Cortinarius section Bicolores and section Saturnini (Basidiomycota, Agaricales), a morphogenetic overview of European and North American species

Abstract

Cortinarius
is the largest genus of ectomycorrhizal fungi worldwide. Recent molecular studies have shown high levels of morphological homoplasy within the genus. Importantly, DNA phylogenies can reveal characteristics that have been either over- or underemphasized in taxonomic studies. Here we sequenced and phylogenetically analysed a large set of pan-European and North American collections taxonomically studied and placed inCortinariussect.Bicoloresand sect.Saturnini, according to traditional morpho-anatomical criteria. Our goal was to circumscribe the evolutionary boundaries of the two sections, to stabilize both the limits and nomenclature of relevant species, and to identify described taxa which, according to our current understanding, belong to other lineages. Our analysis resolves two clades: /Bicolores, including 12 species, one of which is new to science, and /Saturnini, including 6 species. Fifteen binomials, traditionally treated in these two sections based on morphology, do not belong to the above two phylogenetic clades. Instead, six of these latter are clearly placed in other clades that represent sect.Bovini, sect.Sciophylli, sect.Duraciniand sect.Brunneotincti. The presence or absence of blue pigments and the detection of specific odours emerge as clearly misleading taxonomic features, but more surprisingly, spore size and ecology can be misleading as well. A total of 63 type specimens were sequenced, 4 neotypes and 2 epitypes are proposed here, and 1 new combination is made.

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Persoonia 39, 2017: 175–200 ISSN (Online) 1878-9080
www.ingentaconnect.com/content/nhn/pimj https://doi.org/10.3767/persoonia.2017.39.08
RESEARCH ARTICLE
INTRODUCTION
Cortinarius is the largest genus of ectomycorrhizal fungi world-
wide, with no less than 4 701 reported taxa (3 360 species,
1 341 infraspecific taxa, http://www.catalogueoflife.org, 28 Sept.
2016 release). However, the number of species greatly varies
depending on the morphological species concept accepted
by classical authors. Currently, the two major monographs
dedicated to the genus are Cortinarius, Flora Photographica
(CFP), which includes ± 300 species, mostly from northern
Europe (Brandrud et al. 2014), and the Atlas des Cortinaires
(ADC), still on-going and which so far recognizes ± 2 500 spe-
cies, varieties and forms, mostly from France (Bidaud et al.
2015). Recent molecular studies have unveiled high levels of
morphological homoplasy as well as numerous cryptic species
within the genus, and as a result, do not support the broad
species concept of Scandinavian authors or the narrow one of
French authors (e.g., Liimatainen et al. 2014a). Importantly, by
identifying evolutionary units that are independent of morpho-
anatomical and ecological traits, DNA phylogenies revealed
characters that have been overemphasized in monographic
studies but also uncovered significant taxonomic information
that has been neglected by previous investigators (Bellanger et
al. 2015, Loizides et al. 2016). The use of these modern tools a
posteriori, to test the autonomy of previously defined morpho-
logical species, has been instrumental in delineating objective
boundaries to taxa, and when applied to type material, stabi-
lizes taxonomy and nomenclature at the genus level (Frøslev
et al. 2007, Liimatainen et al. 2014b, Cripps et al. 2015). The
next challenge of this nascent integrative systematics era is
undoubtedly to synchronize the two sources of knowledge, so
that on-going monographs introduce morphogenetic species,
i.e., taxa that are both assigned formal diagnosis and a unique
molecular signature.
Historically, mycologists have attempted to tackle the complexity
of Cortinarius by organizing species in hierarchical infra generic
taxa defined on supposedly stable sets of characteristics
(Kühner & Romagnesi 1953, Moser 1967, Melot 1990, Moënne-
Loccoz & Reumaux 1990). In spite of their practical application,
most of these lower level taxonomic divisions have proven to
be artificial when placed under evolutionary scrutiny (Garnica
et al. 2005). Subgenus Telamonia, however, breaks this rule
as most of the numerous species known to date that produce
dry-capped basidiomata lacking vivid colours – the morphologi-
cal definition of the subgenus and excluding a few sections as
sect. Obtusi, Balaustini, Illumini – form a strongly supported
monophyletic clade in all published molecular studies (Peintner
et al. 2004, Stensrud et al. 2014). Recently, several sections
within Telamonia have been phylogenetically revised, such as
Cortinarius section Bicolores and section Saturnini
(Basidiomycota, Agaricales), a morphogenetic overview
of European and North American species
K. Liimatainen1, X. Carteret2, B. Dima3,4, I. Kytövuori5, A. Bidaud6,
P. Reumaux7, T. Niskanen1, J.F. Ammirati8, J.-M. Bellanger9
1 Jodrell Laboratory, Royal Botanic Gardens, Kew, Surrey, TW9 3AB, United
Kingdom.
2 68, rue Alexis Maneyrol, F-92370 Chaville, France.
3 Department of Plant Anatomy, Institute of Biology, Eötvös Loránd University,
Pázmány Péter sétány 1/c, H-1117 Budapest, Hungary.
4 Department of Biosciences, Plant Biology, P.O. Box 65, FI-00014 University
of Helsinki, Finland.
5 Botanical Museum, University of Helsinki, Helsinki, P.O. Box 7, FI-00014
Finland.
6 2436, route de Brailles, F-38510 Vézeronce-Curtin, France.
7 84, avenue de Wagram, F-75017 Paris, France.
8 Department of Biology, University of Washington, Box 351800, Seattle, WA
98195-1800, USA.
9 CEFE UMR5175, CNRS, Université de Montpellier, Université Paul-Valéry
Montpellier, EPHE, INSERM, 1919, route de Mende, F-34293 Montpellier
Cedex 5, France;
corresponding author e-mail: jean-michel.bellanger@cefe.cnrs.fr.
Key words
Bicolores
Cortinarius phylogeny
integrative taxonomy
Saturnini
Telamonia
Abstract Cortinarius is the largest genus of ectomycorrhizal fungi worldwide. Recent molecular studies have shown
high levels of morphological homoplasy within the genus. Importantly, DNA phylogenies can reveal characteristics
that have been either over- or underemphasized in taxonomic studies. Here we sequenced and phylogenetically
analysed a large set of pan-European and North American collections taxonomically studied and placed in Cortinarius
sect. Bicolores and sect. Saturnini, according to traditional morpho-anatomical criteria. Our goal was to circumscribe
the evolutionary boundaries of the two sections, to stabilize both the limits and nomenclature of relevant species,
and to identify described taxa which, according to our current understanding, belong to other lineages. Our analysis
resolves two clades: /Bicolores, including 12 species, one of which is new to science, and /Saturnini, including
6 species. Fifteen binomials, traditionally treated in these two sections based on morphology, do not belong to the
above two phylogenetic clades. Instead, six of these latter are clearly placed in other clades that represent sect.
Bovini, sect. Sciophylli, sect. Duracini and sect. Brunneotincti. The presence or absence of blue pigments and the
detection of specific odours emerge as clearly misleading taxonomic features, but more surprisingly, spore size
and ecology can be misleading as well. A total of 63 type specimens were sequenced, 4 neotypes and 2 epitypes
are proposed here, and 1 new combination is made.
Article info Received: 13 October 2016; Accepted: 1 May 2017; Published: 10 August 2017.

176 Persoonia – Volume 39, 2017
sect. Armillati, Brunnei, Bovini and Disjungendi and more are on
their way to morphogenetic redefinition (Niskanen et al. 2009,
2011, 2013, Liimatainen et al. 2014a).
Here we deal with Cortinarius sect. Bicolores and Cortinarius
sect. Saturnini, which encompass Cortinarius evernius, C. sa-
turninus and their lookalikes. Initially, the two sections were
distinguished by the extent of veil remnants on the stipe, a
character considered by some authors to segregate subg.
Hydrocybe from subg. Telamonia (Moënne-Loccoz & Reumaux
1990). However, this morphological feature may not be support-
ed phylogenetically, justifying the revision of the two sections
altogether (Niskanen et al. 2012). Eight to thirty-three species
have been described in sect. Bicolores and sect. Saturnini in
the major European monographs, from the pioneering work of
Kühner & Romagnesi (1953) to the latest two releases of the
ADC (Bidaud et al. 2014, 2015), in which part of the results
presented here have been incorporated (Table 1). The specific
goals of the present work are:
1. to circumscribe the phylogenetic boundaries of the two
sections, through the analysis of a large internal tran-
scribed spacer (ITS) rDNA sequence dataset built from
pan-European and North American vouchered collections;
2. to stabilize the nomenclature and species limits of morpho-
genetic Bicolores and Saturnini, through sequencing type
material and designating neotype or epitype when oppor-
tune;
3. to assign a molecular signature to the numerous collections
taxonomically placed in these two sections in contemporary
monographs, but that do not belong in the two clades.
This study Bidaud et al. (1992, 2014, 2015) Brandrud et al. (1990, 1994, Moser (1967) Kühner & Romagnesi
1998), Niskanen et al. (2012) (1953)
Sect. Bicolores Sect. Bicolores Sect. Bicolores Key 3.11.7.6.11 Sect. Bicolores
Cortinarius cagei C. minicolor, C. cagei C. bicolor? C. bicolor?
C. periodolens ad. int.
C. dolabratoides sp. nov.
C. dolabratus C. imbutoides
C. evernius C. evernius, C. parvulior ad. int. C. evernius C. evernius, C. scutulatus C. evernius
C. glaphurus C. tubulosus, C. paranomalus (Sat.)
C. hircinosmus C. livor C. livor ?
C. plumulosus C. fundatus C. bicolor ? C. bicolor?
C. refectus C. refectus, C. testaceoviolaceus C. bicolor? C. bicolor?
C. sp1
C. sp2
C. tortuosus C. tortuosus C. tortuosus C. plumbosus C. tortuosus, C. plumbosus
C. turgidipes
C. cinnamoviolaceus C. cinnamoviolaceus, C. parevernius, C. imbutus C. cinnamoviolaceus, C. parevernius
C. basicyaneus C. parevernius
C. disjungendus C. cyanosterix
C. mattiae C. mattiae C. mattiae C. subviolascens
C. parevernioides C. parevernioides
C. salicinus C. salicinus, C. deceptivoides
C. quadricolor
Sect. Saturnini Sect. Saturnini Sect. Firmiores + sect. Telamonia Key 3.11.7.6.11 Sect. Bicolores
C. confirmatus C. confirmatus
C. cyprinus C. cyprinus
C. imbutus C. imbutus C. vilior C. imbutus
C. lucorum C. lucorum C. lucorum C. lucorum, C. umidicola
C. saturninus C. saturninus C. saturninus, C. subtorvus C. saturninus,
C. deceptivus,
C. subtorvus
C. stuntzii
C. cypriacoides C. cypriacoides C. cypriacus C. cypriacus
C. furiosus C. furiosus
C. nefastus C. nefastus
C. serratissimus* C. saturninoides C. serratissimus C. saturninus
C. sciophylloides C. sciophylloides
C. subbulliardioides* C. illepidus
C. subfirmus C. subfirmus
C. suboxytoneus C. suboxytoneus, C. fuscocinctus
C. sciophyllus C. sciophyllus
C. castaneus C. castaneus
C. calopus
C. torvus
C. impennis
C. myrtillinus
Bold names indicate sequenced species. Dotted lines separate morphogenetic species included in /Bicolores and /Saturnini (upper parts) from those (morphological species, lower parts) phylo-
genetically unrelated to the two clades. (Sat.), Saturnini. Asterisk indicates unpublished data of nomenclatural significance.
Table 1 Cortinarius species classified in sections Bicolores and Saturnini by the main European authors.

