Cart-
eret, Chevassut, Consiglio, Daniele, Eyssartier, A. Favre,
J. Favre, Fellner, Ferville, Fillion, Henry, Hesler, Høiland,
Hongo, Karsten, Kauffman, Kühner, Landa, Lindström,
Matheny, McKnight, Moser, Moënne-Loccoz, Murril, Nes-
piak, Orton, Pearson, Peck, Ramm, Reumaux, Sasia, Seidl,
Smith, Schwöbel, Soop, Svrček, Velenovský, and Vialard
were sampled as well as all the Telamonia collections pub-
lished and illustrated in Brandrud etal. (1989, 1992, 1994,
1998). A total of 482 types are included here. An additional

Fungal Diversity
1 3
183 previously published sequences of Telamonia types
were added to our dataset for the best overview of current
available data.
We aimed to have at least two sequences per species in
our study. Therefore, some additional sequences, either our
own unpublished ones or from databases GenBank and
UNITE, were included. Information on the sequences of
type specimens is available in Supplementary Table1 and
information on other sequences included in the phylogenetic
analysis is available in Supplementary Table2. Fungarium
acronyms follow Index Herbariorum (Thiers 2013).
Species concept
Based on criteria mentioned in the introduction we have used
1 % (5 differences) as a cut-off value for species. When type
sequences differ in at least 5 sites from one another we have
treated them as different species. We are not claiming that all
the variation below 1 % is automatically intraspecific. Separat-
ing species below the 1% cut-off value, however, does require
careful study. Therefore, we have added ‘aff.’ prefix to the Latin
name in cases where there are 3 to 4 differences to another type
sequences. With 2 differences we have used the s. lato nota-
tion in the Fig.1 and Supplementary Tables1 and 2. Using
this approach indicates places where determining taxonomic
synonyms might be problematic and require further study. Also,
when macroscopic, microscopic and/or ecology data differ con-
siderably although the ITS sequences are the same, we have not
placed the taxa in synonomy. Furthermore, in cases where a
species complex has previously been shown to include several
species supported by morphology and small, but constant bar-
code gaps, we have avoided making synonymys.
One cannot emphasize enough that using a small cut-off
value requires good quality sequences. In this study all the
specimens have been sequenced from both directions and the
chromatograms of the sequences were checked and edited
manually before any preliminary analyses. When small, less
than ten base or indel changes and/or odd differences are found
between sister species or within species those differences have
been confirmed by combining the relevant chromatograms
and checking manually the base sites that differ. Also, base or
length polymorphisms sites are not counted as a difference and
an indel is counted as one difference despite its length.
Molecular analyses
DNA was extracted from a few milligrams of dried material
(a piece of lamella) with the NucleoSpin Plant kit (Mach-
erey-Nagel, Düren, Germany). The same protocol was used
for all materials. Primers ITS 1F and ITS 4 (Gardes and
Bruns 1993; White etal. 1990) were used to amplify ITS
regions. The same primer pairs were used in direct sequenc-
ing. For problematic material the primer combinations ITS
1F/ITS 2 and ITS 3/ITS 4 were also used. PCR amplifica-
tions were performed in a 25 µl reaction mix with about 70
ng extracted DNA, 1 U Phusion High-Fidelity DNA poly-
merase and 1× HF buffer (ThermoFisher), 200 mM of each
dNTP and 0.5 µM of each primer. The PCR were run on
a MBS 0.2 G Thermal Cycler (Thermo Hybaid) with the
following settings: denaturation for 30 s at 98 °C, followed
by 35 cycles of denaturation for 10 s at 98 °C, annealing
for 30 s at 50 °C, and extension for 30 s at 72 °C. The PCR
products were purified using an ExoSAP-IT purification kit
(Amersham Biosciences). Sequencing was performed on
both strands using a BigDye Terminator v1.1 Sequencing
kit (Applied Biosystems). Reactions were performed in 10 µl
with 1 µl of PCR product, 1.3 mM of primer (ITS 1F or ITS
4), 1 µl 5X sequencing buffer, and 1 µl of Terminator Ready
Reaction Mix. Reactions were run for 1 min at 96 °C, fol-
lowed by 30 cycles of 30 s at 96 °C, 15 s at 50 °C, and 4 min
at 60 °C. Unincorporated dye terminators and primers were
removed by Sephadex G-50 DNA Grade Fine (Amersham
Biosciences) purification system, and the reactions were
analysed by ABI 3730 DNA Analyzer (Applied Biosystems)
automatic sequencer. Sequences were assembled and edited
with Sequencher 4.1 (Gene Codes, Ann Arbor, Michigan,
USA). A total of 755 new ITS sequences were produced for
this study. Collections and GenBank sequences used for the
phylogenetic analysis are given in Supplementary Tables1
and 2.
The short ITS sequences of type specimens were excluded
from the phylogenetic analysis. To improve the resolution
of phylogenetic analyses we included 146 published LSU
sequences from GenBank to our dataset. The chosen LSU
sequences are from different parts of Telamonia and they
were mostly obtained from Garnica etal. (2005), Harrower
etal. (2011), and Stensrud etal. (2014). Sequences from
section Dermocybe Pers. were selected as an outgroup based
on Stensrud etal. (2014). A total of 919 ITS and 146 LSU
sequences were aligned separately for both regions using
MAFFT 7 (Katoh and Standley 2013) with the G-ING-i
algorithm (Katoh etal. 2005). The alignments were then
manually improved in SeaView (Galtier etal. 1996). The
phylogenetically informative indels in the ITS region were
coded as characters following the simple indel coding algo-
rithm (Simmons and Ochoterena 2000) with FastGap 1.2
(Borchsenius 2009). The binary and aligned nucleotide data
were concatenated in Mesquite 3.2 (Maddison and Maddison
2017). The alignment is 2008 nucleotides long (including
gaps) and is available at TreeBASE under S26824 (http://
www.treeb ase.org/treeb ase-web/home.html). A phylogenetic
tree was generated from the concatenated dataset using max-
imum likelihood (ML) analyses with 1000 bootstrap repli-
cates under the GTRGAMMA model for nucleotide parti-
tions (ITS+LSU) and the default setting for binary (indel)
data in RAxML 8 (Stamatakis 2014).

Fungal Diversity
1 3
Fig. 1 A phylogram resulting
from the RAxML analysis of
the concatenated dataset includ-
ing 919 aligned ITS and 146
LSU sequences and the binary
data resulting from the coding
of the phylogenetically informa-
tive insertions/deletions in the
ITS alignment. Bootstrap values
greater than 50% are indicated
above branches. The current
names of species are in boldface
C. diasemospermus CFP657 Sweden
C. udolivascens var. lilacinostipitatus 2001-112 TYPE France
C. conocyboides 99102504 TYPE France
C. altipes 01-10-134 TYPE France
C. distinctus 2004-165 TYPE France
C. pilatii 655887 TYPE Czech
C. pelargoniobtusus RH70579 TYPE France
100
99
92 C. fragrantissimus 10MWB111913 TYPE USA, WA
C. fragrantissimus GQ159881 Canada, BC
C. subparvannulatus 1728 TYPE France
C. subparvannulatus UDB002161 Sweden
C. rusticellus 13512 TYPE Switzerland
84
C. rusticellus 6029429 Finland
99
C. plicatus 91-08-51 TYPE France
C. vulpicolor 83/355 TYPE USA, MT
C. glandicolor var. exilis 13282 TYPE Switz.
99
88
83
65
60
98
100
C. subdepressus 4705 TYPE France
C. difficillimus 97.12.28.08 TYPE France
C. goniosporus 2001-138 TYPE France
C. ammophilus 109694 TYPE UK
C. desertorum 154750 TYPE Czech
C. mimicus 2003-57 TYPE France
C. friesianus 97112425 TYPE France
C. pertristis 13308 TYPE Switzerland
C. impolitus 10366 TYPE USA, MI
C. violilamellatus CFP1304 Sweden
C. aff. cucumisporus IK01-046 Sweden
C. umbrinolutescens 2858 TYPE France
C. umbrinolutescens JQ347072 China
001001
55
C. pelargoniostriatulus 07-08-42 TYPE France
C. pelargoniostriatulus TN09-080 USA, WA
C. sp. GQ159914 Canada, BC
100
100
69
94 C. fagetorum GQ159777 Canada, BC
C. fagetorum MM1948-0743 TYPE Austria
C. fagetorum TN07-179 Canada, ON
C. violaceopapillatus 06-10-196 TYPE France
C. violaceopapillatus IK11-020 Norway
C. geraniolens 95-09-88 TYPE France
C. subcarcharias 03-11-74 TYPE France
C. furfuraceus 1240 TYPE France
C. fuscoruber 909 TYPE France
100
68
C. gurdus 2006-170 TYPE France
C. flabellus CFP672 TYPE Sweden
C. flabellus f. iners 00-09-81 TYPE France
C. pseudodepressus 2009-73 TYPE France
C. furfuraceus AY669678
100
10
0
100
55
97 C. flabellus CFP690 Swedenflexipes var.
C. CFP626 TYPE Swedenlindstroemii
C. flabellus f. biolens 01-09-121 TYPE France
C. lindstroemii FJ039709 Canada, BC
C. pseudoflabellus 96-08-16 TYPE France
C. pseudoflabellus 6001842 Finland
C. flexipes CFP802 TYPE Sweden
C. paleifer 655884/a TYPE Czech
C. paleifer 655884/b TYPE Czech
C. flexipes KC842395 Norway
sect.
Flexipedes
88

Fungal Diversity
1 3
C. andreae CFP1309 TYPE Sweden
sect. Hinnulei
sect. Sporagniti
C. andreae TN07-392 USA, WA
C. sporagnitus 92-11-418 TYPE France
98
98
72
73
C. lux-nymphae CFP984 Sweden
C. pinisquamulosus 6029807 TYPE Finland
C. heterosporus TN05-162 Finland
C. heterosporus 14370 TYPE Germany
100 sect. Heterospori
100
100
93
81
99
98
C. psammocephalus CFP485 TYPE Sweden
C. quercoconicus TN03-1305 TYPE Sweden
C. psammocephalus AY669672
C. olivaceomarginatus 99.10.09.03 TYPE France
C. pseudobavaricus 1184 TYPE France
C. castaneopallidus 2001-124 TYPE France
C. sp FJ039534 Canada, BCC. sp FJ039534 Canada, BC
C. sp. GQ159913 Canada, BC
C. angustisporus KP165560 TYPE Finland
C. angustisporus KP165561 Finland
sect. Ochrovelati
C. flavoperonatus 97-10-384 TYPE France
C. pseudofusisporus 92-10-262 TYPE France
100
100
100
98
96
97
C. privus 4550 TYPE France
C. subfuscodiscus 5254 TYPE France
C. olivacobrunneus 2009-79 TYPE France
C. flexibilifolius 2003-58 TYPE France
C. flexibilifolius AY669677
C. megacystidiosus TN12-042 Finland
C. megacystidiosus 2009-81 TYPE France
C. semivestitus TN09-121 USA, WA
C. fusisporus var. olivaceodepressus 860 TYPE France
C. tubulipes 13317 TYPE Switzerland
C. angelesianus GQ159864 Canada, BC
C. strobilaceus 53/24 TYPE Austria
C. angelesianus CFP359 Sweden
96
93
88
97
75
100
100
94
99
57
89
97
70
100
C. pseudohinnuleus 4224 TYPE France
C. salicum 4479 TYPE France
C. lepidus 765 TYPE France
C. aciculisporus 257 TYPE France
C. herculinus 4195 TYPE France
C. ochraceoplicatus 378 TYPE France
C. hinnuleoscitus RH9221 TYPE France
C. luridus
C. subrigidipes MM1991-0309 TYPE USA, WY
C. pangloius MM1964-0110 TYPE Sweden
C. paraphaeochrous MM1991-0323 TYPE USA, WYC. paraphaeochrous MM1991-0323 TYPE USA, WY
C. ferrugineifolius MM1991-0305 TYPE USA, WY
C. hinnuleus var. minutalis CFP969 Sweden
C. subulatus 4229 TYPE France
C. griseascens 4263 TYPE France
C. nauseosmus 4214 TYPE France
C. semiodoratus RH91.6 TYPE France
C. semiodoratus AY669665
C. solidus 4253 TYPE France
C. albolens TN12-045 Finland
C. albolens 97-10-368 TYPE France
C. roseonudipes 37 TYPE France
C. hinnuleovelatus 4203 TYPE France
C. buxiolens 3300 TYPE France
C. subfilamentosus 1196 TYPE France
C. carcharias 4276 TYPE France
C. badioflavidus KU041724 USA, CA
C. badioflavidus KU041723 TYPE USA, WA
C. hinnuleocervinus TN12-175 TYPE USA, CA
C. JFA13502 USA, WAhinnuleocervinus
C. hinnuleoarmillatus RH88-6 France
C. aureifer 351 TYPE France
C. tigris 4269 TYPE France
C. hinnuleus CFP332 TYPE Sweden
C. hinnuleus AY669667
C. hinnuloides var. phaeopus 4285 TYPE France
C. hinnuloides var. phaeopus TN04-1139 Italy
sect. Flexibilifolii
Fig. 1 (continued)

Fungal Diversity
1 3
sect. Leiocastanei
sect. Rubricosi
100
100
100
63
C. fulvoisabellinus KJ206485 TYPE France
C. radicosissimus 142 TYPE France
C. subhelvolus 470 TYPE France
C. squamulifer 4260 TYPE France
C. speciosior 4280 TYPE France
sect. Hinnulei
C. carbunculus CFP465 TYPE Sweden
C. carbunculus AB828023 Japan
C. lacustris 4232 TYPE France
C. altae-herbae 99669 TYPE France
100
100
66
99
C. aavae NR120095 TYPE Canada, QC
C. aavae FJ717539 Canada, BC
C. leiocastaneus EU259286 Finland
C. leiocastaneus NR119678 TYPE Finland
C. umbrinolens CFP609 Sweden
C. umbrinolens AY669658
C. sericeofibrillosus 5158 TYPE France
95
97
66
50
99
100
88
90
100
89
100
99
100
88
100
55
54
77 100
100
100
98
100
85
88
100
100
C. milvinicolor 4217 TYPE France
C. milvinicolor AY669673
C. elaphinicolor 2001-136 TYPE France
C. alboadustus 92-10-252 TYPE France
C. parhonestus 3657 TYPE France
C. parhonestus TN03-1552 Sweden
C. minusculus TN12-031 Finland
C. minusculus UDB026212 Estonia
C. sp. GQ159898 Canada, BC
C. scotoides 13312 TYPE Switzerland
C. scotoides KC842394 Norway
C. castaneoruber 604 TYPE France
C. helobius CFP542 Sweden
C. sublucorum 2005-240 TYPE France
C. subscotoides TN12-010 TYPE Finland
C. subscotoides TN12-015 Finland
C. subexitiosus KP165574 TYPE USA, WA
C. subexitiosus KP165575 Finland
C. exitiosus 3933 TYPE France
C. safranopes var. bulbosus 748 TYPE France
C. safranopes 364 TYPE France
C. calcareophilus 4293 TYPE France
C. conicus CFP1087 France
C. rubricosus 983 TYPE France
C. comptulus MM1965-0250 TYPE Switzerland
C. inolens var. parvinolens 99-11-369 TYPE France
C. griseosulcatus 99102503 TYPE France
C. fistularioides 1031 TYPE France
C. sublatisporus 625955 TYPE Czech
C. laniatus RH70964 TYPE France
C. comptulus CFP663 Sweden
C. jacobii 4104 TYPE France
C. comptulus FJ039543 Canada, BC
C. pilatii CFP605b Sweden
C. nigrocuspidatus FJ039708 Canada, BC
C. adalbertii var. turritus MM1979-0539 TYPE Sweden
C. striatulorufus 3022 TYPE France
C. russulaespermus 2003-46 TYPE France
C. subobtusus 10417 TYPE USA, MI
C. subobtusus IK11-007 Norway
C. subobtusus GQ159883 Canada, BC
C. pallescens 347 TYPE France
C. pseudosafranopes 4238 TYPE France
C. subgrisescens 4234 TYPE France
C. hinnuleoradicatus 4282 TYPE France
C. epipurrus RH3377 TYPE France
C. mucicola TN09-205 USA, WA
C. mucicola TN12-197 USA, CA
C. uraceoarmillatus 05-10-272 TYPE France
C. rubrocinctus IK01-044 Finland sect. Rubrocincti
syn. sect. Safranopede
s
Fig. 1 (continued)

Fungal Diversity
1 3
sect. Paleacei
sect. Bibuli
sect. Helvelloides
sect. Verni
100
98
100
80
69
80
82
72
C. hemitrichus CFP662 TYPE Sweden
C. fusisemen 3970 TYPE France
C. milvinoides 2009-84 TYPE France
C. hemitrichus AY669680
C. roseohemitrichus 99.10.23.09 TYPE France
C. subpaleaceus IK08-995 TYPE Finland
C. subpaleaceus TN09-029 USA, WA
C. pilatii CFP605a Sweden
C. boreotrichus 6032724 TYPE Finland
C. expallens MM1991-0322 TYPE USA, WY
C. mucronatus 1983-0342 USA, WY
C. gossypinus CFP1344 Sweden
C. hemitrichus f. improcerus 13291/150 TYPE Czech
C. incisior TN04-1041 Finland
C. incisior TN12-043 Finland
C. jacobi-langei 98-11-417 TYPE France
C. mallaensis 6033461 Finland
C. mallaensis 6033453 TYPE Finland
C. saniosus DQ102671 TYPE Sweden
C. bavaricus MM1967-0102 TYPE Germany
C. luteolateritius 154749 TYPE Czech
C. rufoanuliferus 1983/386 TYPE USA, WY
C. saniosus AY669660
100
60
100
90
100
100
96
69
78
100
96
100
100
54
86
100
100
70
100
99
69
100
93
60
63
93
100
66
93
C. subaurantiomarginatus 5249 TYPE France
C. saniosus GQ159783 Canada, BC
C. TN10-177 Canada, QCaff. gentilissimus
C. aureovelatus DQ102652 TYPE Norway
C. af IK01-041 Finlandf. gentilissimus
C. cf. gentilissimus JFA11923 Costa Rica
C. cf. gentilissimus JFA12032 Costa Rica
C. saniosus var. paludophilus 1182 TYPE France
C. paludosaniosus TN07-342 TYPE USA, WA
C. paludosaniosus AY669621
C. fuscogracilescens 00274011 TYPE France
C. fuscogracilescens TN03-1348 Sweden
C. corvinus 1188 TYPE France
sect. Saniosi
C. americanus GQ398244 USA, OR
C. americanus TN07-473 USA, WA
C. americanus 10314 TYPE USA, MI
C. sphagnicola 3934 TYPE France
C. americanus HE979238 China
C. bibulus UDB002795 Estonia
C. bibulus TN05-119 TYPE Finland
C. fuscodiscus FJ039555 Canada, BC
C. fuscodiscus TN09-069 USA, WA
C. sphagnoravus AY669683
C. sphagnoravus CFP746 TYPE Sweden
C. atripes 436 TYPE France
C. griseocarneus 2004-163 TYPE France
C. alnetorum CFP339 Sweden
C. griseocarneus AY669695
C. helvelloides AY669684
C. helvelloides TN05-002 TYPE Finland
C. helvelloides CFP340 Sweden
C. helodes FJ039707 Canada, BC
C. helodes MM1997-0311 TYPE USA, WY
C. lacorum 10370 TYPE USA, MI
C. olivaceofulvus 10384 TYPE USA, MI
C. badiovestitus MM1965-0040 TYPE Austria
C. erythrinus var. russulisporus 58132 TYPE Hungary
C. petroselineus var. radicipes FN429003 TYPE France
C. vernus UDB000742 TYPE Sweden
C. sp. FJ039541 Canada, BC
sect. Alnicolarum
sect. Ravi
sect. Pseudoduracini
sect. Incisiores
Fig. 1 (continued)