177
K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini
(text continues on p. 185)
MATERIAL AND METHODS
DNA extraction, amplification and sequencing
The material analysed in the present work was made available
to us by the public herbaria of the University of Helsinki (H,
Finland), the Muséum National d’Histoire Naturelle de Paris
(PC, France), the Swedish Museum of Natural History (S,
Sweden), the Conservatoire et Jardin botaniques de la Ville
de Genève (GK & G, Switzerland), the Universität Innsbruck
(IB, Austria), the University of Michigan (MICH, USA), and the
University of Washington (WTU, USA), as well as by European
field mycologists (Table 2). Scandinavian, North American,
and part of the French material was extracted, amplified, and
sequenced following Liimatainen et al. (2014b). DNA extraction
and PCR amplification of most of the French and south Euro-
pean material was conducted with the REDExtract-N-Amptm
Plant PCR Kit (Sigma-Aldrich, St. Louis, MO, USA), following
the manufacturer’s instructions. The internal transcribed spac-
ers and 5.8S rDNA (ITS) was amplified from each collection,
with the ITS-1F/ ITS-4b primer pair (Gardes & Bruns 1993) as
described in Richard et al. (2015). When no band was detected
by agarose-gel electrophoresis analysis, one microliter of the
PCR product was used as template in a second PCR using
the ITS1F/ ITS4 primer pair (White et al. 1990). The remaining,
most problematic extracts, were submitted to separate ITS1F/
ITS2 and ITS3/ ITS4 PCRs (White et al. 1990). Amplicons were
purified and sequenced by Eurofins Genomics, Ebersberg,
Germany. Raw sequence data were edited and assembled with
Codon Code Aligner 4.1.1 (CodonCode Corp., Centerville, MA,
USA) and deposited in GenBank under the accession numbers
indicated in Table 2.
Datasets
Out of the 348 sequences analysed in the present study, 290
(83 %) have been newly generated from vouchered material
collected and taxonomically studied by expert field mycolo-
gists, biased towards French authors. In an effort to stabilize
nomenclature, 63 sequences were obtained from type collec-
tions, which, together with 26 additional publically available
sequences, represent more than a quarter of type material
(89 out of 348) within the whole dataset. Also, to further contri-
bute to fix the usage of some well-known binomials, especially
when reference material was not available or not amenable to
successful sequencing, we included in the dataset 24 Species
Hypothesis representative sequences (‘SH repseq’) from the
UNITe database (Kõljalg et al. 2013). These phylogenetic spe-
cies can be labelled or not and their name may be misapplied,
but because they are built from sequences of wide origins,
their occurrence in a subclade often extends our knowledge
of the biogeographical distribution and sometimes the ecology,
of the corresponding species. Dataset 1 (analysed in Fig. 1)
includes 343 Telamonia sequences that belong in the /Bicolores
and /Saturnini clades as well as collections phylogenetically
or morphologically related to species traditionally treated in
the two sections, as well as 5 sequences from sect. Anomali
and subg. Phlegmacium as outgroup. We intended to define
phylogenetic boundaries and robustness of the two sections
and to reveal phylogenetically positions of species that were
formerly classified in the morphological sections Bicolores and
Saturnini, but are not part of the phylogenetic clades /Bicolores
or /Saturnini. Datasets 2 and 3 (analysed in Fig. 2 and 3, re-
spectively) focus on the species content of the revised sections
and include, respectively, 124 and 131 sequences.
Phylogenetic analyses
Phylogenetic analyses were all performed online at phylogeny.
lirmm.fr (Dereeper et al. 2008) and on the CIPRES Science
Gateway (www.phylo.org/index.php/). Multiple sequence align-
ment was carried out with MUSCLE 3.7 (Edgar 2004) using full
processing mode and 16 iterations. When required, alignments
were edited with Gblocks 0.91b, set to lowest stringency in
the selection of conserved blocks (Castresana 2000, Talavera
& Castresana 2007). Maximum likelihood (ML) phylogenetic
analyses were performed with PhyML 3.0 (Guindon et al. 2010),
using the GTR + I + Γ model of evolution. Branch support was
assessed using the non-parametric, Shimodaira-Hasegawa,
version of the approximate likelihood-ratio test (SH-aLRT),
implemented in the latest release of PhyML and which ensures
high accuracy when SH-aLRT > 0.8 (Anisimova et al. 2011,
Bellanger et al. 2015). Bayesian inference of phylogeny was
performed using MrBayes 3.1.2 (Ronquist & Huelsenbeck
2003). Two runs of four Monte Carlo Markov Chains each were
performed for 1 000 000 generations, with stationarity conver-
gence estimated by the Potential Scale Reduction Factor = 1
(Gelman & Rubin 1992). Trees and parameters were sampled
every 1 000 generations (1 000 trees). The initial burn-in was
set to 25 % (250 trees). A 50 % majority-rule consensus phy-
logram was computed from the remaining trees with Bayes-
ian posterior probabilities (BPP) reported as percentages on
supported branches of the phylograms. Trees were visualized
using FigTree 1.4.2 (http://tree.bio.ed.ac.uk/software/figtree/)
and edited with Inkscape 0.91 (https://inkscape.org/fr/).
Morpho-anatomic analyses
Microscopic characteristics were observed from dried mate-
rial mounted in Melzer’s reagent. The pileipellis structure was
studied from both freehand radial and scalp sections from the
pileus centre. The measurements of the elements of pileipellis
were made from scalps. Basidiospores were measured from
the veil or top of the stipe. Sporograms depicted in Fig. 4 have
been mounted following the method of the ADC, described in
Bidaud et al. 1994. Briefly, spores have been observed and
measured at the 1 000× magnification and 8 of them drawn
and aligned by increasing length order (0.5 µm step).
RESULTS
Our analysis resolved two strongly supported clades, referred
to as /Bicolores (BPP = 99 %, SH-aLRT = 0.92) and /Saturnini
(BPP = 100 %, SH-aLRT = 0.88) in the present work, and that
include most representative European species described in
sect. Bicolores and sect. Saturnini, respectively (Fig. 1, Table 2).
In its current sampling, /Bicolores includes 12 species, each
represented by 1 to 23 sequences (Fig. 2, Table 2). Sequenc-
ing existing type material and designating 1 neotype (C. cagei)
and 2 epitypes (C. dolabratus and C. refectus), we stabilized
9 names and identified 8 synonymous binomials at the spe-
cies rank. In addition, we describe C. dolabratoides as a new
species akin to C. dolabratus and so far found in Finland and
France. We postponed naming the North American C. sp1
and the Finnish C. sp2, awaiting further sampling to formally
describe them. Overall, our work confirms C. cagei, C. ever-
nius, C. plumulosus, C. refectus and C. tortuosus as genuine
members of the revised sect. Bicolores, but it also reveals that
C. dolabratus, C. glaphurus, C. hircinosmus and C. turgidipes,
previously reported in other sections of Telamonia, actually
belong in the section as well.
Intraspecific ITS variability in /Bicolores was generally low, with
a maximum number of changes Dintra max = 3 nucleotide (nts)
in the case of C. dolabratus, representing 0.5 % of sequence
divergence. Most species in the clade do not vary at all or only
by one substitution and one or two indels in spite of transcon-

178 Persoonia – Volume 39, 2017
/Bicolores
C. cagei CFP 1260 cagei (neotype) T.E. Brandrud, H. Lindström, 1994 Sweden CFP: D48 (1998) S KX964295
H. Marklund, S. Muskos
AB 04-09-266 minicolor A. Bidaud 2004 France AC 22: f1419 (2014) ADC private KX964296
AB 92-10-256 minicolor A. Bidaud & R. Fillion 1992 France AC 22: f1419 (2014) ADC private KX964297
PML 738 minicolor R. Fillion 1987 France AC 22: f1419 (2014) ADC private KX964298
XC 2014-02 periodolens ad int. A. Ferville 1993 France AC 22: f1417 (2014) ADC private KX964299
PML 3588 basicyaneus A. Ferville 1993 France this study ADC private KX964300
PML 1057 basicyaneus R. Fillion 1988 France this study ADC private KX964301
SH188634.07FU (2 sequences) cagei na na Germany/ Italy na na AY669676
C. dolabratoides sp. nov. H:6033567 sp. (holotype) I. Kytövuori 2008 Finland this study H KX964302
AB 07-08-48 marcellae cf. A. Bidaud & R. Fillion 2007 France this study ADC private KX964303
H:6033615 sp. I. Kytövuori 2004 Finland this study H KX964304
H:6033575 sp. I. Kytövuori 2008 Finland this study H KX964305
H:6033570 sp. I. Kytövuori 2008 Finland this study H KX964306
IK 04-051 ‘smell-of-viola’ I. Kytövuori 2004 Finland this study H KX964307
IK 01-062 ‘smell-of-viola’ I. Kytövuori 2001 Finland this study H KX964308
C. dolabratus CFP 990 dolabratus (epitype) T.E. Brandrud, H. Lindström, 1990 Sweden CFP: D52 (1998) S KX964309
H. Marklund, S. Muskos
AB 04-09-186 imbutoides (holotype) A. Bidaud 2004 France AC 22: f1409 (2014) PC KX964310
RH 80814 phaeoruber (holotype) G. Chevassut 1980 France DM 12(47): 52 (1982) PC KX964311
AB 13-10-120 saturninus cf. A. Bidaud 2013 France this study ADC private KX964312
AB 04-09-169 armillariellus cf. A. Bidaud 2004 France this study ADC private KX964313
AB 01-09-41 privignus sensu Quélet cf. A. Bidaud 2001 France this study ADC private KX964314
AB 98-09-94 saturninus cf. A. Faurite 1998 Canada this study ADC private KX964315
AB 89-11-309 orastriatus A. Bidaud 1989 France this study ADC private KX964316
H:6033519 dolabratus I. Kytövuori 2001 Finland this study H KX964317
IK 02-033 dolabratus I. Kytövuori 2002 Finland this study H KX964318
IK 95-1576 dolabratus I. Kytövuori 1995 Finland this study H KX964319
IK 95-347 dolabratus I. Kytövuori 1995 Finland this study H KX964320
KS CO1576 imbutoides K. Soop 2005 Sweden this study K. Soop private KX964321
KS CO1290 imbutoides K. Soop 2001 Sweden this study K. Soop private KX964322
TN 12-200 dolabratus T. Niskanen 2012 USA this study H KX964323
TN 11-246 dolabratus T. Niskanen 2011 USA this study H KX964324
TN 09-196 dolabratus T. Niskanen 2009 USA this study H KX964325
TN 09-139 dolabratus T. Niskanen 2009 USA this study H KX964326
TN 03-1713 dolabratus T. Niskanen 2003 Slovakia this study H KX964327
TN 02-1095 dolabratus T. Niskanen 2002 Finland this study H KX964328
TN 02-959 dolabratus T. Niskanen 2002 Finland this study H KX964329
XC 2013-103 privignus sensu Quélet P. Reumaux 1998 France this study ADC private KX964330
SH188528.07FU (10 sequences) dolabratus na na NA/ FS/Slovakia na na UDB018659
C. evernius CFP 792 evernius (neotype) T.E. Brandrud, H. Lindström, 1988 Sweden CFP: A11 (1990) S KX964331
H. Marklund, S. Muskos
AB 00-09-83 evernius f. pseudoscutulatus (holotype) A. Bidaud 2000 France AC 22: f1407 (2014) PC KX964332
PML 1727 evernius f. fragrans (holotype) D. Mazuir 1990 France AC 22: f1406 (2014) PC KX964333
AB 96-09-47 parvulior ad int. M. Martin 1996 France AC 22: f1418 (2014) ADC private KX964334
AB 91-08-42 evernius f. pseudoscutulatus A. Bidaud & C. Blanc 1991 France AC 22: f1407 (2014) ADC private KX964335
PML 622 evernius f. fragrans P. Moënne-Loccoz 1987 France AC 22: f1406 (2014) ADC private KX964336
AB 09-07-44 evernius var. insignis A. Bidaud & A. Faurite 2009 France AC 22: f1405 (2014) ADC private KX964337
PML 3469 evernius var. evernius A. Bidaud 1993 France AC 22: f1404 (2014) ADC private KX964338
Table 2 Specimens included in phylogenetic analyses.
Species Voucher/SH Voucher/SH annotation Leg. Collection Country Taxonomy Herbarium Accession*
C. = Cortinarius date