Fungal Diversity
1 3
sect. Verni
sect. Atroalbi
100
85
56
66
99
99
61
100
100
100
80
C. sp. GQ159826 Canada, BC
C. sp. GQ159811 Canada, BC
C. suberythrinus 056 TYPE France
C. majorinus 219 TYPE France
C. tenebricus 13514/168a TYPE Switz.
C. suberythrinus AY669690
C. psammocola IK99-722 TYPE Finland
C. ammophiloides 57443 TYPE Hungary
C. salicetophilus 96-10-169 TYPE France
C. ochrorufus 2174 TYPE France
C. punctatorum 2172 TYPE France
C. subfistularis 2719 TYPE France
C. tener 4813 TYPE France
C. helobius CFP542 Sweden
C. nigromaculatus 98.06.05.10 TYPE France
C. puellaris TN16-504 UK
C. puellaris KT591586 TYPE Norway
C. nucicolor KP137510 TYPE Sweden
C. intempestivus 1157 TYPE France
C. cristatosporus 1149 TYPE France
C. parinsignis 18 TYPE France
C. subtilior 13316/167 TYPE Switz.
C. subtilior FJ039542 Canada, BC
C. sp. GQ159878 Canada, BC
C. furvoumbrinus KP137497 TYPE Sweden
C. subcastaneus 5092 TYPE France
C. tenebrosus 5100 TYPE France
C. castaneus var. nigrescens 20599 TYPE France
C. caliginosus 4046 TYPE France
C. caliginosus TN03-1346 Sweden
C. subobtusobrunneus 1691 TYPE France
C. subobtusobrunneus HM189747
100
100
90
100
52
100
89
52
66
74
71
66
78
96
98
98
68
81
66
64
C. hirtus 154753 Czech
C. similigenus 2626 TYPE France
C. querculus 3885 TYPE France
C. punctatoides 2009-72 TYPE France
C. hirtus 154756 TYPE Czech
C. substemmatus 2716 TYPE France
Telamonia sanguinescens 154758 TYPE Czech
C. casimiri CFP656 Sweden
C. hirtus FJ039548 Canada, BC
C. hirtus FJ039552 Canada, BC
C. rubellopes var. pudoricolor 373 TYPE France
C. decipientoides 367 TYPE France
C. casimiri CFP918 Sweden
C. derelictus 4107 TYPE France
C. hemitrichoides 2157 TYPE France
sect. Megaspori
C. roseobrunneus UDB026042 Estonia
C. roseobrunneus 4707 TYPE France
C. denigratus KU041734 TYPE USA, WA
C. denigratus OC2090812-03 USA
C. denigratus GQ159815 Canada, BC
C. sp. GQ159818 Canada, BC
C. sp. AY669685
C. atroalbus AY669687
C. paludophilus 4974 TYPE France
C. ignotissimus 96-10-96 TYPE France
C. atroalbus var. nigripes MM1987-0317 TYPE USA, WY
C. biformis var. dilatus 08-09-245 TYPE France
C. famatus 4099 TYPE France
C. atroalbus AY669687
C. fuscoalbus UDB016059 Estonia
C. fuscoalbus
CFP760 TYPE Sweden
sect. Megaspori
sect. Caliginosi
sect. Nucicolores
Fig. 1 (continued)

Fungal Diversity
1 3
sect. Duristipedes
sect. Castanei
100
sect. Atroalbi
99
90
100
100
70
100
99
100
64
100
100
100
100
53
C. bohemicus f. subheterosporus 92-11-419 TYPE France
C. heterodepressus 6031902 TYPE Finland
C. roseomyceliosus IK94-1255 Finland
C. roseomyceliosus 01-10-144 TYPE France
C. duristipes KP165559 USA, AK
C. duristipes KP165557 TYPE Finland
C. nigrellus IK01-063 Finland
C. nigrellus TN05-173 Finland
C. punctatiformis CFP1479 Italy
C. punctatiformis 99.10.03.03 TYPE France
C. magus 3006 TYPE France
C. fulguritans 3887 TYPE France
C. poirieri 800 TYPE France
C. rufescentipes 5095 TYPE France
C. sensibilis 00-09-114 TYPE France
C. praestigiosus TN12-028 TYPE Finland
C. fumosifolius 10354 TYPE USA, MI
C. subtabularis 10428 TYPE USA, MI
C. inconspicuus IK01-043 Finland
C. inconspicuus KF617576 USA, AK
C. roseobasilis KU041741 USA, WA
C. roseobasilis KU041740 TYPE USA, WA
C. roseocastaneus KP114461 TYPE Finland
C. roseocastaneus TN08-072 Finland
100
C. fuscoumbrinus TN03-1553 Sweden
C. fuscoumbrinus KP137495 TYPE Sweden
C. vinaceobrunneus GQ159893 Canada, BC
C. vinaceobrunneus KU041742 TYPE USA, WA
100
100
97
64
83
100
71
76
94
C. ferrugineovelatus GQ159884 Canada, BC
C. ferrugineovelatus KP137508 TYPE Sweden
C. umbrinobellus KP137507 TYPE Sweden
C. umbrinobellus AB831855 Japan
C. anthracinoides 74 TYPE France
C. decipiens 366 TYPE France
C. decipiens var. decipiens CFP589 Sweden
C. decipiens f. saliceticola 97112412 TYPE France
C. argumentosus 4092 TYPE France
C. maculatophyllus 94-10-220 TYPE France
C. decipiens var. atrocaeruleus CFP561a Norway
C. fuscoflexipes MM1983-0384 TYPE USA, WY
C. fuscoflexipes TN11-420 USA, WA
C. griseophyllus 2625 TYPE France
C. falsosus 3886 TYPE France
C. griseovioleipes 889 TYPE France
C. recedens 890 TYPE France
55
89
60
94
98
64
99
87
C. carminipes 463 TYPE France
C. robertii 051 TYPE France
C. fallaciosus 4423 TYPE France
C. kunicensis var. caespitosus 69 TYPE France
C. castaneus 275 TYPE France
C. minutulus 13297/a TYPE Switzerland
C. minutulus 13297/b TYPE Switzerland
C. subodoratus 5162 TYPE France
C. atrocaeruleus GQ159796 Canada, BC
C. insignitus 3831 TYPE France
C. atrocaeruleus FN428992 TYPE Austria
C. sertipes FN429001 TYPE France
C. contrarius FN429000 TYPE Netherlands
C. erythrinellus 553 TYPE France
C. sp. GQ159816 Canada, BC
C. sp. FJ039539 Canada, BC
C. sp. GQ159855 Canada, BC
C. sp. GQ159824 Canada, BC
sect. Vinaceobrunnei
sect. Friesiorum
sect. Praestigiosi
sect. Punctatiformes
Fig. 1 (continued)

Fungal Diversity
1 3
sect. Parvuli
sect. Anthracini
sect. Crassispori
sect. Saturnini
100
100
81
C. tuolumnensis JQ906762 TYPE USA, CA
C. tuolumnensis GQ159813 Canada, BC
C. gualalaensis FJ039536 Canada, BC
C. gualalaensis JX501775 TYPE USA, CA
C. ohlone JQ906760 USA, CA
C. ohlone JQ906757 TYPE USA, CA
89
94
89
100
92
92
98
100
98
100
82
71
100
99
C. parvannulatus s. Funga Nordica CFP491 Sweden
C. parvannulatus UDB001025 Greenland
C. parvannulatus s. lato AY669664
C. incisopunctatus 892 TYPE France
C. neofallax KF048129 TYPE France
C. pseudofallax NR131831 TYPE France
C. pseudofallax KF048131 Finland
C. cistopulchripes 00.10.226 TYPE France
C. cistopulchripes JFA10095 USA, WA
C. sp. KC842397 Norway
C. occidentalisagacitas TN09-118 TYPE USA, WA
C. occidentalisagacitas EU821662 Canada, BC
C. sagacitas 6033517 TYPE Finland
C. urdaibaiensis FN428983 TYPE Spain
C. subcoronatus FN428985 Spain
C. subcoronatus 5157 TYPE France
C. bombycinus FN428987 TYPE Spain
C. albosericeus GQ159846 Canada, BC
C. albosericeus KU041721 TYPE USA, WA
C. albosericeus KU041722 TYPE USA, WA
98
100
97
C. campester 3883 TYPE France
C. malefidus 3035 TYPE France
C. porphyreticus 3866 TYPE France
C. auroripes 4798 TYPE France
C. subgracilior 95-10-126 TYPE France
C. gallurae KU953934 Spain
C. gallurae FN428979 TYPE Italy
C. miwok FJ039540 Canada, BC
C. miwok JQ906753 TYPE USA, CA
C. fulvopaludosus 6033460 TYPE Finland
C. fulvopaludosus 6033534 Finland
100
74
100
65
63
100
100
100
88
sect. Squalidi
100
56
98
97
67
C. josserandii 3625 TYPE France
C. subanthracinus 3037 TYPE France
C. procalans 3051 TYPE France
C. danili 540 TYPE France
C. anthracinus s. Funga Nordica CFP804 Sweden
C. anthracinus AY669670
C. colorius 3601 TYPE France
C. col CFP488 Norwayus
C. subminiatopus 6031919 TYPE Finland
C. miniatopus 6041343 TYPE Finland
C. miniatopus FJ039535 Canada, BC
C. miniatopus IK08-997 Finland
C. chevassutii CFP1277a Hungary
C. chevassutii RH70955 TYPE France
C. duboisensis KU041735 TYPE USA, WA
C. duboisensis KU041736 USA, WA
C. crassisporus KP165556 USA, AK
C. crassisporus NR131882 TYPE Sweden
C. sefendens RH70947 TYPE France
C. semivelatus RH2590 TYPE France
C. squalidus 10409 TYPE USA, MI
C. assiduus var. plesiocistus AM713178 TYPE Spain
C. bulbosovolvatus KX964440 TYPE Italy
C. confirmatus AM713179 Spain
C. confirmatus KX964438 TYPE France
sect. Bombycini
sect. Castanei
Fig. 1 (continued)

Fungal Diversity
1 3
sect. Politi
sect.
Boulderenses
sect. Brunneocalcari
sect. Brunneifolii
sect. Urbici
sect. Rustici
sect. Lanigeri
sect. Exsulares
sect. Saturnini
100
100
76
86
97
82
100
54
96
99
82
99
90
87
66
90
93
100
100
100
100
100
98
98
71
100
71
100
65
54
100
100
100
100
C. betulaecomes KX964479 TYPE France
C. imbutus KX964498 TYPE Finland
C. stuntzii KX964558 TYPE USA, WA
C. gramineus KX964522 TYPE France
C. saturninus KX964584 TYPE Sweden
C. incarnatolilascens KX964508 TYPE France
C. lucorum KX964585 TYPE Norway
C. lucorum EU821673 Canada, BC
C. politus NR131829 TYPE USA, WA
C. politus KC608587 USA, WA
C. furvus KP137497 TYPE Finland
C. sp. FJ039561 Canada, BC
C. hepaticus TN12-052 TYPE Finland
C. hepaticus IK04-051 Finland
C. fuscescens KP165549 Finland
C. fuscescens NR131879 TYPE Finland
C. boulderensis var. pallidulus 13271/635a TYPE Switz.
C. boulderensis var. pallidulus 13271/635b TYPE Switz. C. rubrovioleipes
C. boulderensis DQ499466 TYPE USA, WA
C. boulderensis FJ039558 Canada, BC
C. fuscovelatus NR131888 TYPE Sweden
C. pseudobovinus DQ499465 TYPE USA, WY
C. roseivelatus NR131868 TYPE Finland
C. aff. roseivelatus TN09-155 USA, WA
C. brunneocalcarius JQ746603 Finland
C. brunneocalcarius NR120096 TYPE Canada, QC
C. uraceisporus KP165544 Finland
C. uraceisporus KP165543 TYPE Finland
C. brunneifolius TN09-153 USA, WA
C. brunneifolius EU259284 TYPE Finland
C. brunneifolius TN09-153 USA, WA
C. diosmus JFA11456 USA, WA
C. diosmus 17782 TYPE France
C. urbicus TN09-011 Canada, NL
C. urbicus KP866158 TYPE Sweden
C. diosmoides 0093370 TYPE France
C. murinascens KP165572 Finland
C. murinascens NR131886 TYPE Finland
C. canabarba CFP473 Sweden
C. rusticus PK3802 TYPE Finland
C. umidicola f. coeruleus MM1991-0391 TYPE USA, WY
C. rusticus FJ039562 Canada, BC
C. pseudocalopus 2010-104 TYPE France
C. solis-occasus CFP1084 France
C. parasuillus 806 TYPE France
C. subcurtipes 02-10-89 TYPE France
C. distortus 10345 TYPE USA, WA
C. submelleopallens RH1207-2029 TYPE France
C. laniger CFP1136a Sweden
C. distortus AY669696
C. laniger KC842398
C. laniger 6029897 TYPE Finland
C. laniger f. macrosemen 2145 TYPE France
C. laniger CFP1136b Sweden
C. bidaudii 2010-100 TYPE France
C. cremeolaniger 5353 TYPE UK
C. lanigeroides 387 TYPE UK C. pearsonii
C. exsularis TN11-305 Canada, AB
C. exsularis KX234735 TYPE Spain
C. vinaceogrisescens KU041744 USA, WA
C. vinaceogrisescens
KU041743 TYPE USA, WA
sect. Furvi
Fig. 1 (continued)

Fungal Diversity
1 3
sect. Telamoni
a
100
100
98
78
99
98 100
100
96
96
98
100
100
81
87
100
100
100
98
62
60
98
100
98
100
97
99
100
100
99
100
100
90
100
98
91
C. bivelus AY669682
C. quietus RH3689 TYPE France
C. suilloclarus 4393 TYPE France
C. bivelus KP866159 TYPE Sweden
C. bivelus f. sulcatocephalus 97-10-330 TYPE France
C. bivelosimilis TN10-014 TYPE Canada, QC
C. bivelosimilis IK98-1356 Finland
C. impennoides 4971 TYPE France
C. aptecohaerens EU821657 Canada, BC
C. aptecohaerens RH71359 TYPE France
sect. Athabasci
C. serratissimus MM1960-0004 TYPE Switzerland
C. serratissimus IK00-018 Norway
C. subserratissimus KP165553 Sweden
C. subserratissimus NR131881 TYPE Sweden
sect. Sciophylli
C. cisqhale JF795387 TYPE USA, CA
C. cisqhale TN12-065 USA, CA
C. rheubarbarinus CFP848 Belgium C. phaeosmus
C. subpulchrifolius 10419 TYPE USA, MI
C. subpulchrifolius 19.07.09 Canada, NL
C. fructuodorus NR131827 TYPE USA, WA
C. fructuodorus TN12-308 USA, CA
C. tigrinipes TN04-1120 Italy
C. tigrinipes KC842400
C. tigrinipes 874 TYPE France
C. agathosmus KC608590 TYPE Sweden
C. agathosmus IK98-858 Sweden
C. agathosmus TN10-082 Canada, NL
C. agathosmus TN10-005 Canada, QC
C. leucophaeatus IK97-138 Finland
C. leucophaeatus UDB002228 Sweden
C. odoritraganus MT112154 TYPE Canada, QC
C. pulchrifolius var. odorifer 10398 TYPE USA, NC
C. niveotraganus NR131842 TYPE Finland
C. niveotraganus JFA9927 USA, WY
C. calopus CFP439 Sweden
C. venustus FJ039571 Canada, BC
C. venustus PAK3234 TYPE Finland
C. calopus PAK314 TYPE Finland
C. traganulus 4349 TYPE UK
C. fragrans 17713 TYPE USA, WA
C. ionema MM1980-0389 TYPE USA, WA
C. calopus PAK314 TYPE Finland
C. calopus FJ039572 Canada, BC
C. ionophyllus IK97-137 Finland
C. ionophyllus MM1949-0052 TYPE Austria
C. ultimiionophyllus TN07-303 TYPE Canada, NL
C. ultimiionophyllus IK95-507 Finland
C. subionophyllus TN06-050 TYPE Norway
C. ionophyllus CFP974 Sweden
C. sp. GQ159889 Canada, BC
C. torvovelatus 4145 TYPE France
C. torvus CFP778 TYPE Sweden
C. torvus AY669668
C. traganus CFP763 TYPE Sweden
C. traganus f. ochraceus JFA9578 TYPE USA, WA
C. traganus KC842399 Norway
C. venustissimus TN03-938 Sweden
C. venustissimus 5371 TYPE Sweden
sect. Tragani
sect. Phaeosmi
sect. Cisqhale
Fig. 1 (continued)

Fungal Diversity
1 3
sect. Firmiores
91
96
64
60
88
100
100
100
82
100
53
85
64
100
80
100
56
91
88
79
65
98
84
99
95
77
62
C. subumidicola 3982 TYPE France
C. pseudoprivignus RH2883 TYPE France
C. triformis f. strenuus 3773 TYPE France
C. boletiformis 1741 TYPE France
C. turgidoides 431 TYPE France
C. bucknallii RH71385 TYPE France
C. renidentoides 71560 TYPE France
C. hydrotelamonioides RH3372 TYPE France
C. castanearum 71309 TYPE France
C. macropodius
C. subcompar 56933 TYPE Hungary
C. subsedens RH324 TYPE France
C. compressus 10338 TYPE USA, TN
C. kauffmanianus TN09-085 USA, WA
C. biformis CFP665 Sweden
C. biformis CFP614 Sweden
C. kauffmanianus 10369 TYPE USA, MI
C. kauffmanianus AY669688
C. kauffmanianus FJ039574 Canada, BC
C. melitosarx F12652 TYPE Sweden
C. circinans RH908 TYPE France
C. millaresensis KU953933 TYPE Spain
C. submilvinus 000861 TYPE France
C. substriatus s. lato GQ159851 Canada, BC
C. substriatus s. lato TN07-362 USA, WA
C. violaceostriatus 1130 TYPE France
C. propinquus 5404 TYPE France
C. quarciticus CFP765 TYPE Sweden
C. substriatus TN07-426 USA, WA
C. substriatus 10427 TYPE USA, WA
C. pseudobiformis TN00-091 Finland
C. pseudobiformis 00-09-67 TYPE France
C. pseudophlegma TN10-178 Canada, QC
C. pseudophlegma RH70487 TYPE France
C. mixtus var. foetulentus 0093368 TYPE France
C. fuscoviolacens PC977 TYPE France
C. mystagogi 98102404 TYPE France
C. alboviolaceus CFP432 TYPE Sweden
C. mixtus 925 TYPE France
C. fuscoviolacens G977 TYPE France
C. griseoviolaceus 17228 TYPE USA, WA
C. alboviolaceus AF325597
C. obliquus TN14-137 USA, FL
C. obliquus TYPE USA, NY
C. geophilus var. subauroreus 4532 TYPE France
C. acutispissipes 2002-21 France
C. acutispissipes 1610 TYPE France
C. acutispissipes AY669657
C. paralbocyaneus UDB011349 Estonia
C. paralbocyaneus TN10-136 Canada, QC
C. caesioarmeniacus KP137498 TYPE Canada, NL
C. caesioarmeniacus KP137501 Finland
C. ornithopus RH2397 TYPE France
C. turgidus AY669689
C. isabellae 70728 TYPE France
C. cuteclarus 412 TYPE France
C. productus RH81-175 TYPE France
C. subadelphus RH80848 TYPE France
C. turgidus AY669663
C. turgidus CFP846 TYPE Belgium
C. albolilascens RH363 TYPE France
syn. sect. Biveli
syn. sect.
Hydrotelamonia
syn. sect. Armeniaci
syn. sect. Sericeocybe
Fig. 1 (continued)