179
K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini
PML 212 evernius var. evernius P. Moënne-Loccoz 1982 France AC 22: f1404 (2014) ADC private KX964339
AB 04-09-212 evernius A. Bidaud 2004 France this study ADC private KX964340
IK 00-038 evernius I. Kytövuori 2000 Finland this study H KX964341
IK 97-123 evernius I. Kytövuori 1997 Finland this study H KX964342
PML 376 evernius P. Moënne-Loccoz 1986 France this study ADC private KX964343
PML 230 evernius P. Moënne-Loccoz 1984 France this study ADC private KX964344
TN 10-074 evernius T. Niskanen 2010 Canada this study H KX964345
TN 10-055 evernius T. Niskanen 2010 Canada this study H KX964346
TN 10-054 evernius T. Niskanen 2010 Canada this study H KX964347
TN 07-328 evernius T. Niskanen 2007 Canada this study H KX964348
TN 07-312 evernius T. Niskanen 2007 Canada this study H KX964349
TN 07-223 evernius T. Niskanen 2007 Canada this study H KX964350
TN 05-238 evernius T. Niskanen 2005 Norway this study H KX964351
SH188514.07FU (11 sequences) evernius na na Canada / FS/ Germany na na AY669686
C. glaphurus RH 71421 glaphurus (holotype) G. Chevassut 1978 France DM 12(47): 78 (1982) PC KX964352
AB 03-11-87 tubulosus (holotype) A. Bidaud & A. Faurite 2003 France AC 22: f1414 (2014) PC KX964353
AB 92-10-350 cedriosmus (holotype) A. Bidaud 1992 France AC 19: f1144 (2010) PC KX964354
XC 2009-41 violaeolens (holotype) A. & R. Bardet 1996 France AC 19: f1112 (2010) PC KX964355
GK1142 paranomalus (holotype) G. Redeuilh 1987 France K&R: 305 (1953, invalid), GK KX964356
AC 4: f163 (1992)
AB 08-11-445 tubulosus M. Martin 2008 France AC 22: f1414 (2014) ADC private KX964357
AB 92-10-332 turibulosus R. Fillion 1992 France AC 19: f1108 (2010) ADC private KX964358
AB 91-11-360 turibulosus A. Bidaud 1991 France AC 19: f1108 (2010) ADC private KX964359
PML 2390 turibulosus P. Moënne-Loccoz 1991 France AC 19: f1108 (2010) ADC private KX964360
PML 1067 turibulosus P. Moënne-Loccoz 1988 France AC 19: f1108 (2010) ADC private KX964361
AB 14-11-138 minicolor cf. P.-Y. Courio 2014 France this study ADC private KX964362
AB 03-10-56 sciophyllus cf. A. Bidaud 2003 France this study ADC private KX964363
AB 99-11-345 livor cf. M. Martin 1999 France this study ADC private KX964364
TN 12-221 sp. T. Niskanen 2012 USA this study H KX964365
XC 2011-212 laetior cf. X. Carteret 2011 France this study ADC private KX964366
XC 2009-64 paranomalus cf. X. Carteret 2009 France this study ADC private KX964367
SH094444.07FU (3 sequences) turibulosus na na NA/ France na na GQ159774
SH094485.07FU (2 sequences) sp. na na Poland na na HQ115588
C. hircinosmus PML 334 hircinosmus (holotype) P. Moënne-Loccoz 1986 France AC 12: f575 (2002) PC KX964368
AB 02-09-32 livor A. Bidaud 2002 France AC 23: f1459 (2015) ADC private KX964369
F44390 sp. K. Soop na Sweden FN: 849 (2012) S KX964370
H:6033565 hircinosmus I. Kytövuori 2009 Finland FN: 849 (2012) H KX964371
AB 97-10-341 scriptor G. Chamonaz 1997 France AC 19: f1109 (2010) ADC private KX964372
AB 04-10-357 imbutus cf. A. Bidaud 2004 France this study ADC private KX964373
C. plumulosus RH 3417 plumulosus (holotype) R. Henry 1972 France SMF 93(3): 359 (1977) PC KX964374
AB 10-09-183 fundatus A. Bidaud & R. Fillion 2010 France AC 22: f1411 (2014) ADC private KX964375
AB 98-09-119 fundatus E. & A. Bidaud, A. Faurite 1998 Canada AC 22: f1411 (2014) ADC private KX964376
PML 657 fundatus P. Moënne-Loccoz 1987 France AC 22: f1411 (2014) ADC private KX964377
PML 3308 perscutulatus A. Bidaud 1992 France this study ADC private KX964378
IK 98-1612 sp. I. Kytövuori 1998 Finland this study H KX964379
TN 04-730 sp. T. Niskanen 2004 Finland this study H KX964380
C. refectus AB 96-09-73 refectus (epitype) A. Bidaud 1996 Germany AC 22: f1410 (2014) PC KX964385
AB 05-09-138 refectus A. Bidaud 2005 France AC 22: f1410 (2014) ADC private KX964382
AB 04-10-321 refectus A. Bidaud 2004 France AC 22: f1410 (2014) ADC private KX964383
AB 99-09-121 refectus A. Bidaud 1999 France AC 22: f1410 (2014) ADC private KX964384
PML 2159 refectus A. Bidaud 1990 France AC 22: f1410 (2014) ADC private KX964386
PML 769 refectus P. Moënne-Loccoz 1987 France AC 22: f1410 (2014) ADC private KX964387
PML 17 refectus P. Moënne-Loccoz 1985 France AC 22: f1410 (2014) ADC private KX964388
AB 92-10-293 testaceoviolaceus A. Bidaud 1992 France AC 22: f1402 (2014) ADC private KX964389
AB 94-10-268 scriptor A. Bidaud 1994 France AC 19: f1109 (2010) ADC private KX964390
IK 96-1031 refectus I. Kytövuori 1996 Germany this study H KX964381

180 Persoonia – Volume 39, 2017
C. tortuosus IB 79/533 tortuosus (neotype) D. Lamoure 1979 Sweden Opera Botanica 100: 182 IB KX964391
(1989)
XC 2008-43 flabelloides (holotype) M. Pèlerin 2008 France AC 19: f1136 (2010) PC KX964392
PAK 354 laetior (holotype) P.A. Karsten 1879 Finland BFNF 32: 387 (1879) H KX964393
AB 01-09-19 tortuosus A. Bidaud 2001 France AC 22: f1413 (2014) ADC private KX964394
AB 96-08-19 tortuosus A. Bidaud 1996 France AC 22: f1413 (2014) ADC private KX964395
AB 95-09-34 tortuosus C. Blanc 1995 France AC 22: f1413 (2014) ADC private KX964396
PML 3551 tortuosus A. Bidaud & R. Fillion 1993 France AC 22: f1413 (2014) ADC private KX964397
PML 1225 tortuosus P. Moënne-Loccoz 1989 France AC 22: f1413 (2014) ADC private KX964398
PML 1214 tortuosus P. Moënne-Loccoz 1989 France AC 22: f1413 (2014) ADC private KX964399
PML 386 tortuosus P. Moënne-Loccoz 1986 France AC 22: f1413 (2014) ADC private KX964400
CFP 493 tortuosus T.E. Brandrud, H. Lindström, 1986 Norway CFP: A06 (1990) S KX964401
H. Marklund, S. Muskos
AB 02-09-41 saturninus cf. A. Bidaud 2002 France this study ADC private KX964402
AB 96-10-124 saturninus cf. C. Blanc 1996 France this study ADC private KX964403
IK 99-709 tortuosus I. Kytövuori 1999 Finland this study H KX964404
TN 10-087 tortuosus T. Niskanen 2010 Canada this study H KX964405
TN 09-046 tortuosus T. Niskanen 2009 USA this study H KX964406
TN 07-307 tortuosus T. Niskanen 2007 Canada this study H KX964407
TN 05-006 tortuosus T. Niskanen 2005 Finland this study H KX964408
SH094369.07FU (7 sequences) tortuosus na na USA /U /Japan na na AY669669
C. turgidipes AB 93-10-425 turgidipes (holotype) A. & E. Bidaud 1993 France AC 17(1): f885 (2008) PC KX964409
C. sp1 TN 12-217 sp. T. Niskanen 2012 USA na H KX964410
UBCOGTR194 sp. (ectomycorrhiza) na na Canada na na EU597034
C. sp2 TN 05-033 sp. T. Niskanen 2005 Finland na H KX964411
other (morphological) Bicolores
C. cinnamoviolaceus IB 48/590 cinnamoviolaceus (holotype) M. Moser 1948 Austria Nova Hedwigia XIV(2-4): IB KX964412
514 (1967)
RH 70942 basicyaneus (holotype) M. Trimbach 1976 France FAMM 25: 38 (2004) PC KX964413
RH 4000 cylindratus (holotype) R. Henry 1972 France SMF 99(1): 91 (1983) PC KX964414
RH 526 subparevernius (holotype) R. Henry 1956 France SMF 85(4): 442 (1969) PC KX964415
RH 1240 contractus (holotype) R. Henry 1960 France SMF 85(4): 387 (1969) PC KX964416
RH 3258a78 parevernius (holotype) R. Henry 1955 France K&R: 303 (1953, invalid) PC KX964417
AB 02-10-71 dolabratus A. & M. Burat 2002 France AC 17(1): f817 (2008) ADC private KX964418
CFP 574 imbutus T.E. Brandrud, H. Lindström, 1987 Sweden CFP: D60 (1998) S KX964419
H. Marklund, S. Muskos
AB 12-11-240 imbutus A. Bidaud 2012 France this study ADC private KX964420
TN 05-198 imbutus sensu Funga Nordica T. Niskanen 2005 Finland this study H KX964421
TN 05-051 imbutus sensu Funga Nordica T. Niskanen 2005 Finland this study H KX964422
SH188640.07FU (2 sequences) imbutus na na Sweden/Italy na na UDB001160
C. cyanosterix RH 338 cyanosterix (holotype) R. Henry 1952 France SMF 71(3): 259, 261 (1956) PC KX964423
(= C. disjungendus)
C. mattiae KS CO1936 mattiae (isotype) K. Soop 2009 Sweden JEC 13(12): 3 (2010) S KX964424
AB 13-08-35 mattiae A. Bidaud, F. Armada & R. Fillion 2013 France AC 22: f1415 (2014) ADC private KX964425
AB 99-09-77 subviolascens A. Bidaud 1999 France AC 12: f565 (2002) ADC private KX964426
PML 650 subviolascens P. Moënne-Loccoz 1987 France AC 12: f565 (2002) ADC private KX964427
CFP 1204 mattiae T.E. Brandrud, H. Lindström, 1993 Sweden CFP: D30 (1998) S KX964428
H. Marklund, S. Muskos
AB 06-09-153 licinipes/ poecilopus aff. A. Bidaud, F. Armada & R. Fillion 2006 France this study ADC private KX964429
H:6029375 mattiae T. Niskanen 2004 Finland this study H KX964430
Table 2 (cont.)
Species Voucher/SH Voucher/SH annotation Leg. Collection Country Taxonomy Herbarium Accession*
C. = Cortinarius date

181
K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini
H:6000560 mattiae I. Kytövuori 2007 Finland this study H KX964431
IK 01-039 mattiae I. Kytövuori 2001 Sweden this study H KX964432
IK 98-1127 mattiae I. Kytövuori 1998 Finland this study H KX964433
PML 3989 umbrinoconnatus forma A. Bidaud 1993 France this study ADC private KX964434
PML 2298 oxytoneus A. Bidaud 1991 France this study ADC private KX964435
SH009438.07FU (1 sequence) sp. na na Canada na na FJ039684
C. parevernioides AB 02-09-50 parevernioides (holotype) C. Gérard 2002 France AC 22: f1408 (2014) PC KX964436
SH188502.07FU (15 sequences) malachius na na NA/U na na KF617653
C. salicinus XC 2014-03 salicinus (holotype) C. Hugouvieux 2005 France AC 22: f1416 (2014) PC KX964437
/saturnini
C. confirmatus RH 3195 confirmatus (holotype) R. Henry 1970 France SMF 99(1): 67 (1983) PC KX964438
JVG 990125-31 assiduus var. plesiocistus (isotype) X. Llimona & J. Vila 1999 Spain Mycotaxon 101: 140 (2007) J. Vila private AM713178
MES 3541 assiduus (holotype) R. Mahiques 1999 Spain FMDS 162: 42 (2001) MES KX964439
RH 84/159 bulbosovolvatus (isotype) M. Contu & L. Curreli 1984 Italy DM 26 (61): 32 (1985) PC KX964440
XC 2013-160 confirmatus ‘asp. subcylindratus’ na 2013 France AC 23: f1441 (2015) PC KX964441
AB 13-10-97 confirmatus ‘asp. kuehneri’ A. Bidaud 2013 France AC 23: f1440 (2015) ADC private KX964442
XC 2011-199 confirmatus ‘asp. spurcatocephalus’ X. Carteret 2011 France AC 23: f1439 (2015) ADC private KX964443
XC 95-10-04-06 confirmatus ‘asp. spurcatocephalus’ X. Carteret 1995 France AC 23: f1439 (2015) ADC private KX964444
AB 09-11-452 confirmatus ‘asp. rubricosissimus’ A. Bidaud 2009 France AC 23: f1438 (2015) ADC private KX964445
AB 00-10-193 confirmatus ‘asp. rubricosissimus’ A. Bidaud 2000 France AC 23: f1438 (2015) ADC private KX964446
AB 11-11-324 confirmatus ‘asp. paracohabitans’ F. Armada, A. Bidaud & J. Pardo 2011 France AC 23: f1437 (2015) ADC private KX964447
PML 4722 confirmatus ‘asp. imbutus’ P. Reumaux 1990 France AC 23: f1436 (2015) ADC private KX964448
XC 2012-171 confirmatus ‘asp. imbutus’ A. Lantz 2012 France AC 23: f1436 (2015) ADC private KX964449
AB 09-11-514 confirmatus ‘asp. assiduus’ A. Bidaud 2009 France AC 23: f1435 (2015) ADC private KX964450
AB 05-11-423 confirmatus ‘asp. assiduus’ A. & E. Bidaud 2005 France AC 23: f1435 (2015) ADC private KX964451
AB 02-11-201 confirmatus ‘asp. assiduus’ F. Lopez 2002 France AC 23: f1435 (2015) ADC private KX964452
XC 2013-156 confirmatus ‘asp. assiduus’ F. Valade 2013 France AC 23: f1435 (2015) ADC private KX964453
AB 03-11-78 confirmatus ‘asp. confirmatus’ A. Faurite 2003 France AC 23: f1434 (2015) ADC private KX964454
AB 92-11-422 cistoadelphus ad int. A. Bidaud 1992 France FAMM 6: 41 (1994) ADC private KX964455
AB 09-11-450 cohabitans cf. A. Bidaud 2009 France this study ADC private KX964456
FR2016052 assiduus J.-M. Ourcival 2016 France this study CEFE private KX964457
FR2012405 assiduus P.-A. Moreau 2012 France this study CEFE private KX964458
FR2012089 assiduus F. Richard 2011 France this study CEFE private KX964459
FR2012076 assiduus E. Taschen 2011 France this study CEFE private KX964460
XC 2006-204 bresadolae cf. na 2006 France this study ADC private KX964461
XC 2005-249 saturninus cf. X. Carteret 2005 France this study ADC private KX964462
SH094374.07FU (6 sequences) sp. na na U/Iran na na HQ204652
C. cyprinus XC 2012-26 cyprinus (holotype) G. Redeuilh 1993 France AC 23: f1443 (2015) PC KX964463
AB 11-11-251 cyprinus A. Bidaud 2011 France AC 23: f1443 (2015) ADC private KX964464
AB 11-10-192 cyprinus A. Bidaud 2011 France AC 23: f1443 (2015) ADC private KX964465
AB 06-09-144 cyprinus A. Bidaud 2006 France AC 23: f1443 (2015) ADC private KX964466
PML 344 cyprinus P. Moënne-Loccoz 1986 France AC 23: f1443 (2015) ADC private KX964467
PML 81 cyprinus P. Moënne-Loccoz 1981 France AC 23: f1443 (2015) ADC private KX964468
XC 2013-15 cyprinus P. Reumaux 2013 France AC 23: f1443 (2015) ADC private KX964469
XC 2007-103 cyprinus X. Carteret 2007 France AC 23: f1443 (2015) ADC private KX964470
AB 04-09-167 sciophyllus cf. A. Bidaud 2004 France this study ADC private KX964471
JMB 2014111802 circumvelatus cf. P.-A. Moreau 2014 France this study CEFE private KX964472
PAM 13092901 circumvelatus P.-A. Moreau 2013 France this study CEFE private KX964473
PML 425 myrtillinus P. Moënne-Loccoz 1986 France this study ADC private KX964474
XC 2007-95 mutabilis cf. na 2007 France this study ADC private KX964475
TEB 348-10 saturninus aff. T.E. Brandrud na Norway this study na KX964476
TAAM 128765 sp. A. Kollom 2008 Estonia na na UDB016164
C. imbutus IK 97-1162 imbutus (neotype) I. Kytövuori 1997 Finland this study H KX964498
PML 4557 laccatus (holotype) P. Reumaux 1978 France SMF 98(4): 348 (1982) PC KX964478
RH 3123 betulaecomes (holotype) R. Henry 1976 France SMF 93(3): 347 (1977) PC KX964479

182 Persoonia – Volume 39, 2017
XC 2013-13 imbutus ‘asp. laetior’ P. Reumaux 1998 France AC 23: f1447 (2015) ADC private KX964480
XC 2014-77 imbutus ‘asp. saturnalis’ P. Reumaux 1978 France AC 23: f1446 (2015) ADC private KX964481
XC 2014-61 imbutus ‘asp. saturnalis’ P. Reumaux 1986 France AC 23: f1446 (2015) ADC private KX964482
XC 2007-104 imbutus ‘asp. vilior’ X. Carteret 2007 France AC 23: f1445 (2015) ADC private KX964483
AB 10-10-237 imbutus ‘asp. imbutus’ A. Bidaud 2010 France AC 23: f1444 (2015) ADC private KX964484
AB 09-11-471 imbutus ‘asp. imbutus’ A. Bidaud & R. Fillion 2009 France AC 23: f1444 (2015) ADC private KX964485
AB 04-09-228 imbutus ‘asp. imbutus’ A. Bidaud & A. Faurite 2004 France AC 23: f1444 (2015) ADC private KX964486
AB 98-10-358 imbutus ‘asp. imbutus’ A. Bidaud 1998 France AC 23: f1444 (2015) ADC private KX964487
PML 375 imbutus ‘asp. imbutus’ P. Reumaux 1986 France AC 23: f1444 (2015) ADC private KX964488
XC 2002-122 imbutus ‘asp. imbutus’ X. Carteret 2002 France AC 23: f1444 (2015) ADC private KX964489
XC 2002-108 imbutus ‘asp. imbutus’ X. Carteret 2002 France AC 23: f1444 (2015) ADC private KX964490
XC 2002-107 imbutus ‘asp. imbutus’ X. Carteret 2002 France AC 23: f1444 (2015) ADC private KX964491
XC 2002-106 imbutus ‘asp. imbutus’ X. Carteret 2002 France AC 23: f1444 (2015) ADC private KX964492
AB 08-10-307 cohabitans J. Garin 2008 France this study ADC private KX964493
AB 02-10-106 cohabitans M. Renard 2002 France this study ADC private KX964494
AB 02-09-58 cohabitans A. Bidaud 2002 France this study ADC private KX964495
AB 00-09-127 cohabitans cf. A. Bidaud 2000 France this study ADC private KX964496
IK 98-2242 sp. I. Kytövuori 1998 Sweden this study H KX964497
IK 94-1236 sp. I. Kytövuori 1994 Finland this study H KX964477
JMB 2008092703 salicis cf. J.-M. Bellanger 2008 France this study CEFE private KX964499
RH 71030 betulaecomes R. Henry 1976 France this study (Rob. Henry, ined.) PC KX964500
TN 11-257 sp. T. Niskanen 2011 USA this study H KX964501
TN 11-252 sp. T. Niskanen 2011 USA this study H KX964502
TN 11-151 sp. T. Niskanen 2011 USA this study H KX964503
TN 11-150 sp. T. Niskanen 2011 USA this study H KX964504
TN 05-167 sp. T. Niskanen 2005 Finland this study H KX964505
XC 2012-96 laetior forma X. Carteret 2012 France this study ADC private KX964506
XC 2002-109 renidentoides cf. X. Carteret 2002 France this study ADC private KX964507
SH188563.07FU (6 sequences) saturninus na na Canada /Estonia /China na na UDB018346
C. lucorum CFP 490 lucorum (neotype) T.E. Brandrud, H. Lindström, 1986 Norway CFP: C10 (1994) S KX964585
H. Marklund, S. Muskos
RH 71502 incarnatolilascens (holotype) R. Henry 1979 France AC 23: f1431 (2015), PC KX964508
SMF 97(3): 170 (1981)
PML 4142 montis-dei (holotype) P. Reumaux 1980 France AC 23: f1430 (2015), PC KX964509
SMF 96(3): 357 (1980)
PML 34 circumvelatus (holotype) P. Reumaux 1976 France AC 23: f1429 (2015), PC KX964510
SMF 96(3): 355 (1980)
10433 umidicola (syntype) C.H. Kauffman 1903 USA Bull. Torrey Bot. Club 32(6): MICH KX964511
322 (1905)
PML 4143 lucorum ‘asp. montis-dei’ P. Reumaux 1980 France AC 23: f1430 (2015) ADC private KX964512
PAM 14090808 lucorum ‘asp. circumvelatus’ P.-A. Moreau 2014 France AC 23: f1429 (2015) ADC private KX964513
IK 89-748 lucorum I. Kytövuori 1989 Finland this study H KX964514
KS CO513 diabolicus K. Soop na Sweden this study na KX964515
TN 10-002 lucorum T. Niskanen 2010 Canada this study H KX964516
TN 03-1169 lucorum T. Niskanen 2003 Sweden this study H KX964517
SH188495.07FU (21 sequences) lucorum na na NA / FS na na UDB019872
C. saturninus CFP 514 saturninus (neotype) T.E. Brandrud, H. Lindström, 1986 Sweden CFP: C09 (1994) S KX964584
H. Marklund, S. Muskos
PML 4578 urbicus var. sporanotandus (holotype) A. Bidaud 1996 France AC 23: f1455 (2015), PC KX964518
AC 12: f560 (2002)
Table 2 (cont.)
Species Voucher/SH Voucher/SH annotation Leg. Collection Country Taxonomy Herbarium Accession*
C. = Cortinarius date