Fungal Diversity
1 3
sect. Malachii
sect. Badiolaeves
sect. Biformes, syn. sect. Testaceofolii
sect. Subbalaustini
sect. Uracei
sect. Firmiores
99
86
100
50
99
57
100
100
88
98
99
86
100
95
95
92
61
84
100
100
67
100
98
97
81
88
100
84
77
64
100
84
100
100
99
88
87
65
C. armeniacus CFP809 TYPE Sweden
C. hydrobivelus 1213 TYPE France
C. privignus 1273 TYPE France
C. armeniacus KC842403 Norway
C. carneinatus F22032 TYPE Sweden
C. biveloides RH475 TYPE France
C. gymnopus 2451 TYPE France
C. subcarneinatus MF379637 TYPE Norway
C. subcarneinatus 6033545 Finland
C. malachius 1273 TYPE France
C. ochraceus 2138 TYPE USA, NYC. ochraceus 2138 TYPE USA, NY
C. malachius AY669681
C. malachius CFP1198 SwedenC. malachius CFP1198 Sweden
C. malachius f. crinitus 4665 TYPE France
C. suberi F16406 TYPE Sweden
C. brunneogriseus F14331 TYPE Sweden
C. quarciticus CFP765a Sweden
C. suberi EU821680 Canada, BC
C. badiolaevis TN06-301 Finland
C. badiolaevis NR131812 TYPE Sweden
C. glabrellus 10357 TYPE USA, MI
C. testaceofolius CFP583 TYPE Sweden
C. biformis TN07-422 USA, WA
C. glabrellus s. lato IK04-053 Finland
C. glabrellus s. lato IK97-756 Sweden
C. glabrellus s. lato AY669692
C. subbalaustinus EU821669 Canada, BC
C. subbalaustinus TN09-204 USA, WA
C. subbalaustinus CFP926 TYPE Sweden
C. centrirufus JV16764 Finland
C. centrirufus AY669693
C. centrirufus NR131889 TYPE Finland
C. phaeochrous 13309/162 TYPE Switz.
C. phaeochrous TN03-1068 Sweden
C. nolaneiformis KJ206491 Sweden
C. nolaneiformis NR131833 TYPE Czech
C. vernalisierraensis KX882653 USA, CA
C. vernalisierraensis KX882652 TYPE USA, CA
C. ahsii KX882644 TYPE USA, WY
C. ahsii KX882651 USA, WY
C. uraceomajalis NR131835 TYPE Hungary
C. uraceomajalis 857044 Czech Republic
C. colymbadinus KC842404 Norway
C. colymbadinus NR131819 TYPE Sweden
C. colymbadinus KC608592 USA, WA
C. subargyronotus KM576360 Austria
C. subargyronotus NR131871 TYPE Sweden
C. argyronotus 04-09-84 TYPE France
C. rigidipes KJ206506 Germany
C. rigidipes KJ206505 France
C. uraceonemoralis KJ206518 Sweden
C. uraceonemoralis NR131836 TYPE Italy
C. uraceus KC608595 USA, OR
C. uraceus NR131837 TYPE Finland
C. nodosisporus NR131870 TYPE Norway
C. rumoribrunsii KX882669 USA, CA
C. rumoribrunsii KX882668 USA, CA
C. flavobasilis KX882665 USA, MT
C. flavobasilis KX882664 TYPE USA
C. vernalishastensis KX882670 TYPE USA, CA
C. bridgei KX882661 USA, WA
C. bridgei
KX882655 TYPE USA, WA
3
3 subsect. Flavobasiles
2
2 subsect. Uracei
1 subsect. Colymbadini
1
syn. sect. Miniatopode
s
syn. sect. Colymbadini
syn. sect. Cinnabarini
syn. sect. Bulliardii
Fig. 1 (continued)

Fungal Diversity
1 3
sect. Pseudobicolores
sect. Disjungendi
sect. Hydrocybe
sect. Bicolores
sect. Uracei
98
70
98
100
100
94
74
62
92
100
98
98
75
85
94
100
100
81
99
52
100
50
90
100
90
94
89
100
100
62
100
100
96
100
100
100
99
97 100
100
C. hesleri NR131818 TYPE USA, MI
C. hesleri JQ974388 USA, TN
C. sp. FJ039678 Canada, BC
C. coracis KJ206497 France
C. coracis NR131834 TYPE Finland
C. cinnabarinus s. FN NR120163 Sweden
C. cinnabarinus s. FN AY669662 Germany
C. sp. AY669659
C. bulliardii JX114942 TYPE Sweden
C. bulliardii JF907860
C. bulliardii AF389154
C. colus var. occidentalis MM1997-0260 TYPE USA, WY
C. flammeouraceus CFP767 Sweden
C. nauseosouraceus NR131828 TYPE USA, WA
C. nauseosouraceus KC608584 USA, OR
C. pyrophyllus RH2556 TYPE France
C. duracinellus RH1232 TYPE France
C. imbutus CFP1204 Sweden
C. mattiae KX964424 TYPE Sweden
C. mattiae FJ039684 Canada, BC
C. montebelloensis KP114459 TYPE Canada, QC
C. montebelloensis KP114460 Canada, QC
C. planodepressus KP013185 TYPE France
C. claroplaniusculus NR131844 TYPE France
C. olididisjungendus KP013197 Hungary
C. olididisjungendus NR131839 TYPE Canada, ON
C. piceidisjungendus KP013206 Finland
C. piceidisjungendus NR131840 TYPE USA, WA
C. orasericeus KP013204 Norway
C. orasericeus KP013203 TYPE France
C. disjungendus KP013190 TYPE Finland
C. reumauxii KP013193 TYPE France
C. disjungendulus NR131838 TYPE Sweden
C. disjungendulus KP013187 Finland
C. pallidostriatus 6001972 Finland
C. pallidostriatus 23.09.10 Canada, QC
C. aff. FJ039566 Canada, BCpallidostriatus
C. TN09-150 USA, WAaff. pallidostriatus
C. fulminans HQ336686 Germany
C. fulminans 2993 TYPE France
C. sp. AF388778
C. duracinus KX964582 TYPE France
C. subdubius RH2527 TYPE France
C. duracinus CFP1143 Sweden
C. submutabilis 93-09-286 TYPE France
C. subduracinoides 1033 TYPE France
C. subduracinus 1044 TYPE France
C. subradicans 99103009 TYPE France
C. pseudorigens 34158/b TYPE Hungary
C. diabolicorigens 50147 TYPE Hungary
C. caesiostipitatus 4639 TYPE France
C. sp. AY669675
C. spisnii 96140 TYPE Italy
C. spisnii AY669674
C. badioflammeus 99-10-294 TYPE France
C. laceratomarginatus 2007-25 TYPE France
C. evernius KC842401 Norway
C. evernius KX964347 Canada, QC
C. evernius KX964331 TYPE Sweden
C. plumulosus KX964374 TYPE France
C. refectus KX964381 Germany
C. cagei KX964295 TYPE Sweden
C. cagei AY669676
4 subsect. Bulliardii
4
syn. sect. Duracini
Fig. 1 (continued)

Fungal Diversity
1 3
sect. Micro-ornati
sect. Cacaodisci
sect. Fuscoperonati
sect. Humicolae
sect. Craticii
sect. Tortuosi
100
100
87
62
99
92
50
88
100
96
72
100
73
77
92
58
98
100
98
100
100
100
90
55
100
88
99
75
76
100
55
66
97
78
100
100
76
100
100
92
98
100
100 C. subcagei EU597034 Canada, BC
C. subcagei KX964410 TYPE USA, CA
C. sp. FJ039565 Canada, BC
C. cystidiobicolor AF325593
C. cystidiobicolor KX964411 TYPE Finland
sect. Bicolores
C. truckeensis JQ937285 USA, CA
C. truckeensis JQ937284 TYPE USA, CA
C. contractus KX964416 TYPE France
C. cylindratus KX964414 TYPE France C. cinnamoviolaceus
C. sejunctifolius TN04-880 Finland
C. triangulus RH80869 TYPE France
C. micro-ornatus RH70678 TYPE France
C. melleopallens CFP433 TYPE Sweden
C. summomaculatus RH47.9035 TYPE France
C. cacaodiscus KP114462 TYPE Finland
C. umbilicatus PK21.8.1892 TYPE Finland
C. tortuosus KX964406 USA, WA
C. flabelloides KX964392 TYPE France
C. tortuosus AY669669
C. paranomalus KX964356 TYPE France
C. glaphurus MT15-004 UK
C. sp. GQ159774 Canada, BC
C. turgidipes KX964409 TYPE France
C. dolabratus KX964323 USA, CA
C. dolabratus KX964309 TYPE Sweden
C. dolabratus EU821665 Canada, BC
C. dolabratoides KX964304 Finland
C. dolabratoides KX964302 TYPE Finland
C. hircinosmus KX964368 TYPE France
C. hircinosmus KX964370 Sweden
C. aprinus CFP382 Sweden
C. suillonigrescens 724 TYPE France
C. implexobrunnescens 00274013 TYPE France
C. fuscoperonatus EU433390 Sweden
C. fuscoperonatus JX407330 France
C. humicola KP866157 TYPE France
C. humicola AF388785
C. scaurotraganoides RH70538 TYPE France
C. scaurotraganoides DB3117 Hungary
C. subrimosus 10424 TYPE USA, MI
C. aff. scaurotraganoides HRL0578 Canada, QC
C. subheterocyclus IK97-1306 Finland
C. subheterocyclus MF379631 TYPE Finland
C. heterocycloideus MF379632 TYPE Norway
C. heterocycloideus IK99-338 Finland
C. heterocyclus IK95-608 Finland
C. heterocyclus F14330 TYPE Sweden
C. rubicundus 3189 TYPE France
C. danicus 72581 TYPE Denmark
C. basivinosus MM1968-0147 TYPE France
C. craticius TN05-069 TYPE Finland
C. basivinosus MM1968-0147 TYPE France
C. caput-medusae HQ845170 TYPE Sweden
C. francescae 2354 TYPE France
C. fillionii TN09-043 USA, WA
C. fillionii HQ845171 TYPE France
C. subfillionii TN11-158 USA, AK
C. subfillionii MF379630 TYPE Sweden
C. boreasensis FJ039677 Canada, BC
C. boreasensis 970 TYPE USA, NY
C. ionosmus MM1967-0063 TYPE Austria
C. scriptor 388261 TYPE France
C. ionosmus CFP801 Sweden
sect. Ionosmi
sect. Pseudotragani
sect. Sordescentes
sect. Sejunctifolii
sect. Vi
liores
Fig. 1 (continued)

Fungal Diversity
1 3
sect. Bovini
sect. Niveoglobosi
sect. Brunneotincti
sect. Pholidei
sect. Privignati
sect. Armillati
100
86
79
55
100
100
92
70
54
90
100
100
100
59
100
100
77
88
100
100
95
73
90
81 100
100
100
99
94
89
100
97
65
100
100
100
75
100
55
100
100
100
100
C. raphanoides MM1974-0434 TYPE Sweden
C. raphanoides TN09-197 USA, WA
C. raphanoides CFP956 Sweden
C. panellus F129865 TYPE Sweden
C. panellus KC842407 Norway
C. pholideus CFP602 TYPE Sweden
C. pholideus KC842406Norway
C. subpholideus RH70649 TYPE France
C. subpenicillatus XC2005-18 TYPE France
C. orbiculozonarius RH70617 TYPE France
C. rheubarbarinus 1862 TYPE France
C. fuliginosus 109660 TYPE UK
C. depexus var luminosus 98101804 TYPE France
C. olivaceostipitatus 98102507 TYPE France
C. valgus MM1970-0190 TYPE Sweden
C. valgus CFP652 Sweden
C. privignipallens TN09-195 USA, WA
C. privignipallens NR131885 TYPE Finland
C. privignatus TN07-360 USA, WA
C. privignatus F148849 TYPE Sweden
C. alboambitus NR131825 TYPE USA, WA
C. alboambitus FJ717499 Canada, BC
C. luteo-ornatus NR119930 TYPE Austria
C. luteo-ornatus 12.09.01 Canada, BC
C. rossicioenochelis MF379633 TYPE Russia
C. suboenochelis NR119929 TYPE Finland
C. suboenochelis TN09-038 USA, WA
C. paragaudis NR131814 TYPE Norway
C. paragaudis JFA11514 USA, WA
C. pinigaudis HQ845162 TYPE Finland
C. pinigaudis HQ845168 Finland
C. armillatus KC842408 Norway
C. armillatus NR131891 TYPE Sweden
C. armillatus TN11-207 USA, AK
C. roseoarmillatus HQ845117 TYPE Finland
C. roseoarmillatus HQ845118 Sweden
C. alboglobosus NR131841 TYPE Finland
C. alboglobosus AY669661
C. alboglobosus KM273100 USA, WA
C. pseudorusticus 5329 TYPE France
C. niveoglobosus CFP831 TYPE Sweden
C. bovinus NR120189 TYPE Finland
C. bovinus AY669691
C. bovarius NR131830 TYPE USA, AK
C. bovarius KC905159 Canada, AB
C. oulankaensis JX407292 TYPE Finland
C. oulankaensis FJ039672 Canada, BC
C. bovinatus NR131821 TYPE Finland
C. bovinatus JX407269 Sweden
C. bovinaster JX407264 TYPE Finland
C. bovinaster JX407266 Sweden
C. fuscobovinaster JX407310 France
C. fuscobovinaster JX407316 TYPE Norway
C. fuscobovinus NR131824 TYPE Finland
C. fuscobovinus TN11-172 USA, AK
C. sp. FJ039671 Canada, BC
C. bubulus DQ139983 TYPE Austria
C. terribilis 5268 TYPE France
C. terribilis TN03-1387 Sweden
C. terribilis s. lato 4061 France
C. pseudobulbosus XC2010-101 TYPE France
1 subsect. Bovini
2 subsect. T
erribiles
2
1
sect. Alboambiti
sect. Valgi
Fig. 1 (continued)

Fungal Diversity
1 3
C. cacaocolor DW9212010 USA, WA
C. cacaocolor 17194 TYPE USA, WA
C. ectypus EU266689 TYPE Switzerland
C. ectypus EU26668 Sweden
99100
62
98
C. subglandicolor MF379639 TYPE Norway
C. subglandicolor IK99-721 Finland
C. glandicolor NR119683 TYPE Finland
C. subbrunneus f. exannulatus 2332 TYPE France
C. glandicolor AY033134
sect. Brunnei
C.brunneus DQ117927 TYPE Sweden
C.brunneus TN07-193 Canada, ON
100
100
100
100
100
100
99
100
55
100
100
100
99
100
100
70
100
100
100
100
52
87
98
100
57
52
94
50 88
100
100
100
74
69
100
94
79
C. sp. UDB018761 Estonia
C. sp. AJ889942 Denmark
C. wahkiacus KU041745 TYPE USA, WA
C. wahkiacus KU041746 USA, WA
C. sordescens IK03-006 Sweden
C. sordescens RH1687 TYPE France
C. anisochrous JX407297 TYPE Estonia
C. anisochrous JX407298 Finland
C. turgidulus 5333 TYPE France
sect. Bovini
C. diffamatus TN03-748 Sweden
C. diffamatus 99.10.13.03 TYPE France
C. eldoradoensis GQ159869 Canada, BC
C. eldoradoensis JQ906746 TYPE USA, CA
C. sordidemaculatus DQ139984 TYPE France
C. sordidemaculatus TN11-450 USA, WA
C. anthracinicolor 3881 TYPE France
C. neofurvolaesus NR131789 TYPE Sweden
C. neofurvolaesus TN09-104 USA, WA
C. anisatus NR131788 TYPE Sweden
C. anisatus EU821671 Canada, BC
C. anisatus TN11-223 USA, AK
C. neocolus TN11-285 Canada, AB
C. neocolus 3091 TYPE France
C. subbrunneoideus KP165551 Norway
C. subbrunneoideus NR131880 TYPE Finland
C. aleuriodor RH1692 TYPE France
C. umbrinoconn. var. brunnescen. 4789 TYPE Switz.
C. repertus 452421 TYPE France
C. repertus CFP1240 Sweden
C. subbulliardioides RH2476 TYPE France
C. subbulliardioides TN04-989 Sweden
C. torvoides 4342 TYPE France
C. bulbosoides CFP382 Sweden
C. bulbosoides 09-10-198 TYPE France
C. hillieri 336 TYPE France
C. hillieri IK11-019 Norway
C. clarobrunneus NR131805 TYPE Sweden
C. clarobrunneus FJ039682 Canada, BC
C. clarobrunneus EU266672 Slovakia
C. pallidibrunneus MT112157 TYPE Finland
C. clarisordidus MT112158 TYPE Finland
C. clarisordidus TN07-449 USA, WA
C. sp. FJ039680 Canada, BC
C. fibrillosibrunneus MT112159 TYPE Finland
C. brunneovernus NR131826 TYPE USA, WA
C. brunneovernus KC608580 USA, WA
C. grosmorneensis JQ746598 Canada, NL
C. grosmorneensis JQ746596 TYPE Canada, NL
C. caesiobrunneus EU266653 TYPE Finland
C. caesiobrunneus EU266652 Sweden
3 subsect. Brunnei
3
2
2 subsect. Furvolaesi
1
1 subsect. Terribiles
Fig. 1 (continued)

Fungal Diversity
1 3
Molecular results
The phylogenetic tree resulting from the analysis, ITS and
LSU regions including binary data from gap coding of the
ITS region, is shown in FIG.1 and a schematic drawing
of the relationships of the sections based on the phyloge-
netic analysis in Fig.2. Altogether, we recognize 80 sec-
tions which all form monophyletic groups in our analysis
and examples of the sections are shown in Supplementary
Figs.1–11. We almost entirely used section names with clear
identity, i.e., the concept of the type species of a section was
well known.
A total of 482 types representing 184 species were suc-
cessfully sequenced. Of these, about half of the species had
one or more synonyms. A significant number of old types
were successfully sequenced, 105 types over 50 years old
and 18 types over 100 years. All the major Cortinarius tax-
onomists have described new species that already had an
older name, but the portion of younger taxonomic synonyms
in terms of the total number of described species (synonym
rate) varies among the different authors. Here are synonym
rates for authors who have described most of the Telamo-
nia species, based only on morphological characters: Peck
17%, Kauffman 7%, Smith 40%, Moser 47%, Henry 55%,
and French Atlas team 72%. Current names of Cortinarius
species used in this study with their synonyms are listed in
Table1. All the names of the types are listed in alphabetical
order in Supplementary Table1, followed by the current
name.
Sometimes it was only possible to amplify part of the
ITS region, in most cases it was then the ITS1 region that
was succesful. Often in Cortinarius the ITS1 region alone is
enough for a proper identification, but especially in the case
of small Telamonia species several sister species can have
an identical or almost identical ITS1 region. Therefore, all
of the unclear cases are marked in Supplementary Tables1
and 2 with a prefix ‘cf.’ in the Latin name under the current
name.
Taxonomy
Neo‑ andepitypications
All older names without a type specimen that are included
in the Cortinarius subgen. Telamonia key in Funga Nordica
(Niskanen etal. 2012) and not yet typified are typified here
with the exceptions of C. paleaceus Fr. and C. miniatopus
J.E. Lange (not included in Niskanen etal. 2012) and C.
psammocephalus (Bull.) Fr. (nomen dubium, no type pro-
posed). In addition, for C. colus Fr. an epitype that differs
from the current use of the name is proposed, and in the
case of C. alboviolaceus (Pers.) Fr., C. flabellus (Fr.) Fr.
and C. hinnuleus Fr. the best fitting candidate from two or
more avalaible ones in a species group was selected. For
C. anthracinus Fr. and C. cinnabarinus Fr. neotypes have
been chosen by Høiland (1983) but were not sequenced in
this study. Altogether, neotypes for 11 species originally
described by Fries, Liljeblad and Persoon are proposed as
well as epitypes for 9 species described by Fries, Lange,
Persoon, Quélet and Schaeffer. Citations of descriptions and
illustrations of the species are provided. Notes under the
name have only been added if our typification does not fit
with the protologue and/or differs from the current use of
the name in Niskanen etal. (2012). Synonyms are based on
0.07
sect. Ochropallescentes
sect. Brunnei
84
100
100
100
100
92
99
99
100
100
64
100
100
100
100
100
100
99
71
C. ominosus FJ157107
C. ominosus FJ039598
C. birkebakii FJ039592
C. birkebakii FJ039591
C. neosanguineus FJ157116
C. neosanguineus NR120149
C. ochropallens NR131832 TYPE USA, WA
C. ochropallens FJ039673 Canada, BC
C. pardinipes KJ206494 Finland
C. pardinipes KJ206492 TYPE France
C. gentilis EU266692 TYPE Norway
C. gentilis TN09-015 Canada, NL
C. albogaudis EU266634 Finland
C. albogaudis EU266635 TYPE Finland
C. adustorimosus RH883 TYPE France
C. adustorimosus TN11-393 Canada, AB
C. coleoptera EU266684 TYPE Sweden
C. coleoptera JQ888163 UK
C. cicindela TN07-294 Canada, NL
C. cicindela NR119680 TYPE Norway
C. carabus EU266661 TYPE Finland
C. carabus EU266658 Finland
C. subcarabus EU266656 TYPE Finland
C. subcarabus EU266655 Sweden
1
1 subsect. Carabi
Dermocybe
Fig. 1 (continued)