183
K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini
PML 2215 marginatosplendens (isotype) P. Reumaux 1978 France AC 23: f1453 (2015), G KX964519
SMF 96(3): 356 (1980)
XC 2007-14 fulvorimosus (holotype) A. & R. Bardet 1992 France AC 23: f1452 (2015), PC KX964520
AC 17: f869 (2008)
RH 3758 denseconnatus (holotype) na 1973 France SMF 99(1): 65 (1983) PC KX964521
RH 81181 gramineus (holotype) R. Henry 1981 France SMF 99(1): 64 (1983) PC KX964522
RH 71682 rastetteri (holotype) V. Rastetter 1980 France SMF 97(3): 177 (1981) PC KX964523
PR 258 dissidens (holotype) P. Reumaux 1978 France SMF 96(3): 370 (1980) PC KX964524
RH 2623 salicis (holotype) R. Henry 1968 France SMF 93(3): 364 (1977) PC KX964525
RH 476 umbrinoconnatus (holotype) R. Henry 1955 France SMF 73(1): 53 (1957) PC KX964526
AB 02-10-179 saturninus ‘asp. urbicoides’ A. Faurite 2002 France AC 23: f1457 (2015) ADC private KX964527
AB 95-11-144 saturninus ‘asp. urbicoides’ R. Fillion 1995 France AC 23: f1457 (2015) ADC private KX964528
XC 2001-107 saturninus ‘asp. urbicoides’ X. Carteret & P. Reumaux 2001 France AC 23: f1457 (2015) ADC private KX964529
AB 14-11-160 (= AB 14-11-161) saturninus ‘asp. salicis’ A. Bidaud, J. Cavet, 2014 France AC 23: f1454 (2015) ADC private KX964530
R. Fillion & G. Raffini
XC 2014-109 saturninus ‘asp. salicis’ X. Carteret 2014 France AC 23: f1454 (2015) ADC private KX964531
XC 2011-205 saturninus ‘asp. salicis’ X. Carteret 2011 France AC 23: f1454 (2015) ADC private KX964532
XC 2008-55 saturninus ‘asp. salicis’ X. Carteret 2008 France AC 23: f1454 (2015) ADC private KX964533
XC 2007-108 saturninus ‘asp. salicis’ X. Carteret 2007 France AC 23: f1454 (2015) ADC private KX964534
AB 14-09-47 saturninus ‘asp. dionisiae’ E. Bidaud 2014 France AC 23: f1451 (2015) ADC private KX964535
AB 04-10-344 saturninus ‘asp. deceptivus’ A. Bidaud 2004 France AC 23: f1450 (2015) ADC private KX964536
AB 98-10-381 saturninus ‘asp. deceptivus’ Dr. Misermont 1998 France AC 23: f1450 (2015) ADC private KX964537
XC 2014-63 saturninus ‘asp. cohabitans’ M. Pèlerin 1996 France AC 23: f1449 (2015) ADC private KX964538
XC 2014-116 saturninus ‘asp. saturninus’ na 2014 France AC 23: f1448 (2015) ADC private KX964539
XC 2014-114 saturninus ‘asp. saturninus’ L. Tarahu 2014 France AC 23: f1448 (2015) ADC private KX964540
XC 2007-97 saturninus ‘asp. saturninus’ na 2007 France AC 23: f1448 (2015) ADC private KX964541
AB 97-09-187, PML 5347 urbicus E. & A. Bidaud 1997 France AC 12: f560 (2002) ADC private KX964542
PML 3967 salicis var. salicis M. Citérin 1994 France AC 12: f559 (2002) ADC private KX964543
CFP 408 subtorvus T.E. Brandrud, H. Lindström, 1986 Sweden CFP: A04 (1990) S KX964544
H. Marklund, S. Muskos
AB 05-10-273 deceptivus sensu Moser R. Fillion 2005 France this study ADC private KX964545
H:6029320 saturninus I. Kytövuori 1998 Finland this study H KX964546
IK 94-631 saturninus I. Kytövuori 1994 Finland this study H KX964547
JMB 2009101002 cohabitans J.-M. Bellanger 2009 France this study CEFE private KX964548
KH14 subtorvus na 2011 Norway (Svalbard) na na GU234058
O50591 subtorvus na 2011 Norway (Svalbard) na na GU234013
PML 75 urbicus P. Moënne-Loccoz 1984 France this study ADC private KX964549
TN 09-208 saturninus T. Niskanen 2009 USA this study H KX964550
XC 2016-12 euprivignus aff. P. Reumaux 1977 France this study ADC private KX964551
XC 2008-61 salicis X. Carteret 2008 France this study ADC private KX964552
XC 2007-90 mutabilis cf. X. Carteret 2007 France this study ADC private KX964553
XC 2006-194 salicis X. Carteret 2006 France this study ADC private KX964554
XC 2002-167 holophaeus sensu Henry M. Pèlerin 2002 France this study ADC private KX964555
XC 2001-104 mutabilis X. Carteret 2001 France this study ADC private KX964556
XC 96-10-26-09 subprivignus X. Carteret 1996 France this study ADC private KX964557
SH094324.07FU (13 sequences) saturninus na na USA/U na na UDB017613
C. stuntzii Rehner 394 stuntzii (holotype) S.A. Rehner 1981 USA Mycologia 80(6): 903 (1988) WTU KX964558
other (morphological) saturnini
C. cypriacoides PML 1269 cypriacoides ‘asp. cypriacoides’ R. Fillion 1989 France AC 23: f1423 (2015), PC KX964559
(holotype) AC 2: f81 (1990)
PML 3984 cypriacoides ‘asp. cypriacoides’ C. Guyot 1989 France AC 23: f1423 (2015) ADC private KX964560
PML 3979 cypriacoides ‘asp. lucorum’ A. Bidaud 1992 France AC 23: f1424 (2015), ADC private KX964561
AC 9: f419 (1999)
C. furiosus XC 2014-64c furiosus (holotype) D. Brion 2012 France AC 23: f1458 (2015) PC KX964562
LM5411 sp. (Quercus ectomycorrhiza) na na Austria na na KM576363

184 Persoonia – Volume 39, 2017
C. illepidus sensu ADC AB 11-11-331 illepidus A. Bidaud & C. Gérard 2011 France AC 23: f1422 (2015) ADC private KX964563
(= C. subbulliardioides)
AB 11-11-330 illepidus A. Bidaud & C. Gérard 2011 France AC 23: f1422 (2015) ADC private KX964564
C. nefastus XC 2014-60 nefastus (holotype) D. Brion 2012 France AC 23: f1426 (2015) PC KX964565
C. ortovernus JB 604808 ortovernus (holotype) J. Ballará 2008 Spain JEC 12(11): 56 (2009) J. Ballara private KX964566
C. oxytoneus RH 931 oxytoneus (holotype) R. Henry 1957 France SMF 97(3): 277 (1981) PC KX964567
C. saturninoides sensu AB 12-10-93 saturninoides A. Bidaud & M. Renard 2012 France AC 23: f1421 (2015) ADC private KX964568
ADC (= C. serratissimus) AB 00-10-148 saturninoides A. Bidaud 2000 France AC 23: f1421 (2015) ADC private KX964569
RH 3451 oxytoneus R. Henry 1972 France SMF 97(3): 277 (1981) ADC private KX964570
XC 2014-119 saturninoides R. Chalange 2014 France AC 23: f1421 (2015) ADC private KX964571
XC 2014-64b saturninoides D. Brion 2012 France AC 23: f1421 (2015) ADC private KX964572
XC 2013-144 saturninoides F. Valade 2013 France AC 23: f1421 (2015) ADC private KX964573
XC 2010-56 saturninoides X. Carteret 2010 France AC 23: f1421 (2015) ADC private KX964574
XC 2010-29 saturninoides X. Carteret 2010 France AC 23: f1421 (2015) ADC private KX964575
SH188624.07FU (3 sequences) lucorum na na USA /Estonia/ Italy na na UDB016052
C. sciophylloides AB 99-10-254 sciophylloides (holotype) A. Bidaud 1999 France AC 23: f1425 (2015) PC KX964576
AB 91-10-291 sciophylloides J. Garin 1991 France AC 23: f1425 (2015) ADC private KX964577
PML 5446 sciophylloides J. Cavet 1999 France AC 23: f1425 (2015) ADC private KX964578
PML 2381 raphanodiabolicus P. Reumaux 1991 France na ADC private KX964579
SH188568.07FU (6 sequences) valgus na na Canada/U na na UDB002444
C. subfirmus AB 08-10-363 subfirmus (holotype) A. Bidaud & G. Raffini 2008 France AC 23: f1433 (2015) PC KX964580
C. suboxytoneus AB 01-09-56 suboxytoneus (holotype) A. Bidaud 2001 France AC 23: f1442 (2015) PC KX964581
MFT60 sp. (Fagus ectomycorrhiza) na na Germany na na FJ403502
other telamonia
C. alboviolaceus s.lat. SH188487.07FU (26 sequences) alboviolaceus na na NA/ U na na AF325596
C. anisatus CFP 1200 anisatus (holotype) T.E. Brandrud, H. Lindström, 1993 Sweden CFP: E25 (2014) S DQ117931
H. Marklund, S. Muskos
C. anisochrous IK 01-030 anisochrous (holotype) T. Niskanen & I. Kytövuori 2001 Estonia Mycologia 105(4): 988 (2013) H, S, NY JX407297
C. athabascus DBB27618, UC1860905 athabascus (holotype) D. Bojantchev 2011 USA Mycotaxon 123: 382 (2013) UC JN133295
C. biformis SH188479.07FU (41 sequences) biformis na na NA/ U na na UDB002252
C. bovinus IK 04-038 bovinus (neotype) I. Kytövuori 2004 Finland Mycologia 105(4): 981 (2013) H, S, NY JX407276
C. brunneifolius TN 06-146 brunneifolius (holotype) T. Niskanen 2006 Finland Mycol. Progress 7(4): H EU259284
241 (2008)
C. caesioarmeniacus H:7000901 caesioarmeniacus (holotype) K. Liimatainen & T. Niskanen 2007 Canada IF 198: 1 (2014) H KP137498
C. claroplaniusculus RH 2334 claroplaniusculus (holotype) R. Henry 1967 France SMF 99(1): 65 (1983) PC KP013184
C. decipiens PML 366 decipiens f. decipiens (neotype) P. Moënne-Loccoz 1986 France AC 11(1): f507 (2001), G FN428988
AC 2: f52 (1990)
C. disjungendus PAK 4370 disjungendus (lectotype) P.A. Karsten < 1893 Finland ASFFF 9(1): 6 (1893) H KP013190
C. duracinus PML 349 duracinus (neotype) P. Moënne-Loccoz 1986 France AC 2: f76 (1990) G KX964582
C. duracinus s.lat. SH188648.07FU (2 sequences) sp. na na Denmark/Germany na na AJ889943
C. duracinus s.lat. SH094372.07FU (6 sequences) rigens na na NA / Italy na na JF907880
C. fuscescens H:6001898 fuscescens (holotype) K. Liimatainen & T. Niskanen 2008 Finland IF 201: 2 (2014) H KP165546
C. fuscobovinaster IK 09-537 fuscobovinaster (holotype) I. Kytövuori 2009 Norway Mycologia 105(4): 990 (2013) H, S, NY JX407316
C. gallurae CONS 00076 gallurae (holotype) D. & M. Antonini, G. Consiglio 2002 Italy Il genereCortinarius in CONS FN428979
Italia 3: C101 (2005)
C. murinascens IK 08-958 murinascens (holotype) I. Kytövuori 2008 Finland IF 201: 3 (2014) H KP165570
C. neofurvolaesus CFP 1438 neofurvolaesus (holotype) T.E. Brandrud, H. Lindström, 1999 Sweden CFP: E24 (2014) S DQ139999
H. Marklund, S. Muskos
Table 2 (cont.)
Species Voucher/SH Voucher/SH annotation Leg. Collection Country Taxonomy Herbarium Accession*
C. = Cortinarius date

185
K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini
C. niveotraganus SH188538.07FU (8 sequences) niveotraganus na na FS na na KM273103
C. olididisjungendus TN 07-191, H:7000854 olididisjungendus (holotype) na 2007 Canada IF 186: 2 (2014) H KM273091
C. orasericeus RH 70239 orasericeus (holotype) R. Henry 1975 France SMF 99(1): 69 (1983) PC KP013203
C. quarciticus CFP 765 quarciticus (holotype) T.E. Brandrud, H. Lindström, 1988 Sweden CFP: C59 (1994) S UDB000748
H. Marklund, S. Muskos
C. sordidemaculatus RH 1122 sordidemaculatus (holotype) R. Henry < 1981 France SMF 97(3): 196 (1981) PC DQ139984
C. sp. IB 86/172 bovinus cf. M. Moser 1986 Austria na IB DQ139983
C. sp. TF-01-034 aprinus T. Frøslev na Denmark na C AJ889942
C. subserratissimus IK 11-017 subserratissimus (holotype) I. Kytövuori 2011 Sweden IF 201: 4 (2014) H KP165552
C. subturibulosus SH188545.07FU (7 sequences) subturibulosus na na France/ Spain/Portugal na na FJ928484
C. tacitus AB 05-09-72 tacitus (holotype) A. Bidaud 2005 France AC 22: f1400 (2014) PC KX964583
C. torvus SH009362.07FU (10 sequences) torvus na na NA /FS /Germany na na UDB001345
C. urbicus SH188612.07FU (3 sequences) urbicus na na Canada/ FS na na UDB000743
outgroup
C. anomalovelatus JFA13109 anomalovelatus (holotype) J.F. Ammirati 2007 USA IF 93: 1 (2014) WTU FJ717605
C. lepidopus sensu auct. SH196665.07FU (12 sequences) anomalus na na U na na UDB002227
C. caesiocinctus Sa57-13 caesiocinctus (holotype) R. Kühner 1957 France DM 20(77): 92 (1989) G DQ663239
C. flavipallens IK 08-1729, H:6032393 flavipallens (holotype) I. Kytövuori 2008 Finland Persoonia 33 : 125 (2014) H KF732554
C. sannio MM 97/352, IB:1997/0352 sannio (holotype) M. Moser 1997 USA Mycotaxon 72 : 315 (1999) IB KF732420
CFP, Cortinarius, Flora Photographica; AC, Atlas des Cortinaires; FN, Funga Nordica; DM, Documents Mycologiques; SMF, Bulletin de la Société Mycologique de France; FMDS, Bulletin de la Fédération Mycologique Dauphiné-Savoie; JEC, Journal des Journées Européennes du
Cortinaire; K&R, Flore analytique des Champignons supérieurs (Küehner & Romagnesi); IF, Index Fungorum; ASFFF, Acta Societatis pro Fauna et Flora Fennica; BFNF, Bidrag till kännedom av Finlands Natur och Folk; NA, North America (USA, Canada); FS, Fennoscandia (Denmark,
Sweden, Norway, Finland, Estonia, Lithuania, Latvia); U, Europe.
* Sequences generated for the present work are highlighted in bold.
tinental biogeographical distribution in some cases (Table 3).
Minimal interspecific phylogenetic distances Dinter min range
from 3 to 9 substitutions plus 2–4 indels, representing 0.5–2 %
of sequence divergence. Those are, with one exception, longer
than Dintra max for a given pair of sister species clades (Table
3). The topology of /Bicolores strongly supports two distinct
lineages within the section, one including C. cagei, C. evernius,
C. plumulosus, C. refectus, C. sp1 and C. sp2, and another one
including C. dolabratoides, C. dolabratus, C. glaphurus, C. hir-
cinosmus, C. tortuosus and C. turgidipes (Fig. 2).
As sampled here, /Saturnini includes 6 species in Europe
and North America, each represented by 1 to 44 sequences
(Fig. 3, Table 2). Sequencing existing type material revealed
a much higher rate of synonymy when compared to species
in /Bicolores, with 17 binomials identified as later names for
C. confirmatus, C. imbutus, C. lucorum or C. saturninus. A com-
paratively wider species concept has emerged in this section,
as illustrated by the case of C. saturninus, which merged not
less than 9 holotypes previously reported to belong in unrelated
sections. The considerable rise in species polymorphism result-
ing from such finding has been dealt with at the infraspecific
taxonomic level in the last release of the ADC (Bidaud et al.
2015). In order to stabilize the nomenclature and fix the concept
of species widely accepted as genuine members of the Saturnini
section – or interpreted by some authors in sect. Bicolores,
we designated neotypes for C. saturninus, C. imbutus and
C. lucorum (see Taxonomy). Our work also positioned C. stuntzii
and a morphogenetic, widened concept of C. confirmatus in the
revised section, and it unravelled C. cyprinus as an overlooked
species in sect. Saturnini (Fig. 3, Table 2, 3).
Intraspecific phylogenetic distances were considerably larger
in /Saturnini when compared to /Bicolores, with a Dintra max up
to 6 substitutions plus 1 indel, representing 1.2 % of sequence
divergence, only considering sequences with trace files (Table
3). The interspecific genetic distance within the clade is of 3
substitutions plus up to 5 indels, representing 0.5–1.3 % of
sequence divergence, except for C. lucorum, which is more
distantly related to the other species (Dinter min = 16 substitu-
tions plus 3 indels to C. confirmatus, representing 3.1 % of
sequence divergence). Although not significantly lower than
in /Bicolores, these distances exceed Dintra max values only
for C. cyprinus and C. lucorum (Table 3). The topology of the
phylogenetic tree depicted in Fig. 3 indicates that C. lucorum
represents an early-diverging lineage in the section and it
supports C. saturninus, C. cyprinus and C. stuntzii as part of a
distinct lineage within /Saturnini.
The wide survey of subg. Telamonia depicted in Fig. 1 also al-
lows phylogenetic positioning of morphological Bicolores and
Saturnini, i.e., of those species that have been included in the
two sections based on purely morpho-anatomical criteria, but
which evolutionary history is unrelated to that of /Bicolores and
/Saturnini. Eight binomials usually treated in Bicolores could
be assigned to five morphogenetic species (Fig. 1, Table 1):
C. cinnamoviolaceus (incl. C. parevernius, C. subparever-
nius, C. basicyaneus and C. imbutus sensu CFP), C. mattiae,
C. parevernioides, C. salicinus and C. disjungendus. Similarly,
ten species formerly treated in Saturnini based on morphology,
turned out to be phylogenetically distant from /Saturnini. Six
of them could further be assigned to other known sections:
C. cypriacoides, C. subfirmus and C. illepidus in sect. Bovini,
C. saturninoides in sect. Sciophylli, C. oxytoneus in sect. Dura-
cini and C. sciophylloides in sect. Brunneotincti (Fig. 1, Table 1).
(text continues on p. 190)

186 Persoonia – Volume 39, 2017
0.02 substitution per site
Cortinarius parevernius RH3258a78 HOLOTYPE France
Cortinarius fuscobovinaster IK09-537 HOLOTYPE Norway
Cortinarius bovinus IK04-038 NEOTYPE Finland
Cortinarius quarciticus CFP765 HOLOTYPE Sweden
Cortinarius subturibulosus SH188545.07FU France-Spain-Portugal (7 seq)
Cortinarius valgus SH188568.07FU Canada-U (6 seq)
Cortinarius oxytoneus RH931 HOLOTYPE France
Cortinarius basicyaneus RH70942 HOLOTYPE France
Cortinarius mattiae AB06-09-153 France
Cortinarius duracinus PML349 NEOTYPE France
Cortinarius saturninoides XC2010-29 France
Cortinarius saturninoides XC2013-144 France
Cortinarius sciophylloides PML5446 France
Cortinarius nefastus XC2014-60 HOLOTYPE France
Cortinarius lucorum SH188624.07FU USA-Estonia-Italy (3 seq)
Cortinarius malachius SH188502.07FU NA-U (15 seq)
Cortinarius orasericeus RH70239 HOLOTYPE France
Cortinarius athabascus DBB27618 HOLOTYPE USA
Cortinarius contractus RH1240 HOLOTYPE France
Cortinarius sp. SH188648.07FU Denmark-Germany (2 seq)
Cortinarius cypriacoides asp. cypriacoides PML1269 HOLOTYPE France
Cortinarius sp. SH009438.07FU Canada (1 seq)
Cortinarius flavipallens IK08-1729 HOLOTYPE Finland
Cortinarius saturninoides XC2014-119 France
Cortinarius murinascens IK08-958 HOLOTYPE Finland
Cortinarius saturninoides AB00-10-148 France
Cortinarius sannio IB97/352 HOLOTYPE USA
Cortinarius dolabratus AB02-10-71 France
Cortinarius olididisjungendus H7000854 HOLOTYPE Canada
Cortinarius decipiens PML366 NEOTYPE France
Cortinarius saturninoides XC2014-64b France
Cortinarius caesioarmeniacus H7000901 HOLOTYPE Canada
Cortinarius suboxytoneus AB01-09-56 HOLOTYPE France
Cortinarius disjungendus PAK4370 LECTOTYPE Finland
Cortinarius cyanosterix RH338 HOLOTYPE France
Cortinarius mattiae CFP1204 Sweden
Cortinarius anomalovelatus JFA13109 HOLOTYPE USA
Cortinarius torvus SH009362.07FU NA-FS-Germany (10 seq)
Cortinarius biformis SH188479.07FU NA-U (41 seq)
Cortinarius subfirmus AB08-10-363 HOLOTYPE France
Cortinarius illepidus AB11-11-330 France
Cortinarius aprinus TF01-034 Denmark
Cortinarius furiosus XC201464c HOLOTYPE France
Cortinarius sciophylloides AB99-10-254 HOLOTYPE France
Cortinarius sordidemaculatus SH188519.07FU NA-FS-France (9 seq)
Cortinarius oxytoneus RH3451 France
Cortinarius cypriacoides asp. cypriacoides PML3984 France
Cortinarius neofurvolaesus CFP1438 HOLOTYPE Sweden
Cortinarius illepidus AB11-11-331 France
Cortinarius mattiae PML2298 France
Cortinarius subviolascens AB99-09-77 France
Cortinarius claroplaniusculus RH2334 HOLOTYPE France
Cortinarius bovinus cf. IB86/172 Austria
Cortinarius cinnamoviolaceus IB48/590 HOLOTYPE Austria (short)
Cortinarius brunneifolius TN-06146 HOLOTYPE Finland
Cortinarius imbutus SH188640.07FU Sweden-Italy (2 seq)
Cortinarius fuscescens H6001898 HOLOTYPE Finland
Cortinarius alboviolaceus SH188487.07FU NA-U (26 seq)
Cortinarius anisochrous IK01-030 HOLOTYPE Estonia
Cortinarius mattiae H6029375 Finland
Cortinarius subparevernius RH526 HOLOTYPE France (short)
Cortinarius mattiae KS-CO1936 ISOTYPE Sweden
Cortinarius mattiae IK98-001 Finland
Cortinarius sp. (ecto. Quercus) LM5411 Austria
Cortinarius saturninoides XC2010-56 France
Cortinarius saturninoides AB12-10-93 France
Cortinarius anisatus CFP1200 HOLOTYPE Sweden
Cortinarius subserratissimus IK11-017 HOLOTYPE Sweden
Cortinarius lepidopus s. auct. SH196665.07FU U (12 seq)
Cortinarius sciophylloides PML2381 France
Cortinarius ortovernus JB604808 HOLOTYPE Spain
Cortinarius imbutus CFP574 Sweden
Cortinarius sciophylloides AB91-10-291 France
Cortinarius mattiae IK01-039 Sweden
Cortinarius subviolascens PML650 France
Cortinarius salicinus XC2014-03 HOLOTYPE France
Cortinarius cypriacoides asp. lucorum PML3979 France
Cortinarius mattiae H6000560 Finland
Cortinarius parevernioides AB02-09-50 HOLOTYPE France
Cortinarius sp. (ecto. Fagus) MFT60 Germany
Cortinarius caesiocinctus Sa5713 HOLOTYPE France
Cortinarius mattiae PML3989 France
Cortinarius cinnamoviolaceus AB12-11-240 France
Cortinarius cylindratus RH4000 HOLOTYPE France
Cortinarius tacitus AB05-09-72 HOLOTYPE France
Cortinarius niveotraganus SH188538.07FU FS (8 seq)
Cortinarius rigens SH094372.07FU NA-Italy (6 seq)
Cortinarius urbicus SH188612.07FU Canada-FS (3 seq)
Cortinarius mattiae AB13-08-35 France
Cortinarius gallurae CONS00076 HOLOTYPE Italy
63/-
61/-
67/-
71/-
65/
0.80
99/0.75
99/0.92
75/0.65
98/0.66
84/0.7
54/-
73/-
55/-
60/-
97/0.42
91/-
67/
0.65
56/-
94/-
51/-
99/-
100/0.88
100/0.77
/Saturnini (Fig. 3)
/Bicolores (Fig. 2)
Cortinarius cinnamoviolaceus TN05-198 Finland
Cortinarius cinnamoviolaceus TN05-051 Finland
Outgroup
(Sat)
(Sat)
(Sat)
(Sat)
C. serratissimus
(Sat)
(Sat)
(Sat)
C. subbulliardioides
(Sat)
C. cinnamoviolaceus
(Bic)
C. mattiae (Bic)
(Bic)
(Bic)
(Bic)
(Sat)
(Sat)
Sect. Bovini
Sect. Sciophylli
Sect. Disjungendi
Sect. Duracini
Sect. Firmiores
Sect. Urbici
Sect. Telamonia
Sect. Brunneotincti
Sect. Hydrocybe
Cortinarius subgen. Telamonia pp
ITS BI+ML phylogeny
LnL (harmonic mean) = - 5537.69
Parsimony (ML) :688
Tree size (ML) :1.51079
Alignment length :543 nts
Nb of sequences :348 (419)
Nb of species :67
Fig. 1 Sections Bicolores and Saturnini within subg. Telamonia. — Bayesian 50 % majority-rule consensus tree inferred from the analysis of 348 ITS se-
quences (419 represented, due to Species Hypotheses, see Material and Methods) spanning subg. Telamonia plus 5 outgroup sequences, with collapse of
the /Bicolores and /Saturnini clades that are developed in Fig. 2 and 3, respectively. Branches with strong statistical support (BPP ≥ 95 % and SH-aLRT > 0.8)
are highlighted as thick lines, others display support values as % BPP/SH-aLRT. Species excluded from these two clades but morphologically included in sect.
Bicolores and sect. Saturnini and for which molecular data are available, are indicated by (Bic) and (Sat), respectively. Sequences of collections taxonomically
described in these two sections are highlighted in bold. Section assignment follows Niskanen et al. (2012).