Fungal Diversity
1 3
DNA studies of the type specimens and the information on
the types is presented in Table1.
Cortinarius alboviolaceus (Pers.) Fr., Epicr. syst. mycol.
(Upsaliae): 280 (1838) [1836–1838]
Basionym: Agaricus alboviolaceus Pers., Syn. meth.
fung. (Göttingen) 2: 286 (1801): sanctioned in Fr., Syst.
mycol. 1: 218 (1821).
Type: Sweden, Härjedalen, Storsjö sn, Flatruet, in sub-
alpine zone with Betula pubescens, 16 Aug 1986, coll. H.
Lindström etal. CFP 432, F41130 (S, neotypus hic designa-
tus, IF 557454), GenBank No. MT934857 (ITS).
Illustration. Brandrud etal. (1989: pl. A59).
Descriptions of the species. Brandrud etal. (1989: pl.
A59), Niskanen etal. (2012).
Notes—Currently, we know three species in Europe that
fit into Fries’ description of C. alboviolaceus: C. acutispis-
sipes Rob. Henry, C. alboviolaceus s. Brandrud etal. (1989)
& Niskanen etal. (2012), and C. paralbocyaneus Eyssart.
Based on the current data C. alboviolaceus s. auctores has
the widest distribution of the three species and is also the
Flexipedes
Sporagniti
Heterospori
Ochrovelati
Flexibilifolii
Hinnulei
Leiocastanei
Rubricosi
Rubrocincti
Paleacei
Incisiores
Pseudoduracini
Saniosi
Bibuli
Ravi
Alnicolarum
Helvelloides
Verni
Nucicolores
Caliginosi
Megaspori
Atroalbi
Duristipedes
Punctatiformes
Praestigiosi
Friesiorum
Vinaceobrunnei
Castanei
Parvuli
Bombycini
Anthracini
Crassispori
Squalidi
Saturnini
Politi
Furvi
Boulderenses
Brunneocalcari
Brunneifolii
Urbici
Rustici
Lanigeri
Exsulares
Athabasci
Sciophylli
Cisqhale
Phaeosmi
Telamonia
Tragani
Firmiores
Malachii
Badiolaeves
Biformes
Subbalaustini
Uracei
Pseudobicolores
Disjungendi
Hydrocybe
Bicolores
Viliores
Sejunctifolii
Micro-ornati
Cacaodisci
Tortuosi
Sordescentes
Fuscoperonati
Humicolae
Pseudotragani
Cratici
Ionosmi
Brunneotincti
Pholidei
Valgi
Privignati
Alboambiti
Armillati
Niveoglobosi
Bovini
Brunnei
Ochropallescentes
SAINOMALET LASAB
SAINOMALET NWORC
ebycimauqS/ setnecseburE/
ebycimauqsuE/ isodulaP/
88
98
100
93
96
66
77
100
100
98
78
54
70
100
100
99
90
74
100
70
100
100
100
92
71
74
88
76
100
100
100
100
100
100
100
90
99
100
78
88
84
100
100
100
100
100
97
100
100
100
100
100
100
100
100
100
100
100
100
100
84
55
88
90
75
50
88
76
92
98
55
52
73
Fig. 2 A schematic drawing of the relationships of the 80 sections of Cortinarius subgen. Telamonia accepted in this study. The drawing is based
on our phylogenetic analysis with bootstrap values greater than 50% gained in the analysis indicated above branches

Fungal Diversity
1 3
Table 1 Current names of the Cortinarius species accepted in this study with their synonyms
Current name Younger synonyms
C. aavae Liimat. & Niskanen 2012
C. acutispissipes Rob. Henry 1981 C. geophilus var. subauroreus Bidaud, Moënne-Locc. & Reumaux 2002
C. adustorimosus Rob. Henry 1988
C. ahsii McKnight 1975
C. alboadustus Bidaud 2012
C. alboambitus Niskanen, Liimat. & Ammirati 2013
C. albogaudis Kytöv., Niskanen & Liimat. 2009
C. alboglobosus Kytöv., Liimat., Niskanen & Ammirati 2014
C. albolens Bidaud, Carteret & Reumaux 2012
C. albosericeus Ammirati, Beug, Liimat., Niskanen & O. Ceska 2016
C. alboviolaceus (Pers.) Fr. 1838 C. fuscoviolascens Reumaux 1990; C. mixtus Reumaux 2002; C. mixtus
var. foetulentus Carteret, Moënne-Locc. & Reumaux 2002; C. mys-
tagogi Carteret & Reumaux 2002; C. radicatoviolaceus Rob. Henry
1981; C. griseoviolaceus A.H. Sm. 1944
C. aleuriodor Rob. Henry 1981 C. perrinii Rob. Henry & Ramm 1989; C. umbrinoconnatus var. brun-
nescentipes Bidaud, Moënne-Locc., Reumaux & Rob. Henry 2000
C. americanus A.H. Sm. 1934 C. sphagnicola Carteret & Reumaux 2004
C. ammophiloides Bohus 1979 C. ochrorufus Moënne-Locc. & Fillion 2001; C. punctatorum Moënne-
Locc. & Fillion 2012; C. salicetophilus Bidaud & Fillion 2012; C.
subfistularis Fillion 2012; C. tener Carteret 2001; C. nigromaculatus
Carteret 2012
C. andreae H. Lindstr. 1999 s. lato C. sporagnitus Bidaud 2010
C. angelesianus A.H. Sm. 1944 C. angelesianus var. elakatopus M.M. Moser 2001; C. strobilaceus
M.M. Moser 1967; C. tubulipes J. Favre 1960
C. angustisporus Kytöv., Niskanen & Liimat. 2014
C. anisatus H. Lindstr., Kytöv. & Niskanen 2005
C. anisochrous Kytöv., Liimat., Niskanen & H. Lindstr. 2013
C. anthracinicolor Reumaux 2001
C. anthracinus (Fr.) Sacc. 1887 s. Funga Nordica C. danili Rob. Henry 1943; C. ignifluus Bidaud 1994; C. josserandii
Bidaud 1994; C. procalans Moënne-Locc. & Reumaux 1994; C.
subanthracinus Rob. Henry 1944;
C. aptecohaerens Rob. Henry 1983 C. impennoides Bidaud, Moënne-Locc. & Reumaux 2000
C. argyronotus Bidaud 2008
C. armeniacus (Schaeff.) Fr. 1838 C. hydrobivelus Rob. Henry & Reumaux 2010; C. privignorum Rob.
Henry 1985; C. privignus (Fr.) Fr. 1838
C. armillatus (Fr.) Fr. 1838
C. atroalbus M.M. Moser 1993 C. atroalbus var. nigripes M.M. Moser 1993; C. ignotissimus Bidaud
2012; C. paludophilus Carteret & Reumaux 2001
C. atrocaeruleus M.M. Moser 1967 C. carminipes Moënne-Locc. & Reumaux 1988; C. erythrinellus Reu-
maux 1988; C. insignitus Reumaux 2004
C. auroripes Carteret & Reumaux 2001 C. subgracilior Bidaud & Carteret 2008
C. badioflavidus Ammirati, Beug, Niskanen, Liimat. & Bojantchev
2016
C. badiolaevis Niskanen, Liimat., Mahiques, Ballarà & Kytöv. 2011
C. badiovestitus M.M. Moser 1968
C. bibulus Quél. 1881
C. biformis Fr. 1838 C. testaceofolius H. Lindstr. & Soop 1995
C. bistreoides Kauffman 1921
C. biveloides Rob. Henry 1948 C. carneinatus Soop 2002; C. gymnopus Rob. Henry 1983
C. bivelosimilis Kytöv., Niskanen & Liimat. 2017
C. bivelus (Fr.) Fr. 1838 C. amphibalaustius Rob. Henry 1983; C. bivelus f. sulcatocephalus
Bidaud 2010; C. insolitus Rob. Henry 1981; C. quietus Rob. Henry
1977; C. suilloclarus Reumaux 2002

Fungal Diversity
1 3
Table 1 (continued)
Current name Younger synonyms
C. bombycinus Mahiques & Burguete 2001
C. bonamei Rob. Henry 1970
C. boreasensis A.H. Sm. 1944 C. ionosmus M.M. Moser, Nespiak & Schwöbel 1969; C. scriptor
Kühner 1989
C. boreotrichus Kytöv., Niskanen & Liimat. 2017
C. boulderensis A.H. Sm. 1944
C. bovarius Liimat. & Niskanen 2013
C. bovinaster Niskanen, Kytöv. & Liimat. 2013
C. bovinatus Kytöv., Liimat., Niskanen & H. Lindstr. 2013
C. bovinus Fr. 1838
C. bridgei Ammirati, Niskanen, Liimat., Bojantchev, & Fang 2017
C. brunneifolius Kytöv., Niskanen & Liimat. 2008
C. brunneocalcarius Niskanen, Liimat. & Kytöv. 2012
C. brunneoclarus Niskanen, Kytöv. & Liimat. 2020
C. brunneovernus Niskanen, Liimat. & Ammirati 2013
C. brunneus (Pers.) Fr. 1838 C. brunneus var. incommixtus Bidaud 2009
C. bubulus Liimat., Kytöv. & Niskanen 2020
C. bulbosovolvatus Rob. Henry & Contu 1985
C. bulliardii (Pers.) Fr. 1838 C. colus Fr. 1838; C. georgianae Moënne-Locc. 1994
C. cacaocolor A.H. Sm. 1944
C. caesioarmeniacus Kytöv., Niskanen & Liimat. 2014
C. caesiobrunneus Kytöv., Niskanen & Liimat. 2009
C. cagei Melot 1990
C. californicus A.H. Sm. 1939
C. caliginosus Bidaud, Moënne-Locc. & Reumaux 2000
C. calopus P. Karst. 1881 C. fragrans A.H. Sm. 1944; C. ionema M.M. Moser & Ammirati 1996
C. campester Reumaux 2001 C. malefidus Moënne-Locc. 2001; C. porphyreticus Moënne-Locc. 2001
C. canaliculatus Bidaud & Carteret 2008
C. carabus Kytöv., Niskanen & Liimat. 2009
C. carbunculus H. Lindstr. & H. Markl. 2010
C. castaneopallidus Carteret 2004 C. franceschinii Consiglio, D. Antonini & M. Antonini 2006; C.
olivaceomarginatus Carteret 2012; C. pseudobavaricus Bidaud &
Reumaux 2012
C. castaneus (Bull.) Fr. 1838 C. fallaciosus Bidaud 2001; C. kunicensis var. caespitosus Moënne-
Locc. 2001; C. robertii Moënne-Locc. & Reumaux 1988; C. subodo-
ratus Bidaud 2001
C. centrirufus Kytöv., Niskanen & Liimat. 2014
C. chevassutii Rob. Henry 1982 C. subsordescens Rob. Henry 1985
C. cicindela Kytöv., Niskanen & Liimat. 2009
C. cinereobrunneolus Chevassut & Rob. Henry 1982
C. cinnamoviolaceus M.M. Moser 1968
C. circinans Rob. Henry 1985 C. melitosarx Soop 1999
C. cisqhale Bojantchev 2013
C. cistopulchripes Bidaud 2004
C. clarobrunneus (H. Lindstr. & Melot) Niskanen, Kytöv. & Liimat.
2009
C. claroplaniusculus Rob. Henry 1983
C. clarosordidus Niskanen, Kytöv. & Liimat. 2020
C. coccineus Reumaux 1994
C. coleoptera H. Lindstr. & Soop 1999
C. colorius (Bidaud) Niskanen, Dima & Liimat. 2020

Fungal Diversity
1 3
Table 1 (continued)
Current name Younger synonyms
C. colymbadinus Fr. 1838
C. compressus A.H. Sm. 1944
C. comptulus M.M. Moser 1968 C. fistularioides Reumaux, Bidaud & Fillion 2001; C. griseosulcatus
Carteret 2004; C. hemitrichus var. americanus A.H. Sm. 1944; C.
inolens var. parvinolens Bidaud & Carteret 2010; C. jacobii Bidaud,
Moënne-Locc. & Reumaux 2001; C. laniatus Rob. Henry 1983; C.
sublatisporus Svrček 1968
C. confirmatus Rob. Henry 1983
C. coracis Kytöv., Niskanen, Liimat. & Dima 2014
C. corvinus Reumaux 2012
C. crassisporus Kytöv., Niskanen & Liimat. 2014
C. craticius Fr. 1838 C. basivinosus M.M. Moser 1996
C. cucumisporus M.M. Moser 1967
C. cystidiobicolor Liimat. & Niskanen 2017
C. danicus Høil. 1983 C. rubicundus Bidaud, Moënne-Locc. & Reumaux 1994
C. deceptivissimus Reumaux 1984 C. damascenoides Bidaud 2008
C. decipiens (Pers.) Fr. 1838 C. anthracinoides Rob. Henry 2001; C. argumentosus Moënne-Locc.
& Reumaux 2000; C. decipiens f. saliceticola Reumaux & Carteret
2001; C. maculatophyllus Bidaud 2012; C. washingtonensis A.H. Sm.
1939
C. decipientoides Moënne-Locc. & Reumaux 1988 C. alutaceogrisescens Bidaud 1997; C. derelictus Reumaux 2001; C.
rubellopes var. pudoricolor Bidaud, Moënne-Locc. & Reumaux 2001
C. denigratus Ammirati, Beug, Niskanen, Liimat. & O. Ceska 2016
C. desertorum (Velen.) G. Garnier 1991 C. ammophilus A. Pearson 1946; C. diasemospermus var. leptosper-
mus H. Lindstr. 1998; C. difficillimus Carteret 2012; C. goniosporus
Carteret 2004; C. pertristis J. Favre 1955; C. subdepressus Carteret
2012; C. subrigidus Bidaud, Carteret & Reumaux 2010; C. friesianus
Carteret & Reumaux 2001
C. diabolicorigens Bohus 1976 C. caesiostipitatus Reumaux 2008; C. pseudorigens Bohus 1976; C.
subradicans Carteret & Reumaux 2008
C. diffamatus Carteret 2012
C. diffractosuavis Chevassut & Rob. Henry 1978
C. diosmoides Rob. Henry 1989 C. diosmoides Rob. Henry 1989
C. diosmus Kühner 1955
C. disjungendulus Kytöv., Liimat. & Niskanen 2014
C. disjungendus P. Karst. 1893 C. reumauxii Rob. Henry 1996
C. distortus Kauffman 1932 C. parasuillus Reumaux 2010; C. pseudocalopus Reumaux 2010; C.
subcurtipes Bidaud 2010; C. submelleopallens Rob. Henry 1981
C. dolabratoides Kytöv., Carteret, Bidaud, Liimat., Niskanen, Bel-
langer, Dima, Reumaux & Ammirati 2017
C. dolabratus Fr. 1838
C. duboisensis Ammirati, Beug, Niskanen & Liimat. 2016
C. dumetorum J. Favre 1960
C. duracinellus Rob. Henry 1970 C. pyrophyllus Rob. Henry 1970
C. duracinus Fr. 1838 C. subdubius Rob. Henry 1970; C. subduracinoides Moënne-Locc. &
Reumaux 2008; C. subduracinus Bidaud & Moënne-Locc. 2008; C.
submutabilis Bidaud & Carteret 2008
C. duristipes Kytöv., Niskanen & Liimat. 2014
C. ectypus J. Favre 1960
C. elaphinicolor Carteret 2004
C. eldoradoensis Bojantchev 2013

Fungal Diversity
1 3
Table 1 (continued)
Current name Younger synonyms
C. epipurrus Chevassut & Rob. Henry 1978 C. hinnuleoradicatus Bidaud, Moënne-Locc. & Reumaux 1997; C.
pallescens Moënne-Locc. & Reumaux 1997; C. pseudosafranopes
Moënne-Locc. & Reumaux 1997; C. subgrisescens Bidaud, Moënne-
Locc. & Reumaux 1997
C. evernius (Fr.) Fr. 1838
C. exitiosus Bidaud, Moënne-Locc. & Reumaux 2001
C. expallens M.M. Moser 1993 C. mucronatus M.M. Moser & McKnight 1987, nom. illegit.; C. hemi-
trichus f. improcerus J. Favre 1955; C. gossypinus H. Lindstr. 2010
C. exsularis Garrido-Ben., Ballarà & Mahiques 2016
C. fagetorum M.M. Moser 1967 C. fagetorum M.M. Moser 1967
C. falsosus Moënne-Locc. & Reumaux 2001 C. griseophyllus Reumaux 2001; C. griseovioleipes Moënne-Locc. &
Reumaux 2001; C. recedens Bidaud, Moënne-Locc. & Reumaux 2001
C. famatus Moënne-Locc. & Reumaux 2001 C. biformis var. dilatus Bidaud, Carteret & Reumaux 2012
C. ferrugineovelatus Kytöv. Liimat. & Niskanen 2014
C. fibrillosobrunneus Kytöv., Niskanen & Liimat. 2020
C. fillionii Bidaud, Moënne-Locc. & Reumaux 1995
C. flabellus (Fr.) Fr. 1838 C. flabelliformis Bidaud 2010; C. flabellus f. iners Bidaud 2010; C.
furfuraceus Bidaud 1997; C. fuscoruber Reumaux 2001; C. gurdus
Carteret 2012; C. pseudodepressus Carteret & Reumaux 2010
C. flammeouraceus Niskanen, Kytöv., Liimat., Dima & Ammirati 2020 C. colus var. occidentalis M.M. Moser 2002
C. flavobasilis Peintner, Kuhnert-Finkernagel, Cripps & Ammirati
2017
C. flexibilifolius Carteret 2004 C. olivaceobrunneus Reumaux 2012; C. privus Reumaux 2004; C.
subfuscodiscus Reumaux 2004
C. flexipes (Pers.) Fr. 1838 C. paleifer Svrček 1968
C. fragrantissimus Ammirati, Beug, Liimat., Niskanen & O. Ceska
2016
C. francescae Reumaux 1992 C. caput-medusae H. Lindstr. 1998
C. fructuodorus Niskanen, Liimat. & Ammirati 2013
C. fulminans Moënne-Locc. & Reumaux 2008
C. fulvoisabellinus Rob. Henry 1941
C. fulvopaludosus Kytöv., Niskanen & Liimat. 2017
C. furfurellus Peck 1880
C. furvoumbrinus Liimat., Niskanen & Kytöv. 2014
C. furvus Liimat., Niskanen & Kytöv. 2014
C. fuscescens Kytöv., Niskanen & Liimat. 2014
C. fuscoalbus Kytöv., Niskanen & Liimat. 2017
C. fuscobovinaster Kytöv., Liimat., Niskanen & H. Lindstr. 2013
C. fuscobovinus Kytöv., Niskanen & Liimat. 2013
C. fuscodiscus A.H. Sm. 1944
C. fuscoflexipes M.M. Moser & McKnight 1987
C. fuscogracilescens A. Favre 2009
C. fuscoperonatus Kühner 1953
C. fuscoumbrinus Liimat., Niskanen & Kytöv. 2014
C. fuscovelatus Kytöv., Niskanen & Liimat. 2014
C. gallurae D. Antonini, M. Antonini & Consiglio 2005
C. gentilis (Fr.) Fr. 1838
C. gentilissimus A.H. Sm. 1939
C. geraniolens Bidaud 2010 s. lato C. subcarcharias Bidaud 2010
C. glabrellus Kauffman 1907 C. leiopus Rob. Henry 1981
C. glandicolor (Fr.) Fr. 1838 C. subbrunneus f. exannulatus Moënne-Locc. 2009
C. glaphurus Chevassut & Rob. Henry 1982 C. paranomalus Rob. Henry 1992