187
K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini
0.005 substitution per site
Cortinarius evernius IK97-123 Finland
Cortinarius evernius TN10-074 Canada
Cortinarius glaphurus RH71421 HOLOTYPE France
Cortinarius tortuosus AB96-08-19 France
Cortinarius evernius TN07-312 Canada
Cortinarius imbutoides AB04-09-186 HOLOTYPE France
Cortinarius dolabratus TN09-196 USA
Cortinarius tortuosus AB01-09-19 France
Cortinarius hircinosmus AB04-10-357 France
Cortinarius flabelloides XC2008-43 HOLOTYPE France
Cortinarius cedriosmus AB92-10-350 HOLOTYPE France
Cortinarius plumulosus TN04-730 Finland (short)
Cortinarius evernius TN10-055 Canada
Cortinarius dolabratoides H6033567 HOLOTYPE Finland
Cortinarius sp. UBCOGTR194 Canada
Cortinarius tubulosus AB03-11-87 HOLOTYPE France
Cortinarius evernius PML622 France
Cortinarius dolabratus TN11-246 USA
Cortinarius minicolor PML738 France
Cortinarius dolabratoides IK01-062 Finland
Cortinarius refectus PML769 France
Cortinarius paranomalus GK1142 HOLOTYPE France
Cortinarius tortuosus TN09-046 USA
Cortinarius scriptor AB97-10-341 France
Cortinarius glaphurus TN12-221 USA
Cortinarius dolabratus TN12-200 USA
Cortinarius tortuosus AB96-10-124 France
Cortinarius tortuosus TN07-307 Canada
Cortinarius tortuosus AB95-09-34 France
Cortinarius refectus AB05-09-138 France
Cortinarius dolabratus TN03-1713 Slovakia
Cortinarius dolabratus H6033519 Finland
Cortinarius turibulosus AB92-10-332 France (short)
Cortinarius refectus AB99-09-121 France
Cortinarius dolabratus IK02-033 Finland
Cortinarius tortuosus TN10-087 Canada
Cortinarius tortuosus CFP493 Norway
Cortinarius evernius AB04-09-212 France
Cortinarius turibulosus PML2390 France
Cortinarius evernius TN10-054 Canada
Cortinarius dolabratus AB89-11-309 France
Cortinarius imbutoides KS-CO1290 Sweden
Cortinarius glaphurus AB14-11-138 France
Cortinarius fundatus AB98-09-119 Canada
Cortinarius evernius TN05-238 Norway
Cortinarius cagei SH188634.07FU Germany-Italy (2 seq)
Cortinarius periodolens ad int. XC2014-02 France
Cortinarius hircinosmus PML334 HOLOTYPE France
Cortinarius dolabratus SH188528.07FU NA-FS-Slovakia (10 seq)
Cortinarius evernius PML376 France
Cortinarius dolabratus IK95-347 Finland
Cortinarius livor AB02-09-32 France
Cortinarius scriptor AB94-10-268 France
Cortinarius fundatus AB10-09-183 France
Cortinarius glaphurus XC2011-212 France
Cortinarius tubulosus AB08-11-445 France
Cortinarius tortuosus PML3551 France
Cortinarius glaphurus AB03-10-56 France
Cortinarius dolabratus IK95-1576 Finland
Cortinarius evernius f. pseudoscutulatus AB00-09-83 HOLOTYPE France
Cortinarius sp. TN05-033 Finland
Cortinarius plumulosus IK98-1612 Finland
Cortinarius refectus IK96-1031 Germany
Cortinarius violaeolens XC2009-41 HOLOTYPE France
Cortinarius evernius TN07-328 Canada
Cortinarius dolabratoides H6033570 Finland
Cortinarius dolabratus XC2013-103 France
Cortinarius refectus AB96-09-73 EPITYPE Germany
Cortinarius minicolor AB92-10-256 France
Cortinarius dolabratus AB13-10-120 France
Cortinarius imbutoides KS-CO1576 Sweden
Cortinarius dolabratoides H6033575 Finland
Cortinarius refectus PML17 France
Cortinarius glaphurus AB99-11-345 France
Cortinarius evernius f. fragrans PML1727 HOLOTYPE France
Cortinarius tortuosus IK99-709 Finland
Cortinarius tortuosus TN05-006 Finland
Cortinarius evernius PML212 France
Cortinarius plumulosus RH3417 HOLOTYPE France
Cortinarius dolabratus AB98-09-94 Canada
Cortinarius dolabratus TN09-139 USA
Cortinarius evernius var. evernius P ML3469 France
Cortinarius dolabratoides H6033615 Finland
Cortinarius plumulosus PML3308 France
Cortinarius laetior PAK354 HOLOTYPE Finland (short)
Cortinarius dolabratus CFP990 EPITYPE Sweden
Cortinarius dolabratoides AB07-08-48 France
Cortinarius dolabratus AB01-09-41 France
Cortinarius dolabratus AB04-09-169 France
Cortinarius evernius CFP792 NEOTYPE Sweden
Cortinarius tortuosus PML386 France
Cortinarius minicolor AB04-09-266 France
Cortinarius sp. TN12-217 USA
Cortinarius evernius var. insignis AB09-07- 44 France
Cortinarius phaeoruber RH80814 HOLOTYPE France
Cortinarius glaphurus XC2009-64 France
Cortinarius parvulior ad int. AB96-09-47 France
Cortinarius cagei PML3588 France
Cortinarius tortuosus PML1225 France
Cortinarius evernius PML230 France
Cortinarius evernius f. pseudoscutulatus AB91-08-42 France
Cortinarius dolabratoides IK04-051 Finland
Cortinarius tortuosus IB79/533 NEOTYPE Sweden (short)
Cortinarius turibulosus AB91-11-360 France (short)
Cortinarius evernius SH188514.07FU Canada-FS-Germany (11 seq)
Cortinarius turibulosus PML1067 France (short)
Cortinarius turibulosus SH094444.07FU NA-France (3 seq)
Cortinarius cagei CFP1260 Sweden NEOTYPE
Cortinarius dolabratus TN02-1095 Finland
Cortinarius evernius IK00-038 Finland
Cortinarius fundatus PML657 France
Cortinarius sp. SH094485.07FU Poland (2 seq)
Cortinarius hircinosmus F44390 Sweden
Cortinarius tortuosus PML1214 France
Cortinarius refectus PML2159 France
Cortinarius tortuosus AB02-09-41 France
Cortinarius refectus AB04-10-321 France
Cortinarius evernius TN07-223 Canada
Cortinarius tortuosus SH094369.07FU USA-U-Japan (7 seq)
Cortinarius testaceoviolaceus AB92-10-293 France
Cortinarius cagei PML1057 France
Cortinarius dolabratus TN02-959 Finland
Cortinarius turgidipes AB93-10-425 HOLOTYPE France
Cortinarius hircinosmus H6033565 Finland
82/-
85/-
79/-
80/-
67/-
91/0.94
65/0
66/-
95/-
75/-
Cortinarius tortuosus
Cortinarius glaphurus
Cortinarius dolabratus
Cortinarius dolabratoides
Cortinarius refectus
Cortinarius plumulosus
Cortinarius cagei
Cortinarius evernius
Cortinarius hircinosmus
Cortinarius turgidipes
Cortinarius sp1
Cortinarius sp2
Cortinarius sect. Bicolores
ITS BI+ML phylogeny
LnL (harmonic mean) = -1488.69
Parsimony (ML) :71
Tree size (ML) :0.1456
Alignment length :608 nts
Nb of sequences :124 (153)
Nb of species :12
Fig. 2 The morphogenetic Bicolores section. — Bayesian 50 % majority-rule consensus tree inferred from the analysis of the ITS sequence of 124 (153 repre-
sented, due to Species Hypotheses, see Material and Methods) Telamonia sequences nested in /Bicolores. Branches with strong statistical support (BPP ≥ 95%
and SH-aLRT > 0.8) are highlighted as thick lines, others display support values as % BPP/SH-aLRT. Sequences from ‘type’ material are highlighted in bold,
those having nomenclatural priority are further underlined.