Fungal Diversity
1 3
Table 1 (continued)
Current name Younger synonyms
C. griseocarneus Carteret 2010 C. atripes Reumaux 2012
C. grosmorneënsis Liimat. & Niskanen 2012
C. gualalaensis Bojantchev 2013
C. helodes M.M. Moser, Matheny & Daniele 2001
C. helvelloides (Bull.) Fr. 1838
C. hemitrichoides Bidaud & Moënne-Locc. 2010
C. hemitrichus (Pers.) Fr. 1838 C. fusisemen Reumaux 2000; C. milvinoides Carteret & Reumaux 2010;
C. paleaceus Fr. 1838; C. roseohemitrichus Carteret & Reumaux 2010
C. heparinus Kytöv., Niskanen & Liimat. 2020
C. hesleri Ammirati, Niskanen, Liimat. & Matheny 2013
C. heterocycloideus Kytöv., Niskanen & Liimat. 2017
C. heterocyclus Soop 1990
C. heterodepressus Kytöv., Niskanen & Liimat. 2017 C. bohemicus f. subheterosporus Bidaud 2010
C. heterosporus Bres. 1889
C. hillieri Rob. Henry 1938
C. hinnuleoarmillatus Reumaux 1989 C. aureifer Reumaux 1989
C. hinnuleocervinus Niskanen, Liimat. & Ammirati 2017 C. distans var. olympianus A.H. Sm. 1944
C. hinnuleus Fr. 1838 C. tigris Bidaud 1997
C. hircinosmus Moënne-Locc. 2002
C. hirtus (Velen.) G. Garnier 1991 C. querculus Moënne-Locc. & Reumaux 2001; C. punctatoides Reu-
maux 2012; Telamonia sanguinescens Velen. 1939; C. similigenus
Moënne-Locc. & Reumaux 2001; C. subargyropus Bidaud, Moënne-
Locc. & Reumaux 2001; C. substemmatus Moënne-Locc. & Reumaux
2001
C. humicola (Quél.) Maire 1911
C. hydrotelamonioides Rob. Henry 1970 C. boletiformis Bidaud & Reumaux 2010; C. bucknallii Reumaux 2010;
C. castanearum Rob. Henry 1981; C. macropodius Rob. Henry 1961
Nom. inval.; C. pseudoprivignus Rob. Henry 1985; C. renidentoides
Rob. Henry 1981; C. subumidicola Bidaud, Moënne-Locc. & Reu-
maux 2000; C. triformis f. strenuus Rob. Henry 1981; C. turgidoides
Rob. Henry 1981
C. imbutus Fr. 1838
C. impolitus Kauffman 1918 C. subacutus A.H. Sm. 1944
C. incisior Bidaud, Moënne-Locc. & Reumaux 1997 C. perzonatus Reumaux 2004; C. spurcatus Moënne-Locc. & Reumaux
2001
C. inconspicuus J. Favre 1955 C. erubescens M.M. Moser 1968
C. iners (Bidaud) Liimat., Dima & Niskanen 2020
C. intempestivus Moënne-Locc. & Reumaux 2001 C. cristatosporus Reumaux 2004
C. ionophyllus M.M. Moser 1968
C. jacobi-langei Bidaud 2008
C. kauffmanianus A.H. Sm. 1933
C. lacustris Moënne-Locc. & Reumaux 1997 C. altae-herbae Moënne-Locc. & Reumaux 1999
C. laniger Fr. 1838 C. laniger f. macrosemen Bidaud, Carteret & Reumaux 2010
C. leiocastaneus Niskanen, Liimat. & Soop 2008
C. leucophaeatus Rob. Henry 1985
C. lindstroemii Niskanen, Kytöv. & Liimat. 2020 C. flabellus f. biolens Bidaud 2010
C. lucorum (Fr.) Berger 1846

Fungal Diversity
1 3
Table 1 (continued)
Current name Younger synonyms
C. luridis Rob. Henry 1969 C. aciculisporus Moënne-Locc. 1997; C. armillifer Moënne-Locc.
& Reumaux 1997; C. aspilus Moënne-Locc. 1997; C. collybioides
Reumaux 1997; C. conicoides Bidaud 1997; C. herculinus Reumaux
1997; C. hinnuleoscitus Ramm & Rob. Henry 1995; C. hinnuleus
f. parincisus Bidaud, Moënne-Locc. & Reumaux 1997; C. lepidus
Moënne-Locc. 1997; C. ochraceoplicatus Reumaux 1997; C. pseu-
dohinnuleus Bidaud, Moënne-Locc. & Reumaux 1997; C. salicum
Reumaux 1997
C. luteo-ornatus (M.M. Moser) Bidaud, Moënne-Locc. & Reumaux
1995
C. malachius (Fr.) Fr. 1838 C. cinereoviolascens Moënne-Locc. & Reumaux 1988; C. malachius f.
cholagogus Bidaud, Moënne-Locc. & Reumaux 2002; C. malachius f.
crinitus Bidaud & Reumaux 2002; C. ochraceus Peck 1872
C. mallaensis Kytöv., Niskanen & Liimat. 2020
C. mattiae Soop 2010
C. megacystidiosus Reumaux 2012
C. melleopallens (Fr.) Britzelm. 1892 C. micro-ornatus Chevassut & Rob. Henry 1978; C. summomaculatus
Rob. Henry 1995
C. milvinicolor Moënne-Locc. & Reumaux 1997
C. miniatopus J.E. Lange 1940
C. minusculus Liimat. & Niskanen 2019
C. miwok Bojantchev 2013
C. montebelloensis Niskanen & Liimat. 2014
C. mucicola A.H. Sm. 1944
C. murinascens Kytöv., Niskanen & Liimat. 2014
C. nauseosouraceus Niskanen, Liimat. & Ammirati 2013
C. neocolus Reumaux & Sasia 2011
C. neofallax Carteret & Reumaux 2004 C. incisopunctatus Reumaux 2012
C. neofurvolaesus Kytöv., Niskanen, Liimat. & H. Lindstr. 2005
C. nigrellus Peck 1873
C. nigrocuspidatus Kauffman 1921 C. adalbertii var. turritus M.M. Moser 1980; C. inolens (H. Lindstr.)
Bidaud 2010; C. striaepilus J. Favre 1948
C. niveoglobosus H. Lindstr. 1992 C. pseudorusticus Bidaud 2002
C. niveotraganus Kytöv., Niskanen & Liimat. 2014
C. nodosisporus Kytöv., Niskanen & Liimat. 2014
C. nolaneiformis (Velen.) Dima, Niskanen & Liimat. 2014
C. nucicolor Liimat., Niskanen & Kytöv. 2014
C. obliquus Peck 1902
C. ochropallens Liimat., Niskanen & Ammirati 2013
C. ohlone Bojantchev 2013
C. olididisjungendus Liimat., Niskanen, Dima & Kytöv. 2014
C. olivaceofulvus Kauffman & A.H. Sm. 1933 C. lacorum A.H. Sm. 1934
C. orasericeus Rob. Henry 1983
C. oulankaënsis Kytöv., Niskanen, Liimat. & H. Lindstr. 2013
C. pallidostriatus Rob. Henry 1968
C. paludosaniosus Liimat., Niskanen, Dima & Ammirati 2017 C. saniosus var. paludophilus Carteret & Reumaux 2012
C. panellus Soop 2009
C. pangloius M.M. Moser 1969 C. ferrugineifolius M.M. Moser 1993; C. paraphaeochrous M.M. Moser
1993; C. subrigidipes M.M. Moser 1993
C. paragaudis Fr. 1838
C. paralbocyaneus Eyssart. 2002
C. pardinipes Romagn. 1977

Fungal Diversity
1 3
Table 1 (continued)
Current name Younger synonyms
C. parhonestus Reumaux 2012
C. pearsonii P.D. Orton 1958 C. cremeolaniger P.D. Orton 1983; C. lanigeroides P.D. Orton 1983
C. pelargoniostriatulus Bidaud & Fillion 2010
C. phaeochrous J. Favre 1955
C. phaeosmus Rob. Henry 1981
C. pholideus (Lilj.) Fr. 1838 C. pholideoides Bidaud & Reumaux 2005; C. subpenicillatus Carteret
& Reumaux 2005; C. subpholideus Rob. Henry 1992
C. piceidisjungendus Kytöv., Liimat., Niskanen & Ammirati 2014
C. pilatii Svrček 1968 C. altipes Bidaud 2010; C. conocyboides Carteret 2004; C. distinctus
Carteret 2012; C. pelargoniobtusus Rob. Henry 1985; C. pseudo-
rigidus Bidaud, Carteret & Reumaux 2012; C. udolivascens var.
lilacinostipitatus Carteret 2004
C. pinigaudis Niskanen, Kytöv. & Liimat. 2011
C. pinosquamulosus Kytöv., Niskanen & Liimat. 2020
C. plumulosus Rob. Henry 1977
C. politus Niskanen, Liimat. & Ammirati 2013
C. praepallens Peck 1887
C. praestigiosus (Fr.) M.M. Moser 1965 C. fulguritans Reumaux 2000; C. magus Moënne-Locc. 2001; C.
poirieri Reumaux 1988; C. rufescentipes Bidaud 2001; C. sensibilis
Bidaud 2010
C. privignatus Soop 2010
C. privignipallens Kytöv., Niskanen & Liimat. 2014
C. psammocola Kytöv., Niskanen & Liimat. 2017
C. pseudobiformis Bidaud & Carteret 2012
C. pseudobovinus M.M. Moser & Ammirati 1995
C. pseudobulbosus Carteret & Reumaux 2010
C. pseudofallax Carteret 2004
C. pseudoflabellus Bidaud 2010
C. pseudofusisporus Bidaud 2010 C. flavoperonatus Bidaud & Reumaux 2012
C. pseudophlegma Rob. Henry 1981
C. puellaris Brandrud, Bendiksen & Dima 2015
C. punctatiformis Carteret 2012
C. quarciticus H. Lindstr. 1994 C. propinquus Eyssart. & Bidaud 2002; C. violaceostriatus Moënne-
Locc. & Reumaux 2002
C. quercoconicus Liimat., Kytöv. & Niskanen 2017
C. radicosissimus Moënne-Locc. 1997 C. speciosior Bidaud, Moënne-Locc. & Reumaux 1997; C. squamulifer
Bidaud & Reumaux 1997; C. subhelvolus Moënne-Locc. & Reumaux
1997
C. raphanoides (Pers.) Fr. 1838
C. repertus A. Favre & Vialard 2004
C. rigidipes M.M. Moser 1967
C. roseivelatus Kytöv., Liimat. & Niskanen 2014
C. roseoarmillatus Niskanen, Kytöv. & Liimat. 2011
C. roseobasilis Ammirati, Beug, Liimat., Niskanen & O. Ceska 2016
C. roseobrunneus Carteret 2000
C. roseocastaneus Niskanen, Liimat. & Kytöv. 2014
C. roseomyceliosus Bidaud 2009
C. roseonudipes Rob. Henry & Moënne-Locc. 1997 C. buxiolens Bidaud 1997; C. carcharias Bidaud 1997; C. hinnuleovela-
tus Reumaux 1997; C. subfilamentosus Reumaux 1997
C. rossicioenochelis Liimat., Kytöv. & Niskanen 2017
C. rubipes Kauffman 1909

Fungal Diversity
1 3
Table 1 (continued)
Current name Younger synonyms
C. rubricosus (Fr.) Fr. 1838 C. calcareophilus Bidaud 1997; C. crassogriseascens A. Favre 2009; C.
phaeomaculatus Rob. Henry 1989; C. safranopes Rob. Henry 1938;
C. safranopes var. bulbosus Rob. Henry 1997; C. safranopes var.
laevipes Reumaux 1997
C. rubrocinctus Reumaux 1995 C. uraceoarmillatus Bidaud 2012
C. rubrovioleipes Bendiksen & K. Bendiksen 1991 C. boulderensis var. pallidulus J. Favre 1960
C. rumoribrunsii Bojantchev, Ammirati, Niskanen, & Liimat. 2017
C. russulaespermus Carteret 2004 C. striatulorufus Moënne-Locc. 2004
C. rusticellus J. Favre 1955
C. rusticus P. Karst. 1882 C. canabarba M.M. Moser 1966; C. umidicola f. coeruleus M.M.
Moser & Ammirati 1995
C. sagacitas Kytöv., Niskanen & Liimat. 2020
C. sagacito-occidentalis Liimat., Niskanen, Kytöv. & Ammirati 2020
C. saniosus (Fr.) Fr. 1838 C. bavaricus M.M. Moser 1983; C. luteolateritius (Velen.) G. Garnier
1991; C. rufoanuliferus M.M. Moser & McKnight 1987; C. subauran-
tiomarginatus Bidaud & Ferville 2012
C. saturninus (Fr.) Fr. 1838
C. scaurotraganoides Rob. Henry 1986
C. scotoides J. Favre 1955 C. castaneoruber Bidaud & Reumaux 2012; C. sublucorum Carteret
2012
C. sejunctifolius Rob. Henry 1995 C. raphanicus Bidaud & Moënne-Locc. 2008
C. semiodoratus Rob. Henry 1993 C. griseascens Bidaud, Moënne-Locc. & Reumaux 1997; C. immacula-
tus Bidaud 1997; C. nauseosmus Bidaud, Moënne-Locc. & Reumaux
1997; C. subulatus Bidaud, Moënne-Locc. & Reumaux 1997; C.
solidus Bidaud, Moënne-Locc. & Reumaux 1997
C. semivelatus Rob. Henry 1970 C. sefendens Rob. Henry 1983
C. semivestitus M.M. Moser 1968 C. fusisporus var. olivaceodepressus Reumaux 2010
C. serratissimus M.M. Moser 1968
C. sociatus Rob. Henry 1983 C. terribilis Reumaux 2002
C. sordescens Rob. Henry 1944 C. sordescens var. vestitissimus Eyssart. 2002; C. strenuipes var. sub-
acuminatus Reumaux 2002
C. sordidemaculatus Rob. Henry 1981
C. sphagnoravus Liimat., Kytöv., Niskanen & Ammirati 2017
C. spisnii Consiglio, D. Antonini & M. Antonini 2004 C. badioflammeus Bidaud 2008; C. laceratomarginatus Carteret &
Reumaux 2008
C. squalidus A.H. Sm. 1942
C. stipitemirus Rob. Henry 1995
C. stuntzii S.A. Rehner & Ammirati 1989
C. subargyronotus Niskanen, Liimat. & Kytöv. 2014
C. subbalaustinus Rob. Henry 1991
C. subbrunneoideus Kytöv., Liimat. & Niskanen 2014
C. subbulliardioides Rob. Henry 1970
C. subcagei Niskanen & Liimat. 2017
C. subcarabus Liimat., Kytöv. & Niskanen 2017
C. subcarneinatus Niskanen, Kytöv. & Liimat. 2017
C. subcastaneus Bidaud & Reumaux 2000 C. castaneus var. nigrescens Reumaux 1989; C. tenebrosus Reumaux
2001
C. subcoronatus Bidaud 2001 C. urdaibaiensis Fernández Sas. 2003
C. suberi Soop 1990 C. brunneogriseus Soop 1993
C. subexitiosus Liimat., Niskanen, Kytöv. & Ammirati 2014
C. subfillionii Kytöv., Niskanen & Liimat. 2017
C. subglandicolor Niskanen, Liimat. & Kytöv. 2017

Fungal Diversity
1 3
Table 1 (continued)
Current name Younger synonyms
C. subheterocyclus Liimat., Niskanen & Kytöv. 2017
C. subionophyllus Niskanen, Liimat. & Kytöv. 2017
C. submilvinus Bidaud 2010
C. subminiatopus Kytöv., Niskanen & Liimat. 2017 C. miniatopus var. konradii M.M. Moser 1965
C. subobtusobrunneus Bidaud 2004
C. subobtusus Kauffman & A.H. Sm. 1933
C. suboenochelis Kytöv., Liimat. & Niskanen 2011
C. subpaleaceus Kytöv., Niskanen & Liimat. 2017
C. subparvannulatus Moënne-Locc. & Fillion 2010
C. subpulchrifolius Kauffman 1918
C. subrigens Kauffman 1918
C. subrimosus A.H. Sm. & Hesler1944
C. subscotoides Niskanen & Liimat. 2019
C. subsedens Rob. Henry 1956 C. subcompar Bohus 1979
C. subserratissimus Kytöv., Liimat. & Niskanen 2014
C. substriatus Kauffman 1932
C. subtabularis Kauffman 1918 C. fumosifolius A.H. Sm. 1942
C. subtilior J. Favre 1955 C. parinsignis Moënne-Locc. & Carteret 2001; C. percavus J. Favre
1955
C. suillonigrescens Reumaux 2002 C. implexobrunnescens A. Favre 2009
C. tatrensis R. Fellner & Landa 1993
C. tenebricus J. Favre 1955
C. tigrinipes Bergeron 1997
C. tortuosus (Fr.) Fr. 1838
C. torvoides Rob. Henry 2000
C. torvus (Fr.) Fr. 1838 C. bidiscendus Rob. Henry 1985; C. subamethysteus Rob. Henry 2000;
C. testaceofractus Carteret & Reumaux 2000; C. torvovelatus Reu-
maux 2000
C. tragano-odorus Niskanen, Liimat. & Ammirati 2020 C. pulchrifolius var. odorifer Hesler 1944
C. traganus (Fr.) Fr. 1838 C. pyriodorus Kauffman 1932; C. traganus f. ochraceus M.M. Moser,
Ammirati & M.T. Seidl 1995
C. triangulus Rob. Henry 1983
C. truckeensis Bojantchev 2013
C. tuolumnensis Bojantchev 2013
C. turgidipes Bidaud & Carteret 2008
C. turgidulus Bidaud 2002
C. turgidus Fr. 1838 C. albolilascens Rob. Henry 1988; C. cuteclarus Bidaud, Moënne-Locc.
& Reumaux 2008; C. isabellae Rob. Henry 1981; C. ornithopus Rob.
Henry 1970; C. productus Chevassut & Rob. Henry 1988; C. subadel-
phus Rob. Henry 1981
C. ultimiionophyllus Kytöv., Niskanen & Liimat. 2017
C. umbilicatus P. Karst. 1893 C. cacaodiscus Liimat., Niskanen & Kytöv. 2014
C. umbrinobellus Liimat., Niskanen & Kytöv. 2014
C. umbrinolens P.D. Orton 1980 C. sericeofibrillosus Bidaud & Boutev. 2001
C. umbrinolutescens Reumaux 2004
C. uraceisporus Niskanen, Kytöv. & Liimat. 2014
C. uraceomajalis Dima, Liimat., Niskanen & Bojantchev 2014
C. uraceonemoralis Niskanen, Liimat., Dima, Kytöv., Bojantchev &
H. Lindstr. 2014
C. uraceus Fr. 1838 Hydrocybe praecox Velen. 1939
C. urbicus (Fr.) Fr. 1838 C. alsomatii Rob. Henry 1992

Fungal Diversity
1 3
most common one at least in the Nordic countries. There-
fore, we choose to follow the current Nordic concept of the
species and propose the collection H. Lindström etal. CFP
432 as the neotype of the species.
Cortinarius armeniacus (Schaeff.) Fr., Epicr. syst. mycol.
(Upsaliae): 304 (1838) [1836–1838]
Basionym: Agaricus armeniacus Schaeff., Fung. bavar.
palat. nasc. (Ratisbonae) 4: 35 (1774): sanctioned in Fr.,
Syst. mycol. 1: 234 (1821).
= Cortinarius privignus (Fr.) Fr., Epicr. syst. mycol.
(Upsaliae): 304 (1838) [1836–1838]
Types: Schaeff., Fung. bavar. palat. nasc. (Ratisbonae)
1–2: Tab LXXXI, 1774 (lectotypus hic designatus, IF
557455) as Agaricus armeniacus). Sweden, Ångermanland;
Häggdånger sn, Torrom, in spruce forest with blueberry, 26
Sep 1988, coll. H. Lindström etal. CFP 809, F37506 (S, epi-
typus hic designatus, IF 557456), GenBank No. DQ117925
(ITS).
Illustration. Brandrud etal. (1989: pl. A46).
Descriptions of the species. Brandrud etal. (1989: pl.
A46), Niskanen etal. (2012).
Cortinarius bibulus Quél., Compt. Rend. Assoc. Franç.
Avancem. Sci. 9: 666 (1881) [1880]
Types: Quélet, Compt. Rend. Assoc. Franç. Avancem.
Sci. 9: 666, Pl. VIII, fig.7, 1881 (lectotypus hic designa-
tus, IF 557457). Finland, Kainuu, Puolanka, Pihlajavaara
S., old, mossy, mesic grass-herb spruce forest (Picea abies)
with some Betula, Pinus sylvestris and Populus tremula,
240–270 m, 15 Sept 2005, coll. K. Liimatainen & T. Nis-
kanen 05-119, H6031525 (H, epitypus hic designatus, IF
557458, K, isoneotypus), GenBank No. MT934904 (ITS).
Illustration: Brandrud etal. (1992: pl. B25).
Descriptions of the species: Brandrud etal. (1992: pl.
B25), Niskanen etal. (2012) as C. lilacinopusillus P.D.
Orton.
Cortinarius bulliardii (Pers.) Fr. [as ‘bulliardi’], Epicr. syst.
mycol. (Upsaliae): 282 (1838) [1836–1838]
Basionym: Agaricus bulliardii Pers. [as ‘bulliardi’],
Observ. mycol. (Lipsiae) 2: 43 (1800) [1799]: sanctioned in
Fr., Syst. mycol. 1: 221 (1821).
= Cortinarius colus Fr., Epicr. syst. mycol. (Upsaliae):
308 (1838) [1836–1838]
Types: Bulliard, Herbier de la France: pl. 431 Fig.3, 1780
(lectotypus hic designatus, IF 557459, as Agaricus araneo-
sus). Sweden, Västergötland, Österplana sn, Österplana hed,
decideous forest on calcareus soil (Corylus, Quercus, Tilia),
15 Sep 1986, coll. H. Lindström etal. CFP 499, F41127
(S, epitypus hic designatus, IF 557460), GenBank No.
JX114942 (ITS).
Illustration. Brandrud etal. (1989: pl. A37).
Descriptions of the species. Brandrud etal. (1989: pl.
A37), Niskanen etal. (2012).
Cortinarius colus Fr., Epicr. syst. mycol. (Upsaliae): 308
(1838) [1836–1838]
= Cortinarius bulliardii (Pers.) Fr., Epicr. syst. mycol.
(Upsaliae): 282 (1838) [1836–1838]
Types: Paulet, Traité des Champignons: t. 99 spec. solita
majus 1793–1835 (lectotypus hic designatus, IF 557461, as
Hypophyllum colus). Sweden, Västergötland, Österplana sn,
Österplana hed, decideous forest on calcareus soil (Corylus,
Quercus, Tilia), 15 Sep 1986, coll. H. Lindström etal. CFP
499, F41127 (S, epitypus hic designatus, IF 557462), Gen-
Bank No. MT934978 (ITS).
Table 1 (continued)
Current name Younger synonyms
C. wahkiacus Ammirati, Beug, Liimat. & Niskanen 2016
C. valgus Fr. 1838 C. depexus var. luminosus Carteret 2005; C. fuliginosus P.D. Orton
1964; C. olivaceostipitatus Carteret 2012; C. orbiculozonarius Rob.
Henry 1983; C. rheubarbarinus Rob. Henry 1956
C. venustissimus Bidaud 2002
C. venustus P. Karst. 1881 C. traganulus P.D. Orton 1983
C. vernalishastensis Bojantchev, Ammirati, Niskanen, & Liimat. 2017
C. vernalisierraensis Bojantchev, Ammirati, Niskanen, & Liimat.
2017
C. vernus H. Lindstr. & Melot 1994 C. erythrinus var. russulisporus Bohus 1979
C. vinaceobrunneus Ammirati, Beug, Liimat., Niskanen & O. Ceska
2016
C. vinaceogrisescens Ammirati, Beug, Liimat. & Niskanen 2016
C. violaceopapillatus Bidaud 2010
C. vulpicolor M.M. Moser & McKnight 1987 C. glandicolor var. exilis J. Favre 1955; C. plicatus Bidaud 2010

Fungal Diversity
1 3
Illustration. Brandrud et al. (1989: pl. A37) as C.
bulliardii.
Descriptions of the species. Brandrud etal. (1989: pl.
A37), Niskanen etal. (2012) as C. bulliardii.
Notes—The protologue by Fries (1838) has a refer-
ence to Paulet’s illustration of Hypophyllum colus that is
desingnated as the lectotype of the species. It shows a fun-
gus most similar to C. bulliardii although the lamellae are
pale and decurrent. Fries (1838) describes the species with
a pileus about 2.5–5 cm wide, a stipe about 3 mm wide
and growing in Pinus forests near Uppsala. The width of
the stipe is too narrow for C. bulliardii, the species has not
been found from Uppsala and is normally associated with
Quercus, Fagus and Corylus. Due to these contradictions,
Brandrud etal. (1989, 1992) decided to apply the name to
another species that occurs in pine forests which has a nar-
row stipe and a red orange veil, a character emphasized by
Fries (1838). However, in this case the species chosen by
Brandrud etal. (1989) does not fit with the type illustration
or Fries’s measurements of the pileus since C. colus sensu
Brandrud etal. (1989) is a very small and slender species
(pileus 0.5–3 cm wide). Our conclusion is that a species that
would completely fit to Fries’s description and the holotype
does not exist. However, since the name has been widely
used in the Nordic countries after its publication in Brandrud
etal. (1989) it is better to stabilize the name rather than treat
it as nomen dubium. Here, we choose to follow the type of
the species and suggest an epitype for the species that will
make C. bulliardii and C. colus synonyms.
Cortinarius craticius Fr., Epicr. syst. mycol. (Upsaliae): 282
(1838) [1836–1838]
Type: Finland, Satakunta, Ikaalinen, Seitseminen
National Park, Multiharju strict protections area, old, mesic
spruce forest (Picea abies) with some Pinus sylvestris, Bet-
ula and Populus tremula, 8 Sep 2005, coll. K. Liimatainen &
T. Niskanen 05-069, H6029911 (H, neotypus hic designatus,
IF 557463; K, isoneotypus), GenBank No. MT934988 (ITS).
Illustration. Stockholm’s herbarium, S0279; http://herba
rium.nrm.se/speci mens/S0279
Description of the species. Niskanen etal. (2012).
Cortinarius diosmus Kühner, Bull. mens. Soc. linn. Soc.
Bot. Lyon 24: 39 (1955)
Type: France, Haute-Savoie, Environs de Samoëns; forêt
de Bostan, sentier montant au chalet de Bostan, 6 Sep 1948,
coll. R. Kühner 00110647 (G, lectotypus hic designatus, IF
557464), GenBank No. MT935017 (ITS).
Descriptions of the species. Niskanen etal. (2012) as C.
argillaceosericeus ined.
Notes—Kühner (1955) made two collections of C. dios-
mus that are considered syntypes: 00110646 and 00110647
(G). The syntypes represent two different species that have
the following younger names: C. diosmoides Rob. Henry
and C. argillaceosericeus ined. Niskanen etal. (2012). Both
species fit the original concept of C. diosmus, but we choose
the specimen 110647 as the lectotype of C. diosmus (syn. C.
argillaceosericeus ined.) since that represents the more com-
mon and more widespread species based on our current data.
Cortinarius flabellus (Fr.) Fr., Epicr. syst. mycol. (Upsaliae):
300 (1838) [1836–1838]
Basionym: Agaricus flabellus Fr., Syst. mycol. (Lundae)
1: 231 (1821): sanctioned in Fr., Syst. mycol. 1: 231 (1821).
Type: Sweden, Uppland, Lena sn, S of Salsta slott, in rich
coniferous forest on calcarious ground, 23 Sep 1987, coll. H.
Lindström etal. CFP 672, F44866 (S, neotypus hic designa-
tus, IF 557465), GenBank No. MT935053 (ITS).
Illustration. Brandrud etal. (1998: pl. D35).
Descriptions of the species. Brandrud etal. (1998: pl.
D35).
Notes—In Brandrud etal. (1998) three collections of
C. flexipes var. flabellus (Fr.) H. Lindstr. & Melot are pre-
sented. They represent two species D35 and D45/D34. The
basidiomata in plate D35 fit best to Fries’ description of
Agaricus flabellus that has a dark olive to blackish brown
pileus whereas the other species, represented by plates D45
and D34 sometimes has a red brown pileus. Therefore, we
propose coll. H. Lindström etal. CFP 672 as the neotype of
this species. The name of the other species is C. lindstroemii
Niskanen, Kytöv. & Liimat.
Cortinarius gentilis (Fr.) Fr., Epicr. syst. mycol. (Upsaliae):
297 (1838) [1836–1838]
Basionym: Agaricus gentilis Fr., Syst. mycol. (Lundae)
1: 212 (1821): sanctioned in Fr., Syst. mycol. 1: 212 (1821).
Type: Norway, Oppland, Dokka kn, Vest-Torpa, in spruce
forest with blueberry (Picea, Salix), 15 Sep 1983, coll. H.
Lindström etal. CFP 178, F256849 (S, neotypus hic desig-
natus, IF 557466), GenBank No. EU266692 (ITS).
Illustration. Brandrud etal. (1992: pl. B31).
Descriptions of the species. Brandrud etal. (1992: pl.
B31), Niskanen etal. (2012).
Cortinarius helvelloides (Fr.) Fr., Epicr. syst. mycol.
(Upsaliae): 297 (1838) [1836–1838]
Basionym: Agaricus gentilis e helvelloides Fr., Syst.
mycol. (Lundae) 1:213 (1821).
Type: Finland, Uusimaa, Espoo, Hindsby-Svartböle,
under Alnus incana and Alnus glutinosa, among grasses, 17
Aug 2005, coll. anonymous, T. Niskanen 05-002, H6031432
(H, neotypus hic designatus, IF 557467; K, isoneotypus),
GenBank No. MT935110 (ITS).
Illustration. Brandrud etal. (1989: pl. A17).
Descriptions of the species. Brandrud etal. (1989: pl.
A17), Niskanen etal. (2012).

Fungal Diversity
1 3
Cortinarius hemitrichus (Pers.) Fr., Epicr. syst. mycol.
(Upsaliae): 302 (1838) [1836–1838]
Basionym: Agaricus hemitrichus Pers., Syn. meth. fung.
(Göttingen) 2: 296 (1801): sanctioned in Fr., Syst. mycol.
1: 230 (1821).
Type: Sweden, Skåne, Maglehem sn, ”Piraten rasten”, in
birch forest, 21 Sep 1987, coll. H. Lindström etal. CFP 662,
F44875 (S, neotypus hic designatus, IF 557468), GenBank
No. MT935113 (ITS).
Illustration. Brandrud etal. (1989: pl. A31).
Descriptions of the species. Brandrud etal. (1989: pl.
A31), Niskanen etal. (2012).
Cortinarius hinnuleus Fr., Epicr. syst. mycol. (Upsaliae):
296 (1838) [1836–1838]
Types: Sowerby, Col. Fig. Engl. Fungi Mushr.1: tab.
173, 1805 (lectotypus hic designatus, IF 557469). Sweden,
Medelpad, Torp sn, Hussborg, in cultivated grassland under
Betula, 28 Sep 1985, coll. H. Lindström etal. CFP 332,
F37503 (S, epitypus hic designatus, IF 557470), GenBank
No. DQ117926 (ITS).
Illustration. Brandrud etal. (1989: pl. A19).
Descriptions of the species. Brandrud etal. (1989: pl.
A19).
Notes—The name C. hinnuleus has been collectively used
for several deciduous forest species that have a yellowish
brown to reddish brown pileus, distant lamellae with an
earthy odour, white universal veil and strongly verrucose,
subglobose to obovoidly subglobose spores. They collec-
tively more or less fit to the Fries’s protologue (Fries 1838)
that describes a species with fulvous cinnamon pileus, dis-
tant lamellae and a white veil ring on the stipe that grows
early in the season in deciduous forests. The species in the
photograph of Brandrud etal. (1989), plate A19, fits Fries’s
protologue as well as Sowerby’s colour plate and therefore
we propose it as an epitype of the species.
Cortinarius laniger Fr., Epicr. syst. mycol. (Upsaliae): 292
(1838) [1836–1838]
Type: Finland, Joutsa, Koivuranta, W of Rakkolanselkä,
fairly young, mesic to damp, spruce-dominated (Picea abies)
forest with some Betula and Pinus, 30 Aug 2005, coll. K.
Liimatainen, S. Miettinen & T. Niskanen 05-019, 6029897
(H, neotypus hic designates, IF 557471; K, isoneotypus),
GenBank No. MT935187 (ITS).
Illustrations: Brandrud etal. (1994: pl. C53, mixed col-
lection), Fries (1867–1884: pl. 156)
Descriptions of the species: Brandrud etal. (1994: pl.
C53, mixed collection), Niskanen etal. (2012).
Notes — The plate C53 of C. laniger in Brandrud etal.
(1994) is a mixed collection also including C. distortus
Kauffman and therefore another specimen is proposed as
a neotype here.
Cortinarius melleopallens (Fr.) Britzelm., Bot. Zbl. 51(2-
3): 38 (1892)
Basionym: Cortinarius triformis var. melleopallens Fr.,
Epicr. syst. mycol. (Upsaliae): 299 (1838) [1836–1838].
Type: Sweden, Härjedalen, Storsjö sn, Flatruet, in sub-
alpine zone with Betula, Pinus, Picea, 16 Aug 1986, coll.
H. Lindström etal. CFP 433, F44880 (S, neotypus hic
designatus, IF 557472), GenBank No. MT935221 (ITS).
Illustration: Brandrud etal. (1992: pl. B12)
Descriptions of the species: Brandrud etal. (1992), Nis-
kanen etal. (2012).
Notes—Fries’ protologue does not perfectly fit to any
currently known Cortinarius species. Since there is no
clear solution, we decide to follow the Nordic concept of
this name (Brandrud etal. 1992, Niskanen etal. 2012).
For more nomenclatural discussion of this name and the
reasoning for the current interpretation see the booklet of
Brandrud etal. (1992).
Cortinarius miniatopus J.E. Lange, Fl. Agaric. Danic. 5
(Taxon. Consp.): III (1940)
Types: Lange, Fl. Agaric. Danic. 5(Taxon. Consp.):
III, Plate103 Fig. B, 1940 (lectotypus hic designatus,
IF 557473). Finland, Kainuu, Suomussalmi, Näljänkä,
Lohivaara, W of Kiviaro, SW side of the forest road, NE
sloping spruce forest with fairly rich grass-herb depres-
sions, Pinus, Betula, Populus tremula and Salix spp., 230
m, 13 Sep 1997, coll. I. Kytövuori 97-1369, H6041343
(H, epitypus hic designatus, IF 557474), GenBank No.
MT935228 (ITS).
Notes—This species has recently been called C. colus
(see also C. colus above) in the Nordic literature and listed
as a synonym of C. miniatopus in Brandrud etal. (1989).
However, the concept included two species, one with
large spores currently named C. subminiatopus Kytöv.,
Niskanen & Liimat., (photograph Brandrud etal. (1989;
A55)) and a sister species with smaller spores (7.0–9.0
x 4.5–5.5 μm, av.= 7.5–8.2 x 5.0–5.2 μm, Q=1.45–1.70,
Qav.= 1.52–1.62). The macroscopic description of C.
miniatopus by Lange (1940) fits both species well but
the spore size given is 6.5–7 x 4.3–4.5 μm. Although the
spore size in the protologue is even smaller than that of the
small-spored species we conclude that the small-spored
species fits best to the original description and here pro-
pose collection H6041343 as the epitype of the species.
Cortinarius paleaceus Fr., Epicr. syst. mycol. (Upsaliae):
302 (1838) [1836–1838]
current name Cortinarius hemitrichus (Pers.) Fr., Epicr.
syst. mycol. (Upsaliae): 302 (1838) [1836–1838]
Type: Sweden, Skåne, Maglehem sn, ”Piraten rasten”, in
birch forest, 21 Sep 1987, coll. H. Lindström etal. CFP 662,

Fungal Diversity
1 3
F44875 (S, neotypus hic designatus, IF 557475), GenBank
No. MT935265 (ITS).
Illustrations: Brandrud etal. (1989: pl. A31), Fries
(1867–1884: pl. 160)
Descriptions of the species: Brandrud et al. (1989:
pl. A31) as C. hemitrichus, Niskanen etal. (2012) as C.
hemitrichus.
Notes—The name C. paleaceus has often been applied to
C. flexipes (Pers.) Fr. coll. However, no odour, which is very
typical of species of C. sect. Flexipedes Kytöv., Niskanen
& Liimat., is mentioned in Fries’s protologue (Fries 1838)
and the lamellae are described as whitish when young. In
addition, a plate from Fries (1867–1884) illustrates a species
with pale lamellae and context of the stipe, a species that
looks like C. hemitrichus, and not like C. flexipes and rela-
tives that have darker lamellae and stipe context. Based on
this we conclude that our current interpretation of C. hemi-
trichus best represents also this species and a neotype mak-
ing these two names synonyms is suggested. Both names,
C. paleaceus and C. hemitrichus, were described by Fries
(1838) in the Epicrisis. Here we choose to continue the use
of the name C. hemitrichus as the current name of the spe-
cies to avoid a name change and confusion.
Cortinarius pholideus (Lilj.) Fr., Epicr. syst. mycol.
(Upsaliae): 282 (1838) [1836–1838]
Basionym: Agaricus pholideus Lilj., Utkast. Sv. Fl., Edn
3: 645 (1816).
Type: Sweden, Ångermanland, Säbrå sn, Näs, in dry
coniferous forest with blueberry and lichen (Betula, Picea),
29 Aug 1987, coll. H. Lindström etal. CFP 602, F248484
(S, neotypus hic designatus, IF 557476), GenBank No.
MT935303 (ITS).
Illustration: Brandrud etal. (1992: pl. B37).
Descriptions of the species: Brandrud etal. (1992: pl.
B37), Niskanen etal. (2012).
Cortinarius praestigiosus (Fr.) M.M. Moser, Schweiz. Z.
Pilzk. 43(8): 131 (1965)
Basionym: Cortinarius paragaudis var. praestigiosus Fr.,
Hymenomyc. eur. (Upsaliae): 379 (1874)
Type: Finland, Uusimaa, Vantaa, Tammisto, Tammisto
Nature Reserve Area, herb-rich mesic to dryish mixed for-
est (Quercus, Corylus, Betula, Pinus sylvestris and Populus
tremula), 17 Sept 2012, coll. K. Liimatainen & T. Niskanen
12-028, H6083157 (H, neotypus hic designatus, IF 557477;
K, isoneotypus), GenBank No. MT935314 (ITS).
Illustration: Brandrud etal. (2012: pl. E04).
Descriptions of the species: Brandrud etal. (2012: pl.
E04), Niskanen etal. (2012).
Cortinarius psammocephalus (Bull.) Fr., Epicr. syst.
mycol.: 301 (1838) nomen dubium
Basionym: Agaricus psammocephalus Bull., Herb. Fr.
(Paris) 13: 12, tab. 531, fig.2 (1793).
Notes —This species was described by Bulliard (1793)
and the only original material is the painted figure that
has been chosen as a lectotype of the species in Brandrud
etal. (1998). The plate illustrates a rather slender, brown
species with a wide, convex to low convex, sometimes low
umbonate, scaly pileus, and a scaly stipe, the lamellae are
brown. However, it is not obvious that Agaricus psammo-
cephalus would be a Cortinarius. The illustrated basidi-
omata are also reminiscent of species in the genus Inocybe
and the clustered growing habit reminds one of a sapro-
trophic fungus. The epithet psammocephalus was combined
in the genus Cortinarius by Fries (1838), who intepreted it
as a species growing in coniferous forests. Because Bulliard
worked in the Paris region, already Brandrud etal. (1998)
concluded, that Fries’s species most likely is different from
Bulliard’s species that supposedly was growing in a decidu-
ous forest. Currently, the name is applied to a species pair C.
castaneopallidus Carteret/C. quercoconicus Liimat., Kytöv.
& Niskanen that usually have a much narrower, acutely
umbonate pileus (Bidaud etal. 2004, plate481; Brandrud
etal. 1998, plate D57).
Taking into consideration that i) the basidiomata illus-
trated in the Bulliard’s plate do not fit the species for which
the name has currently been used, ii) the plate may represent
a species from another genus, and iii) we have not found
another candidate for the name from the genus Cortinarius,
we refrain to use the name for a species in genus Cortinarius
and treat is as a nomen dubium.
Cortinarius torvus (Fr.) Fr., Epicr. syst. mycol. (Upsaliae):
293 (1838) [1836–1838]
Basionym: Agaricus torvus Fr., Observ. mycol. (Havniae)
2: 80 (1818): sanctioned in Fr., Syst. mycol. 1: 211 (1821).
Types: Bulliard, Herb. Fr. (Paris) 2: Tab. 96, pl. 600, 1782
[1781-82] (lectotypus hic designatus, IF 557478, as Agari-
cus araneosus). Sweden, Skåne, Degeberga sn, Forsakar,
in beech forest on calcareous ground, 17 Sep 1988, coll. H.
Lindström etal. CFP 778, F248482 (S, epitypus hic desig-
natus, IF 557479), GenBank No. MT935556 (ITS).
Illustration. Brandrud etal. (1992: pl. B13).
Descriptions of the species. Brandrud etal. (1992: pl.
B13), Niskanen etal. (2012).
Cortinarius traganus (Fr.) Fr., Epicr. syst. mycol.
(Upsaliae): 281 (1838) [1836–1838]
Basionym: Agaricus traganus Fr., Observ. mycol.
(Havniae) 2: 82 (1818): sanctioned in Fr., Syst. mycol. 1:
217 (1821).
Types: Schaeff., Fung. bavar. palat. nasc. (Ratis-
bonae) 1–2: Tab 56, Fig. I-V, 1774 (lectotypus hic desig-
natus, IF 557480), as Agaricus amethystinus). Sweden,

Fungal Diversity
1 3
Ångermanland, Härnösand, Härnön at Myran, in dry sandy
pine forest, 13 Sep 1988, H. Lindström etal. CFP763,
F248486 (S epitypus hic designatus, IF 557481), GenBank
No. MT935361 (ITS).
Illustration. Brandrud etal. (1994: pl. C04).
Descriptions of the species. Brandrud etal. (1994: pl.
C04), Niskanen etal. (2012).
Notes—The protologue by Fries (1818) is very short but
mentions the main characteristics of the species currently
considered as C. traganus (Brandrud etal. 1994; Niskanen
etal. 2012): Basidiomata with a smell. Pileus pale lilac, stipe
whitish purplish and bulbous, context yellow. Fries (1818)
also refers to an illustration of Schaeffer (1774) that then
becomes the type of the species. A majority of the figures
in the illustration represent our interpretation of C. traganus
(Fig. I-V), but Fig. VII clearly shows a typical characteristic
of C. cyanites Fr.: the context of the stipe and pileipellis have
become vinaceous red on exposure. In Fig. IX the spores
are round which does not fit either of the above species,
a potential species could be found from C. sect. Anomali
where species with round spores and bluish colours occur. It
seems that the type of C. traganus is a mixed illustration, but
since the majority of the figures and the protologue fit the
current concept of C. traganus, we here choose an epitype
to support this interpretation.
Cortinarius turgidus Fr., Epicr. syst. mycol. (Upsaliae): 278
(1838) [1836–1838]
Types: Battarra, Fungorum agri Arimensis historia: tab. 9
fig. C, 1755 (lectotypus hic designatus, IF 557482; as Mono-
myces ventricosus). Sweden, Bohuslän, Sotenäs, Tossene, E
of Bovallstrand, Hogsäms bokskog, Fagus forest with some
Betula and Populus, seashells on ground, 29 Sep 2004, coll.
K. Liimatainen & T. Niskanen 04-1020, H7017832 (H, epi-
typus hic designatus, F 557483; K, isoepitypus), GenBank
No. MT935565 (ITS).
Illustration: Brandrud etal. (1992: pl. B58).
Descriptions of the species: Brandrud etal. (1992: pl.
B58), Niskanen etal. (2012).
New combinations
Cortinarius colorius (Bidaud) Niskanen, Dima & Liimat.
comb. nov.
IF 557484
Basionym: Cortinarius ignifluus var. colorius Bidaud, in
Bidaud etal., Atlas des Cortinaires (Meyzieu) 6: 190 (1994)
Cortinarius iners (Bidaud) Liimat., Dima & Niskanen
comb. nov.
IF 557485
Basionym: Cortinarius duracinus f. iners Bidaud, in
Bidaud etal., Atlas des Cortinaires (Meyzieu) 17(1): 1176
(2008)
Discussion
Studies oftype specimens
There are two ways for naming a barcode in a sequence
database: either sequence a named voucher specimen based
on a morphological identification or sequence a type speci-
men. Paradoxically, the first approach is currently the most
widely used although the core reason for using the DNA-
based identification is the unreliability of the morphological
identification. The gold standard should be sequencing the
type specimens to achieve an unambiguous, good quality
identification database, but this unfortunately has thus far
been generally neglected.
To improve the sequence-based identification of the
important ectomycorrhizal genus Cortinarius and create a
solid base for future taxonomic work 482 type specimens
were sequenced. This is more than twice as many as the
largest type study of Cortinarius so far (Liimatainen etal.
2014). We were able to successfully sequence many old type
specimens; 105 types which were over 50 years old and 18
over 100 years old. This shows that most available Corti-
narius type specimens can likely be sequenced regardless of
the age of the specimen. The dataset, including the already
published type sequences in this group, contains a total of
363 species. About half of these species’ names, altogether
184, are published now for the first time in GenBank, thus
doubling the reliably of barcoded species of Cortinarius,
subgen. Telamonia in the public sequence databases. Also
33% of the species represented here have been described
over the last decade using DNA sequences alongside mor-
phology and ecological data. Adding DNA tools for fungal
taxonomy has accelerated the process of discovering and
describing fungus diversity.
Synonyms
Our dataset shows that many species have been described
several times. Of the 363 species recognized in this study
31% have a synonym, the synonym rate is even higher
with species described using only morphological char-
acters (46%). The two main reasons for synonyms are
that the interpretation of the existing names has been
challenging and there have been problems based on the
morphological species concept. The high number of spe-
cies, convergent evolution and the small number of useful
morphological characters for classification have not made
the task any easier. Also, the lack of uniform and stable

Fungal Diversity
1 3
infrageneric classification has made it more difficult to
find potential, already existing descriptions of the species
and thus many species have been subsequently named as
new again. In the future, the problem of synonyms will
be much reduced when sequences from type specimens
are available and the description of new species without
ITS barcodes are strongly discouraged.
One example of the difficulty of interpret existing
names is C. impolitus Kauffman. It was the species
described the most times by multiple authors over dec-
ades, e.g. by Kauffman (1918) and Smith (1944) from
North America and by Velenovský (1939), Pearson
(1946), Favre (1955) and Lindström (Brandrud etal.
1998) from Europe (Table1 and Figure1). The species is
small and brown which partly explains the problem but
it also has two good characters, odour of Pelargonium in
the lamellae and narrow basidiospores, but despite these
characteristics it has been very challenging to recognize
it from the works of different mycologists based on mor-
phology only.
Examples of the second problem, the challenges of
using the morphological species concept, are C. macropo-
dius Rob. Henry and C. luridus Rob. Henry that overall
had the highest number of synonyms, 13 and 9 respec-
tively. In this case, all synonyms come from the French
authors and are due to a too narrow species concept.
Some of the synonyms are also placed in different infra-
generic groups in their classification system. This error
rate and unnatural classification make it very difficult to
use the earlier parts of the Atlas des Cortinaires series
for identification of Cortinarius. However, the individual
descriptions of the species are usually of good quality and
61 species names that have been described by the team are
the oldest names for the species: representing about 15 %
of all the currently known species of C. subgenus Tela-
monia. In recent years they have also included molecular
data into their work which has greatly improved the out-
come (e.g. Bidaud etal. 2017).
When looking at the rate at which the different authors
described synonyms it is self-evident that it was easier
to describe new species earlier when more species were
undescribed. For example, the error rate of Kauffman
is only 7% whereas Smith’s error is double that, most
likely because he was partly describing the species from
the same area where Peck and Kauffman had previously
worked. Half of the Smith’s synonyms are Kauffman’s
species. The error rate of Moser and Henry are rather sim-
ilar, which is a bit surprising since they mainly worked
in different habitats and with a different species concept.
Interpretation andtypication oftheearly names
withouttype materials
Many early names without type specimens have been re-
described by later authors. From all the old names used in
this study only 10 of them are without synonyms: C. armil-
latus (Fr.) Fr., C. bibulus, C. bovinus Fr., C. cinnabarinus
Fr., C. colymbadinus Fr., C. dolabratus Fr., C. evernius (Fr.)
Fr., C. gentilis, C. glandicolor (Fr.) Fr., and C. helvelloides.
About half of them are rather characteristic and easy to inter-
pret so taxonomists after Fries understood his concept and
therefore did not describe those species again, i.e. C. armil-
latus and C. evernius. On the other hand, some of these
species are really difficult to interpret and might not have
been described again just because of the restricted distribu-
tion, infrequent occurrence or just a matter of chance, i.e. C.
bovinus and C. dolabratus.
Interpreting the early names, like those of Fries and Per-
soon, when often no physical specimen is left to study and
the descriptions themselves are short, vague and without
microscopical characters, is extremely difficult. In many
cases their species concept most likely included several spe-
cies and was too generalized. They surely did their best but
the state of knowledge in those times was far from what we
know now. For example, C. paragaudis and C. praestigio-
sus, two species which based on current, widely accepted
concept are far from each other both phylogenetically and
morphologically, were included as varieties of one species
in Fries’ concept (Fries 1874). In Cortinarius sect. Bovini
only one species was described by Fries although the section
includes at least seven species in Sweden (Niskanen etal.
2013). Of course, Fries might not have found all those spe-
cies in the areas he collected or did not have time to work
with them, but it is still rather certain that many Fries’ names
included several species. Thus, due to the broad species con-
cept there often is not any correct one candidate for epi- or
neotypification. And even if there has been a clear concept
behind the early species descriptions, it is often very difficult
to interpret based on short and vague descriptions.
The interpretation of a name based only on morphology
is a demanding, often impossible, task. In this study ca. 80%
of the species described by Fries have been described again.
The poor record can not be explained by a few poor stud-
ies or unprofessional authors—all major Telamonia authors
have misinterpreted Fries’ names or simply overlooked
them. Studying the type specimens of Karsten’s species gave
a similar result. Karsten’s descriptions are somewhat better
than Fries’ since they also include microscopical character-
istics, but the critical difference is that Karsten’s specimens
are available and can be sequenced, thus we really can con-
firm the true identity of his species. The result was that all
the seven Telamonia species described by Karsten, which
we studied, have been redescribed later by other authors

Fungal Diversity
1 3
confirming the conclusion from Fries’ materials. No cur-
rent data supports the claim that the early names could be
interperated correctly and consistently by anyone.
Because of the problems mentioned above the interpreta-
tion of early names in general is not a very meaningful thing
to do and often the outcome is highly questionable. The
majority of early names should probably be treated as nomen
dubium. Therefore, we only typified those early names that
have been widely used, e.g. appear on many national check
lists or are commonly used in books like Funga Nordica
(Niskanen etal. 2012). In these cases, the typification is a
quicker and a more efficient way to stabilize nomenclature
than trying to convince users to stop using the name. Also, it
is important to point out that when typifying early names, we
do not claim that the outcome would be correct, i.e. would
represent the original concept of the author. We simply try to
find the species that would best fit to the original description
and in the case of several equally suitable candidates choose
the most practical solution, i.e. the one that causes fewest
changes in the current use of the name, the species itself
would be the most common and wide spread of the candidate
species and/or the easiest to recognize.
Another problem with the old names is the references
to the illustrations. At those early times authors did not
know that the references would later turn out to be the most
important part of the descriptions—based on the current
International Code of Nomenclature for algae, fungi, and
plants (https ://www.iapt-taxon .org/nomen /main.php), they
are considered as ‘original material’ of the species. At the
time there was not a huge amount of published illustrations
to choose from. It seems that in some cases Fries referred to
an illustration that did not fit perfectly to his concept of the
species but was the closest one with some similarity. This is
e.g. obvious with C. colus and C. turgidus.
Nomenclatural coverage ofthedataset
andconclusions
In this study we tried to sequence all species level type spec-
imens belonging to Cortinarius subgen. Telamonia that have
not been previously studied. Our aim also was to stabilize
all commonly used early names for which a type specimen
does not exist. Obviously, all names in Telamonia are not in
this dataset. Some type specimens could not be sequenced,
especially Hongo’s and Murrill’s types failed almost without
exception. Also, Henry’s material was difficult to sequence
and in addition, many of his type specimens were not found,
the names are nomenclaturally invalid, or had other prob-
lems. Most of the Peck’s material could not be acquired
from NYS during the time of the molecular study of this
paper. Some of Favre’s type specimens were too small to
sample or have already been sequenced but not published
by other authors. Melot’s type specimens are in his personal
collection and despite several attempts to aquire them on
loan, they were not avalaible for molecular study. Unless this
situation changes the identity of the names remains unclear
and it would be better not to use them to avoid confusion
rising from the different interpretations of the names. There
are a few authors whose materials we have not studied, e.g.
Bon and Lamoure, but the number of Telamonia species
they described is relatively small, only some tens of species.
After this study there will only be a few dozen valid
names that have not yet been studied with molecular meth-
ods and where the type specimens are good quality for
sequencing and available for study. Most likely many of
them have an earlier name which already have been studied.
There are a few exceptions, however, for example if one is
working with the sub-alpine Telamonia species the names
described by Favre (e.g. 1955) and Lamoure (1977, 1978)
are relevant, for Eastern North America species described by
C. H. Peck’s should be checked (Burnham 1919; Gilbertson
1962), and for European Mediterranean areas the works of
local authors would be appropriate to study (e.g. Mahiques
and Ortega 2002). Otherwise, if a new sequence does not
have ≥ 99% similarity to any published type sequence it can
be rather certain that it derived from an undescribed species,
given that the quality of the sequence is good.
Overall, our data set contains about 300 species from
Europe and 150 species from North America and many of
which they have in common. There may only be a few hun-
dred more Telamonia species to be found from Europe, but
certainly in North America the quest has just begun. The
situation in Africa, Asia and Central and South America is
praticially unknown, but it would not be an exaggeration to
predict that the world-wide diversity of Cortinarius subgen.
Telamonia would be a four-digit number. Thus far, members
of the subgenus have not been found in the Nothofagus for-
ests of New Zealand (Soop etal. 2019) and from Nothofa-
gus forests of South America only one species is confirmed
(Garnica etal. 2005).
As species are discovered and named the easier the iden-
tification based on ITS will become. Unfortunately, the same
does not apply to morphological identification. All the cur-
rent keys we use would require extensive rewriting and even
though there often are morphological and/or ecological dif-
ferences between the species, identifying many of the spe-
cies of Telamonia using keys without deeper experience and
knowledge of the group will be challenging if not impossi-
ble. Having local keys (i.e. Scandinavian boreal Telamonias
or Telamonias of the Pacific North West) and in certain cases
only trying to identify sections or species complexes rather
than species would be the most realistic approach when
using morphological identification.
Many times, the biggest obstacle for efficient identifi-
cation and naming of alfa diversity are the nomenclatoral
problems, i.e. what is the correct name for the species or is

Fungal Diversity
1 3
it an undesribed one? The species of C. subgenus Telamonia
have been considered one of the most challenging cases in
the Agaricales at the species level. Its high diversity com-
bined with convergent, similar appearing taxa have earned
it a reputation of being an impossible group to study, one
better left in the forest. Our study shows that nomenclatoral
problems, even in difficult groups like Telamonia, can be
solved and identification based on ITS barcodes becomes
an easy task even for non-experts.
Infrasubgeneric classication
Relationships ofthesections withinCortinarius
subgen. Telamonia
The relationships of the sections within Telamonia remain
unclear in our phylogenetic analysis. The grouping of the
sections in the tree, however, does not seem random and
makes sense when compared to the morphological charac-
teristics. Therefore, the main findings that we feel would be
of importance are summarized below and could be used as
starting hypotheses for future studies.
Based on our phylogenetic analysis C. subgen. Telamonia
is roughly divided into two main entities (Fig.2): (i) The
basal groups of the tree (”Basal Telamonias”) that only con-
tain species with medium- to large-sized basidiomata (the
apex of the stipe is > 4 mm wide) with the exception of a
few species in C. sect. Brunnei. (ii) The monophyletic upper
part of the tree (“Crown Telamonias”) that mainly contains
species with small basidiomata (the apex of the stipe is <
4 mm wide), and the following sections including species
with mainly small- to medium-, less commonly large-sized
basidiomata: Hinnulei Melot, Rubricosi Moënne-Locc. &
Reumaux Leiocastanei Niskanen, Kytöv. & Liimat., and
the monotypic sections Pseudoduracini Liimat., Niskanen
& Kytöv., Friesiorum Liimat., Kytöv. & Niskanen, and Vina-
ceobrunnei Ammirati, Niskanen & Liimat.. The most basal
part of this clade also includes sections Anthracini Melot,
Crassispori Kytöv., Niskanen & Liimat. and Squalidi Lii-
mat., Ammirati & Niskanen.
Within the “Crown Telamonias” some further group-
ing can be observed. Brandrud etal. (1989) initially clas-
sified the species with small basidiomata into two sections,
Incrustati Melot and Hydrocybe (Fr. ex Rabenh.) P. Karst.
Although not forming well supported clades, these two ear-
lier groups seem to correlate with the phylogeny to some
extent. In the Fig.2 the groups are named as /Squamicybe
(Incrustati s. Brandrud etal.) and /Erubescentes (Hydrocybe
s. Brandrud etal.). The new names are introduced because
the type species of sect. Hydrocybe, Cortinarius duracinus,
does not belong to “Crown Telamonias” but to “Basal Tel-
amonias” and the type species of sect. Incrustati, C. lux-
nymphae, was not available for study and thus the identity
of the species remains unclear.
The previous members of the C. sect. Incrustati are all
placed in/Squamicybe (Fig.2) in two monophyletic groups/
Eusquamicybe and/Paludosi but the group also includes sec-
tions of species with medium- to large-sized basidiomata.
Many species of this group have a ±scaly pileus, a univer-
sal veil that forms distinct girdles on the stipe and a stipe/
context of the stipe that becomes darker towards the base,
especially with age. No part of the basidiomata turns red-
dish (except in C. sect. Rubrocincti that resembles more the
species in /Erubescentes). Typical examples of this group
are C. flexipes (C. sect. Flexipedes), C. hemitrichus (C.
sect. Paleacei) and C. saniosus (Fr.) Fr. (C. sect. Saniosi
Moënne-Locc. & Reumaux) and from the larger species
C. hinnuleus (C. sect. Hinnulei) and C. rubricosus (Fr.) Fr.
(syn. C. safranopes Rob. Henry, C. sect. Rubricosi). Species
associated with Alnus, i.e. C. bibulus (C. sect. Bibuli), C.
griseocarneus Carteret (C. alnetorum (Velen.) M.M. Moser
sensu Brandrud etal. 1989, C. sect. Alnicolarum) and C.
helvelloides (C. sect. Helvelloides), also belong to this larger
group. Together with C. sect. Saniosi they form a monophy-
letic group /Paludosi, although without support, indicating
that within C. subgen. Telamonia the ability to form mycor-
rhizae with Alnus may only have evolved once.
A majority of the species classified earlier in C. sect.
Hydrocybe are placed in another, monophyletic, group /
Erubescentes (Fig2). They all have small basidiomata and
a smooth pileus and in most species the stipe/context of the
stipe does not become darker towards the base. In addition,
in quite a few species either the base of the stipe, univer-
sal veil and/or basal mycelium turns ± reddish with time.
The universal veil varies from indistinct to forming distinct
girdles on the stipe. Typical examples of this group are C.
fuscoalbus Kytöv., Niskanen & Liimat. (C. sect. Atroalbi
Niskanen, Kytöv. & Liimat.), C. decipiens (Pers.) Fr. (C.
sect. Castanei Moënne-Locc. & Reumaux), C. praestigiosus
(C. sect. Praestigiosi Kytöv., Niskanen & Liimat.) and C.
vernus H. Lindstr. & Melot (C. sect. Verni Kytöv., Niskanen
& Liimat.).
Sections
The aim of this study was not to solve the infrasubgeneric
classification of C. subgen. Telamonia but to show the pre-
liminary placement of the studied species and existing sec-
tions (Figs.1 and 2). Examples of the species belonging
to the sections are shown in Supplementary Fig.1–11. We
included representative photographs for all but the following

Fungal Diversity
1 3
three sections: C. sect. Cacaodisci Kytöv., Niskanen & Lii-
mat., C. sect. Pseudoduracini, and C. sect. Squalidi. In this
study, 80 previously described sections and 9 subsections are
used, and additional 11 section names are considered syno-
nyms. A small number of species are not currently placed in
any of the sections. The “Basal Telamonias” with medium-
to large-sized basidiomata have been easier to study and are
thus better known and only four species, C. hepaticus Kytöv.,
Niskanen & Liimat., C. hillieri Rob. Henry, C. uraceisporus
Niskanen, Kytöv. & Liimat. and one C. sp., remain outside
the currently accepted sections. In the “Crown Telamonias”,
that have been more overlooked mainly due to their small
size, 18 species included in our phylogenetic analysis remain
unclassified. Some of them, like C. denigratus Ammirati,
Beug, Niskanen, Liimat. & O. Ceska and the related C. spp
from North America that form a monophyletic group and
differ > 4% (>20 indels and substitutions) from other spe-
cies of Cortinarius subgen. Telamonia, might be considered
as a new section in the future. Some may be grouped with
existing sections with futher analysis using additional DNA
regions, i.e. C. ferrugineovelatus Kytöv. Liimat. & Niskanen
and C. umbrinobellus Liimat., Niskanen & Kytöv. that share
morphological characteristics with the species in C. sect.
Praestigiosi but were currently placed in a basal position of
the branch containing that section.
We wanted the section names to be as unambiguous as
the species names as far as possible and therefore we only
accepted section names that can be interpreted without a
doubt, i.e. the type specimen of the type species of a section
is sequenced. Exceptions were made with four names: sect.
Anthracini Melot, sect. Brunneotincti M.M. Moser, sect.
Cinnabarini Melot, and sect. Parvuli Melot. For these sec-
tions we have not been able to study the type specimen of
the type species for several reasons or the sequencing failed
but we believe that the concept of the type species is rather
uniform and clear (e.g. Niskanen etal. 2012). Therefore, it
seems acceptable to use these common section names. We
are aware that this kind of approach is risky as the case of
sect. Testaceofolii Liimat., Niskanen & Kytöv. shows. At the
time, it was clear that C. biformis Fr. sensu Funga Nordica
(e.g. Niskanen etal. 2012) was a different species than C.
testaceofolius H. Lindstr. & Soop. However, later it turned
out that the neotype Moser had selected for C. biformis was
in fact an older synonym for C. testaceofolius, an outcome
that no one had previously thought possible. Therefore, sect.
Biformes Moënne-Locc. & Reumaux and sect. Testaceofolii
are now synonyms.
If the species concept is often difficult to apply, then clas-
sification above species becomes even more subjective. In
general, we should try to avoid having too many monotypic
entities since they are less meaningful in classification.
However, the risk with bigger entities is having units which
would have very little, if any, exclusive morphological char-
acters that would define those groups, since one of the main
reasons of having a higher-level classification is to recog-
nize groups with unique character states. For example, C.
armillatus, C. sect. Armillati, C. subgen. Telamonia, genus
Cortinarius represent four levels of classification in which
the species C. armillatus belongs to all four groups that have
their own, unique defining characters that other groups in
higher or lower levels do not have.
In this study, one example of the difficulties of delimiting
a section is C. sect. Uracei. With a wider concept it includes
several previously recognized sections, C. sect. Cinnabarini,
C. sect. Colymbadini, and C. sect. Miniatopodes Moënne-
Locc. & Reumaux, that all form a monophyletic clade with
good support value and morphological differences from C.
sect. Uracei s. str. Therefore, keeping all above-mentioned
sections separate would be an arguable choice, but then
there would be at least more than four monotypic sections
inside the clade Uracei that would need a new name. In this
case we have currently delimited C. sect. Uracei in a broad
sense because the group is also supported by morphologi-
cal characters. The other existing sections are treated at the
subsection level.
We have tried to delimit the sections to be the widest
monophyletic group with a reasonable support value and
with at least some shared morphological character states.
This approach and level of grouping mainly corresponds to
the concepts previously used to delimit the sections in the
genus Cortinarius in the era of molecular data (e.g. Ammi-
rati etal. 2013, 2017; Dima etal. 2014; Liimatainen etal.
2015, 2017, 2020; Niskanen etal. 2009, 2011, 2013; San
Fabian etal. 2018; Soop etal. 2019).
The sections identified here vary from monotypic entities
i.e. C. sect. Brunneocalcarii Niskanen, Liimat. & Kytöv.
to middle-sized groups i.e. C. sect. Armillati and C. sect.
Disjungendi to very diverse groups i.e. C. sect. Bovini and
C. sect. Uracei. The imbalance is unlikely to be solved due
to the speciation history of different groups, likely some of
them have diversified more than the others which has led
to the current species-poor and species-rich groups. Also,
this dataset only contains a fraction of the true diversity of
C. subgen. Telamonia worldwide and therefore the number
and the species diversity of sections will change when more
data are available. Most of the monotypic sections will most
likely turn out to be multi-species sections as shown e.g. by
Soop etal. (2019).
This is the first extensive phylogenetic study of C. sub-
gen. Telamonia. The great majority of sections and species
are shown in a phylogenetic context for the first time. Also,
many sections previously included in phylogenetic studies

Fungal Diversity
1 3
now contain more species and therefore seem to have better
support values. For example, Harrower etal. (2011) used the
same two DNA regions, ITS and LSU, in their study and got
less than BS 50% support for C. sect. Firmiores (Fr.) Hen-
nings when including three species in their analysis. In our
study that contains 20 species the support value for the same
section was BS 88%. For C. sect. Armillati the correspond-
ing values were BS 54% (2 species) and 90% (7 species).
Conclusions
Fig.1 shows our current view of the number of the sec-
tions in Cortinarius subgen. Telamonia and which species
we include in them. The earlier delimitations based on mor-
phology have been partly incorrect and included only a part,
often a small fraction, of the species (Bidaud etal. 2017;
Brandrud etal. 2012; Niskanen etal. 2012). The classifica-
tion presented here is a major step forward and can be used
as a basis for a more thorough revision of morphological
characterstics of the groups in the future.
Now that the nomenclatoral history of the last 100 years
has been sorted out for many taxa, everyone can benefit from
the outcome and continue to improve the understanding of
this diverse group of species. Fortunately, all current Cor-
tinarius taxonomists produce an ITS barcode of the type
specimen of new species and upload and annotate the new
sequence in GenBank. We hope that mycologists working
on Cortinarius and other genera will build on the findings
reported here.
Acknowledgements The help that we received from the curators of
E, FLAS, G, H, IB, K, MICH, NYSM, O, OULU, PC, PRM, S and
WTU was essential to this project. The following curators are warmly
thanked: Bart Buyck, Xavier Carteret, Philippe Clerc, Regina Kühner-
Winkler, Ursula Peintner, and Patricia Rogers. We are grateful to Jesko
Kleine for his invaluable help in the typification process. Mike Beug is
thanked for providing the photos of C. albosericeus and C. vinaceogris-
escens, Geert Schmidt-Stohn for providing the photos of C. phaeosmus
and C. valgus, Jodi Friesen and Marty Kranabetter for providing the
photo of C. ochropallens, Cathy Cripps for providing the photo of C.
flavobasilis, and Andy Overall for providing the photo of C. punctati-
formis (published in Overall (2017): Fungi - Mushrooms & Toadstools
of parks, gardens, heaths and woodlands).This work was supported by
the Ministry of Environment, Finland (YM38/5512/2009), The Finnish
Cultural Foundation, Daniel E. Stuntz Memorial Foundation (Univ. of
Washington, USA), Swedish taxonomy Initiative-project (University of
Gothenburg), Kone Foundation (FinBOL project) and the ELTE Insti-
tutional Excellence Program by the National Research, Development
and Innovation Office of Hungary (NKFIH-1157-8/2019-DT). Lastly,
we would like to thank the reviewers for their constructive comments,
which helped us to improve the manuscript.
Open Access This article is licensed under a Creative Commons Attri-
bution 4.0 International License, which permits use, sharing, adapta-
tion, distribution and reproduction in any medium or format, as long
as you give appropriate credit to the original author(s) and the source,
provide a link to the Creative Commons licence, and indicate if changes
were made. The images or other third party material in this article are
included in the article’s Creative Commons licence, unless indicated
otherwise in a credit line to the material. If material is not included in
the article’s Creative Commons licence and your intended use is not
permitted by statutory regulation or exceeds the permitted use, you will
need to obtain permission directly from the copyright holder. To view a
copy of this licence, visit http://creat iveco mmons .org/licen ses/by/4.0/.
References
Ammirati JF, Hughes KW, Liimatainen K, Niskanen T, Matheny
PB (2013) Cortinarius hesleri from eastern North America and
related species from Europe and western North America. Botany
91(2):91–98
Ammirati JF, Niskanen T, Liimatainen K, Dimitar B, Peintner U, Kuh-
nert-Finkernagel R, Cripps C (2017) Spring and early summer
species of Cortinarius, subgenus Telamonia, section Colymbadini
and /Flavobasilis, in the mountains of western North America.
Mycologia 109(3):443–458
Bidaud A, Bellanger J-M, Carteret X, Reumaux P, Moënne-Loccoz
P (2017) Atlas des Cortinaires. Pars XXIV, Èditions Fèdération
mycologique Dauphiné-Savoie, Lomazzo, France
Bidaud A, Carteret X, Eyssartier G, Moënne-Loccoz P, Reumaux
P (2004) Atlas des Cortinaires. Pars XIV, Èditions Fèdération
mycologique Dauphiné-Savoie, Lomazzo, France
Bidaud A, Moënne-Loccoz P, Reumaux P (1994) Atlas des Corti-
naires. Pars VI, Èditions Fèdération mycologique Dauphiné-
Savoie, France
Bidaud A, Moënne-Loccoz P, Reumaux P, Carteret X, Eyssartier G
(2008) Atlas des Cortinaires. Pars XVII, Èditions Fèdération
mycologique Dauphiné-Savoie, Lomazzo, France
Borchsenius F (2009) FastGap 1.2. Department of Biosciences,
Aarhus University, Denmark. Published online at http://www.
aubot .dk/FastG ap_home.htm
Brandrud TE, Lindström H, Marklund H, Melot J, Muskos S (1989)
Cortinarius Flora Photographica. Vol. I (Swedish version). Cor-
tinarius HB, Matfors, Sweden.
Brandrud TE, Lindström H, Marklund H, Melot J, Muskos S (1992)
Cortinarius Flora Photographica. Vol. II (Swedish version).—
Cortinarius HB, Matfors, Sweden.
Brandrud TE, Lindström H, Marklund H, Melot J, Muskos S (1994)
Cortinarius Flora Photographica. Vol. III (Swedish version).—
Cortinarius HB, Matfors, Sweden.
Brandrud TE, Lindström H, Marklund H, Melot J, Muskos S (1998)
Cortinarius Flora Photographica. Vol. IV (Swedish version).
Cortinarius HB, Matfors, Sweden.
Brandrud TE, Lindström H, Marklund H, Melot J, Muskos S (2012)
Cortinarius Flora Photographica. Vol. V (Swedish version).
Cortinarius HB, Matfors, Sweden.
Bulliard P (1793) Herbier de la France 13. Paris, France.
Burnham SH (1919) Charles Horton Peck. Mycologia 11(1):33–39.
https ://doi.org/10.1080/00275 514.1919.12016 772
Dima B, Liimatainen K, Niskanen T, Kytövuori I, Bojantchev D
(2014) Two new species of Cortinarius, subgenus Telamonia,
sections Colymbadini and Uracei, from Europe. Mycol Progress
13:867–879. https ://doi.org/10.1007/s1155 7-014-0970-6
Favre J (1955) Les champignons supérieurs de la zone alpine du
Parc National suisse. Ergebn. wiss. Unters. schweiz. NatnParks
V 33:1–212
Fries EM (1818) Observationes Mycologicae 2. Bonnier, Copenha-
gen, Denmark
Fries EM (1836–1838) Epicrisis systematis mycologici seu synopsis
Hymenomycetum. Uppsala, Sweden.

Fungal Diversity
1 3
Fries EM (1867–1884) Icones selectae Hymenomycetum nondum
delineatorum. Nordstedt & Son, Uppsala, Sweden.
Fries EM (1874) Hymenomycetum Eur. Uppsala, Sweden
Frøslev TG, Jeppesen TS, Læssoe T, Kjøller R (2007) Molecular
phylogenetics and delimitation of species in Cortinarius sec-
tion Calochroi (Basidiomycota, Agaricales) in Europe. Mol
Phyl Evol 44:217–227
Galtier N, Gouy M, Gautier C (1996) SEAVIEW and PHYLO_WIN:
two graphic tools for sequence alignment and molecular phy-
logeny. Bioinformatics 12:543–548
Gardes M, Bruns TD (1993) ITS primers with enhanced specifity for
basidiomycetes: Application to the identification of mycorrhizae
and rusts. Mol Ecol 2:113–118. https ://doi.org/10.1111/j.1365-
294X.1993.tb000 05.x
Garnica S, Schön ME, Abarenkov K, Riess K, Liimatainen K, Nis-
kanen T, Dima B, Soop K, Frøslev TG, Jeppesen TS, Peint-
ner U, Kuhnert R, Brandrud TE, Saar G, Oertel B, Ammirati
JF (2016) Determining threshold values for barcoding fungi:
Lessons from Cortinarius (Basidiomycota), a highly diverse
and widespread ectomycorrhizal genus. FEMS Microbiol Ecol
92(4):fiw045. https ://doi.org/10.1093/femse c/fiw04 5
Garnica S, Weiß M, Oertel B, Oberwinkler F (2005) A framework for a
phylogenetic classification in the genus Cortinarius (Basidiomycota,
Agaricales) derived from morphological and molecular data. Can
J Bot 83:1457–1477
Gilbertson RL (1962) Index to species and varieties of fungi described by
C.H. Peck from 1909 to 1915. Mycologia 54(5):460–465
Harrower E, Ammirati JF, Cappuccino AA, Ceska O, Kranabetter JM,
Kroeger P, Lim S, Taylor T, Berbee ML (2011) Cortinarius species
diversity in British Columbia and molecular phylogenetic compari-
son with European specimen sequences. Botany 89:799–810. https
://doi.org/10.1139/b11-065
Høiland K (1983) Cortinarius subgenus Dermocybe. Opera Botanica
71:1–113
Kauffman CH (1918) The Agaricaceae of Michigan. Biol. Ser 5
26:314–442
Katoh K, Kuma K, Toh H, Miyata T (2005) MAFFT version 5: improve-
ment in accuracy of multiple sequence alignment. Nucleic Acids Res
33(2):511–518. https ://doi.org/10.1093/nar/gki19 8
Katoh K, Standley DM (2013) MAFFT Multiple Sequence Alignment
Software Version 7: improvements in performance and usability.
Mol Biol Evol 30:772–780
Kirk PM, Cannon PF, Minter DW, Stalpers JA (2008) Dictionary of the
fungi, 10th edn. CAB International, Wallingford, UK
Kõljalg U, Nilsson H, Abarenkov K etal (2013) Towards a unified
paradigm for sequence-based identification of fungi. Mol Ecol
22:5271–5277
Kühner R (1955) Compeléments à la Flore analytique IV. Bull Mens Soc
linn Soc Bot Lyon 24(2):39–54
Lamoure D (1977) Agaricales de la zone alpine. Genre Cortinarius Fr.
sous-genre Telamonia (Fr.) Loud. Première partie. Trav Sci. Parc
Nat Vanoise 8:115–146
Lamoure D (1978) Agaricales de la zone alpine. Genre Cortinarius Fr.
sous-genre Telamonia (Fr.) Loud. Suite I. Trav Sci Parc Nat Vanoise
9:77–101
Lange JE (1940) Flora Agaricina Danica. 5. Copenhagen.
Liimatainen K, Carteret X, Dima B, Kytövuori I, Bidaud A, Reumaux P,
Niskanen T, Ammirati JF, Bellanger J-M (2017) Cortinarius sec-
tion Bicolores and section Saturnini (Basidiomycota, Agaricales), a
morphogenetic overview of European and North American species.
Persoonia 39:175–200
Liimatainen K, Niskanen T, Ammirati JF, Kytövuori I, Dima B (2015)
Cortinarius, section Disjungendi, cryptic species in North America
and Europe. Mycol Progress 14:1016. https ://doi.org/10.1007/s1155
7-014-1016-9
Liimatainen K, Niskanen T, Dima B, Kytövuori I, Ammirati JF, Frøslev
T (2014) The largest type study of Agaricales species to date: bring-
ing identification and nomenclature of Phlegmacium (Cortinarius,
Agaricales) into the DNA era. Persoonia 33:98–140
Liimatainen K, Niskanen T, San-Fabian B, Mujic AB, Peintner U,
Dresch P, Furci G, Nouhra E, Matheny PB, Smith ME (2020)
Cortinarius section Thaumasti in South American Nothofagaceae
forests. Mycologia 112(2):329–341. https ://doi.org/10.1080/00275
514.2019.16897 63
Lindahl B, Nilsson RH, Tedersoo L, Abarenkov K, Carlsen T, Kjøller R,
Kõljalg U, Pennanen T, Rosendahl S, Stenlid J (2013) Kauserud H
(2013) Fungal community analysis by high-throughput sequencing
of amplified markers—a user’s guide. New Phytol 199:288–299.
https ://doi.org/10.1111/nph.12243
Maddison DR, Maddison WP (2017) Chromaseq: a Mesquite package
for analyzing sequence chromatograms. Version 1.3. http://mesqu
itepr oject .org/packa ges/chrom aseq
Mahiques R, Ortega A (2002) Cortinarius erythrofuscus (subgenus Tel-
amonia, section Firmiores), a new species from Spain. Persoonia
17(4):657–660
Moser M (1983) Die Röhrlinge und Blätterpilze. In: Gams H (ed) Kleine
Kryptogamenflora, Band II b/2, 5th edn. Gustav Fischer Verlag,
Stuttgart, Germany
Niskanen T, Kytövuori I, Bendiksen E, Bendiksen K, Brandrud TE,
Frøslev TG, Høiland K, Jeppesen TS, Liimatainen K, Lindström H
(2012) Cortinarius (Pers) Gray. In: Knudsen H, Vesterholt J (eds).
Funga Nordica, 2nd revised edition. Agaricoid, boletoid, clavarioid,
cyphelloid and gastroid genera, Nordsvamp, Copenhagen, Denmark,
pp 762–763.
Niskanen T, Kytövuori I, Liimatainen K (2009) Cortinarius sect. Brun-
nei (Basidiomycota, Agaricales) in North Europe. Mycol Res
113:182–206
Niskanen T, Kytövuori I, Liimatainen K (2011) Cortinarius sect. Armil-
lati in northern Europe. Mycologia 103(5):1080–1101. https ://doi.
org/10.3852/10-350
Niskanen T, Kytövuori I, Liimatainen K, Lindström H (2013) Cortinar-
ius section Bovini (Agaricales, Basidiomycota) in northern Europe,
conifer associated species. Mycologia 105(4):977–993. https ://doi.
org/10.3852/12-320
Pearson AA (1946) New records and observations III. Trans. Br. Mycol.
Soc. 29(4):191–209
Peintner U, Moncalvo J-M, Vilgalys R (2004) Towards a better under-
standing of the infrageneric relationships in Cortinarius (Agaricales,
Basidiomycota). Mycologia 96(5):1042–1058
San-Fabian B, Niskanen T, Liimatainen K, Kooij PW, Mujic AB, Truong
C, Peintner U, Dresch P, Nouhra E, Matheny PB, Smith ME (2018)
New species of Cortinarius sect Austroamericani, sect. nov, from
South American Nothofagaceae forests. Mycologia 110(6):1127–
1144. https ://doi.org/10.1080/00275 514.2018.15154 49
Schaeffer JC (1774) Fungorum qui in Bavaria et Palatinatu circa Ratis-
bonam nascuntur Icones 4. Regensburg
Schoch CL, Robbertse B, Robert V etal (2014) Finding needles in hay-
stacks: linking scientific names, reference specimens and molecular
data for Fungi. Database 1–21. 10.1093/database/bau061
Schoch CL, Seifert KA, Huhndorf A etal (2012) Nuclear ribosomal inter-
nal transcribed spacer (ITS) region as a universal DNA barcode
marker for Fungi. Proc Natl Acad Sci USA 109:6241–6246
Simmons MP, Ochoterena H (2000) Gaps as characters in sequence-
based phylogenetic analysis. Syst Biol 49:369–381. https ://doi.
org/10.1093/sysbi o/49.2.369
Smith AH (1944) New and interesting Cortinarii from North America.
Lloydia 7(3):163–235
Soop K, Dima B, Cooper JA, Park D, Oertel B (2019) A phyloge-
netic approach to a global supraspecific taxonomy of Cortinarius
(Agaricales) with an emphasis on the southern mycota. Persoonia
42:261–290

Fungal Diversity
1 3
Stamatakis A (2014) RAxML Version 8: a tool for phylogenetic
analysis and post-analysis of large phylogenies. Bioinformatics
30:1312–1313
Stensrud Ø, Orr RJS, Reier-Røberg K, Schumacher T, Høiland K (2014)
Phylogenetic relationships in Cortinarius with focus on North Euro-
pean species. Karstenia 54:57–71
Suárez-Santiago VN, Ortega A, Peintner U, López-Flores I (2009)
Study on Cortinarius subgenus Telamonia section Hydrocybe in
Europe, with especial emphasis on Mediterranean taxa. Mycol Res
113:1070–1090
Thiers B (2013) Index Herbariorum: a global directory of public her-
baria and associated staff. New York Botanical Garden’s Virtual
Herbarium. http://sweet gum.nybg.org/ih/
Velenovský J (1939) Novitates Mycologicae. Prague, Czech
White TJ, Bruns T, Lee S, Taylor J (1990) Amplification and direct
sequencing of fungal ribosomal RNA genes for phylogenetics. In:
Michael AJ, Gelfand DH, Sninsky JJ, White TJ (eds) PCR protocols:
a guide to the methods and applications. Academic Press, New York,
pp 315–322
Willis KJ (ed) (2018) State of the world’s fungi 2018. Report. Royal
Botanic Gardens, United Kingdom.