188 Persoonia – Volume 39, 2017
0.004 substitution per site
Cortinarius incarnatolilascens RH71502 HOLOTYPE France
Cortinarius saturninus TN09-208 USA
Cortinarius imbutus asp. imbutus XC2002-107 France
Cortinarius imbutus asp. imbutus AB09-11-471 France
Cortinarius confirmatus XC2006-204 France
Cortinarius saturninus SH188563.07FU Canada-Estonia-China (6 seq)
Cortinarius imbutus IK94-1236 Finland
Cortinarius imbutus asp. imbutus XC2002-122 France
Cortinarius fulvorimosus XC2007-14 HOLOTYPE France
Cortinarius saturninus asp. salicis XC2011-205 France
Cortinarius confirmatus AB09-11-450 France
Cortinarius imbutus JMB2008092703 France
Cortinarius saturninus asp. deceptivus AB04-10-344 France
Cortinarius umbrinoconnatus RH476 HOLOTYPE France (short)
Cortinarius subtorvus CFP408 Sweden
Cortinarius salicis PML3967 France
Cortinarius saturninus SH094324.07FU USA-U (13 seq)
Cortinarius urbicus AB97-09-187/PML5347 France
Cortinarius saturninus asp. dionisiae AB14-09-47 France
Cortinarius betulaecomes RH3123 HOLOTYPE France
Cortinarius lucorum asp. montis-dei PML4143 France
Cortinarius saturninus XC2002-167 France
Cortinarius imbutus TN11-252 USA
Cortinarius gramineus RH81181 HOLOTYPE France
Cortinarius lucorum asp. circumvelatus PAM14090808 France
Cortinarius cyprinus AB04-09-167 France
Cortinarius subtorvus KH14 Norway (Svalbard)
Cortinarius bulbosovolvatus RH84159 ISOTYPE France
Cortinarius imbutus IK97-1162 NEOTYPE Finland
Cortinarius imbutus asp. imbutus AB98-10-358 France
Cortinarius imbutus AB02-10-106 France
Cortinarius subtorvus O50591 Norway (Svalbard)
Cortinarius saturninus AB05-10-273 France
Cortinarius confirmatus XC2005-249 France
Cortinarius cyprinus PML81 France
Cortinarius cyprinus AB11-10-192 France
Cortinarius cyprinus XC2007-95 France
Cortinarius confirmatus FR2012089 France
Cortinarius saturninus asp. salicis XC2007-108 France
Cortinarius rastetteri RH71682 HOLOTYPE France
Cortinarius confirmatus asp. imbutus XC2012-171 France
Cortinarius confirmatus asp. spurcatocephalus XC95-10-04-06 France
Cortinarius cyprinus AB06-09-144 France
Cortinarius saturninus H6029320 Finland
Cortinarius imbutus XC2002-109 France
Cortinarius imbutus IK98-2242 Sweden
Cortinarius confirmatus asp. imbutus PML4722 France
Cortinarius saturninus XC2007-90 France
Cortinarius cyprinus PAM13092901 France
Cortinarius confirmatus asp. assiduus AB09-11-514 France
Cortinarius cyprinus PML425 France
Cortinarius sp. TAAM128765 Estonia
Cortinarius saturninus XC2006-194 France
Cortinarius circumvelatus PML34 HOLOTYPE France
Cortinarius imbutus TN11-257 USA
Cortinarius cistoadelphus ad int. AB92-11-422 France
Cortinarius stuntzii Rehner394 HOLOTYPE USA
Cortinarius saturninus XC96-10-26-09 France
Cortinarius dissidens PR258 HOLOTYPE France
Cortinarius saturninus asp. saturninus XC2014-116 France
Cortinarius umidicola 10433 SYNTYPE USA (short)
Cortinarius cyprinus PML344 France
Cortinarius saturninus JMB2009101002 France
Cortinarius marginatosplendens PML2215 ISOTYPE France
Cortinarius confirmatus asp. subcylindratus XC2013-160 France
Cortinarius betulaecomes RH71030 France
Cortinarius salicis RH2623 HOLOTYPE France
Cortinarius confirmatus asp. assiduus XC2013-156 France
Cortinarius saturninus asp. urbicoides AB95-11-144 France
Cortinarius saturninus asp. urbicoides AB02-10-179 France
Cortinarius saturninus asp. salicis AB14-11-160 France
Cortinarius cyprinus XC2012-26 HOLOTYPE France
Cortinarius lucorum TN03-1169 Sweden
Cortinarius saturninus asp. cohabitans XC2014-63 France
Cortinarius imbutus asp. saturnalis XC2014-61 France
Cortinarius saturninus XC2008-61 France
Cortinarius saturninus asp. saturninus XC2014-114 France
Cortinarius confirmatus asp. rubricosissimus AB09-11-452 France
Cortinarius imbutus TN05-167 Finland
Cortinarius imbutus asp. imbutus XC2002-106 France
Cortinarius confirmatus FR2012405 France
Cortinarius assiduus MES3541 HOLOTYPE Spain
Cortinarius imbutus asp. imbutus AB04-09-228 France
Cortinarius imbutus AB00-09-127 France
Cortinarius confirmatus RH3195 HOLOTYPE France
Cortinarius urbicus PML75 France (short)
Cortinarius urbicus var. sporanotandus PML4578 HOLOTYPE France
Cortinarius sp. SH094374.07FU U-Iran (6 seq)
Cortinarius saturninus asp. deceptivus AB98-10-381 France
Cortinarius saturninus asp. urbicoides XC2001-107 France
Cortinarius cyprinus XC2007-103 France
Cortinarius montis-dei PML4142 HOLOTYPE France
Cortinarius cyprinus JMB2014111802 France
Cortinarius confirmatus asp. spurcatocephalus XC2011-199 France
Cortinarius lucorum IK89-748 Finland
Cortinarius imbutus AB08-10-307 France
Cortinarius imbutus asp. vilior XC2007-104 France
Cortinarius confirmatus asp. paracohabitans AB11-11-324 France
Cortinarius lucorum KS-CO513 Sweden
Cortinarius imbutus asp. imbutus AB10-10-237 France
Cortinarius assiduus var. plesiocistus JVG9 90125-31 ISOTYPE Spain
Cortinarius confirmatus asp. confirmatus AB03-11-78 France
Cortinarius confirmatus asp. assiduus AB02-11-201 France
Cortinarius saturninus asp. salicis XC2008-55 France
Cortinarius confirmatus asp. assiduus AB05-11-423 France
Cortinarius saturninus asp. salicis XC2014-109 France
Cortinarius cyprinus AB11-11-251 France
Cortinarius confirmatus FR2016052 France
Cortinarius confirmatus asp. rubricosissimus AB00-10-193 France
Cortinarius saturninus IK94-631 Finland
Cortinarius imbutus TN11-151 USA
Cortinarius confirmatus FR2012076 France
Cortinarius imbutus asp. laetior XC2013-13 France
Cortinarius imbutus asp. saturnalis XC2014-77 France
Cortinarius saturninus CFP514 NEOTYPE Sweden
Cortinarius lucorum TN10-002 Canada
Cortinarius saturninus XC2001-104 France
Cortinarius saturninus asp. saturninus XC2007-97 France
Cortinarius imbutus AB02-09-58 France
Cortinarius laccatus PML4557 HOLOTYPE France
Cortinarius lucorum CFP490 NEOTYPE Norway
Cortinarius imbutus TN11-150 USA
Cortinarius saturninus XC2016-12 France
Cortinarius imbutus asp. imbutus XC2002-108 France
Cortinarius imbutus XC2012-96 France
Cortinarius confirmatus asp. kuehneri AB13-10-97 France
Cortinarius cyprinus XC2013-15 France
Cortinarius imbutus asp. imbutus PML375 France
Cortinarius denseconnatus RH3758 HOLOTYPE France
Cortinarius lucorum SH188495.07FU NA-FS (21 seq)
56/-
51/-
87/0.3
85/0.84
96/-
92/0.87
79/0.81
58/0.92
*
Cortinarius cyprinus TEB348-10 Norway
Cortinarius imbutus
Cortinarius confirmatus
Cortinarius saturninus
Cortinarius cyprinus
Cortinarius stuntzii
Cortinarius lucorum
Cortinarius sect. Saturnini
ITS BI+ML phylogeny
LnL (harmonic mean) = -1530.69
Parsimony (ML) :65
Tree size (ML) :0.11305
Alignment length :613 nts
Nb of sequences :131 (173)
Nb of species :6
Fig. 3 The morphogenetic Saturnini section. — Bayesian 50 % majority-rule consensus tree inferred from the analysis of the ITS sequence of 131 (173 repre-
sented, due to Species Hypotheses, see Material and Methods) Telamonia sequences nested in /Saturnini. Branches with strong statistical support (BPP ≥ 95 %
and SH-aLRT > 0.8) are highlighted as thick lines, others display support values as % BPP/SH-aLRT. Sequences from ‘type’ material are highlighted in bold,
those having nomenclatural priority are further underlined. The asterisk points to a subclade that segregates a 1 nt intra-individual polymorphism, as XC 2011-
205 (within the subclade) was fruiting from the same mycelium as XC 2007-108 and XC 2014-109 (outside the subclade).

189
K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini
Table 3 Morphogenetic features of C. cinnamoviolaceus and species in sect. Bicolores and sect. Saturnini.
Species Blue hues a Odour(s)b L min Av L L max l min Av l l max Av Q Reported hostc,d Dintra max / difference rate Dinter min / difference rate (incl. indels)c
(incl. indels)a
Cortinarius cinnamoviolaceus + or – R, r, i 8.25 9.65 11.07 4.66 5.21 6.00 1.86 Picea, Abies, Pinus, Tilia, Quercus, Betula, na na
Populus
sect. Bicolores
C. cagei + 0, r, e, I 7.80 9.04 10.50 5.10 5.54 6.18 1.64 Deciduous trees 0 nt / 0 % 3 nts + 3 indels (to C. evernius) / 1 %
C. dolabratoides sp. nov. + or – CE, g 7.50 8.30 9.50 3.50 4.60 5.00 1.82 Picea, Pinus 0 nt / 0 % 3 nts (to C. dolabratus) / 0.5 %
C. dolabratus + or – ce, co 7.42 8.62 9.86 4.41 4.90 5.51 1.76 Pinus, Picea, Betula, Fagus, Quercus 3 nts / 0.5 % 3 nts (to C. dolabratoides) / 0.5 %
C. evernius + 0, ce, r, R 8.75 10.34 11.85 5.35 6.01 6.77 1.72 Picea, Abies 1 nt / 0.2 % (1 nt + 1 indel / 0.3 %) 3 nts + 3 indels (to C. cagei) / 1 %
C. glaphurus + or – ce, r, CE, V 8.03 9.32 10.60 4.82 5.23 5.78 1.78 Pinus, Quercus, Fagus, Abies, Picea, 2 nts + 4 indels / 1 % 4 nts + 2 indels (to C. tortuosus) / 1 %
Populus, Betula
C. hircinosmus + or – 0, r, B 8.00 9.04 10.00 4.70 4.98 5.40 1.82 Picea 2 nts / 0.3 % 9 nts + 2 indels (to C. dolabratus) / 1.8 %
C. plumulosus + or – ca, r, i 8.75 9.78 11.08 4.80 5.53 6.10 1.77 Picea, Abies 1 nt + 4 indels / 0.8 % 7 nts + 3 indels (to C. evernius) / 1.6 %
C. refectus + g, r 8.06 9.50 10.94 5.58 6.30 6.92 1.51 Abies, Picea, Fagus, Quercus 0 nt + 1 indel / 0.2 % 4 nts + 3 indels (to C. evernius) / 1.2 %
C. tortuosus + ce, 0, E 8.00 9.30 10.61 4.83 5.44 6.00 1.71 Tsuga, Abies, Picea, Pinus 1 nt + 1 indel / 0.3 % 4 nts + 2 indels (to C. glaphurus) / 1 %
C. turgidipes (–) 0 7.50 8.50 9.50 5.00 5.30 6.00 1.60 Picea na 3 nts + 4 indels (to C. dolabratus) / 1.2 %
C. sp1 (+) na na na na na na na na na 0 nt / 0 % 8 nts + 4 indels (to C. evernius) / 2 %
C. sp2 na na na na na na na na na na na 8 nts + 4 indels (to C. evernius) / 2 %
sect. saturnini
C. confirmatus + or – 0, ca, r, g 6.91 8.26 9.79 4.27 4.79 5.61 1.73 Quercus, Cistus, Pinus, Betula, Populus, 6 nts + 1 indel / 1.2 % 3 nts (to C. imbutus) / 0.5 %
Picea
C. cyprinus + ca, p 6.90 8.40 9.90 4.18 4.77 5.45 1.76 Deciduous trees 0 nt / 0 % (5 nts / 0.8 %) 3 nts + 2 indels (to C. saturninus) / 0.8 %
C. imbutus + or – 0, g, ca 7.27 8.68 10.21 4.09 4.62 5.41 1.88 Betula, Salix, Alnus, Fagus, Populus, 3 nts + 1 indel (0.7 %) 3 nts (to C. confirmatus) / 0.5 %
Carpinus, Picea
C. lucorum + r, ca, 0 8.07 9.56 11.07 5.36 5.86 6.71 1.63 Populus, Betula, Carpinus, Quercus, Picea, 2 nts + 1 indel (0.5 %) 16 nts + 3 indels (to C. confirmatus) / 3.1 %
Tsuga
C. saturninus + or – 0, ca, g 7.10 8.38 9.59 4.38 4.78 5.39 1.76 Salix, Betula, Corylus, Tilia, Fagus, Quercus, 4 nts + 1 indel / 0.8 % 3 nts + 2 indels (to C. cyprinus) / 0.8 %
Populus, Carpinus, Picea, Abies (7 nts + 3 indels / 1.6 %)
C. stuntzii (+) 0 9.60 11.50 14.40 5.90 6.70 8.50 1.72 Salix na 3 nts + 5 indels (to C. saturninus) / 1.3 %
nt = nucleotide change; indel = insertion or deletion; na = not applicable (single sequence) or not available.
a Brackets mark uncertainty because of single collections (column ‘Blue hues’) or lack of available trace files for public sequences (column ‘Dintra max’).
b 0 = odourless; b = burnt keratin; ca = camphorated; ce = cedar wood; co = coconut; e = earth-like; g = grass-like; i = iodine; p = plum; r = radish. Upper/ lower case relates to odour intensity. Bold indicates the most frequent odour.
c Bold indicates proven interaction (ectomycorrhizal sequences, column ‘Reported host’) or species with Dinter min > Dintra max (column Dinter min).
d Names are in the order of citation frequency.

190 Persoonia – Volume 39, 2017
TAXONOMY
Each morphogenetic (i.e., defined by both morpho-anatomic
features and unique molecular signature) species that belongs
in the two revised sections is here introduced. To keep the
present survey reasonably short, taxonomic descriptions are re-
stricted to the new C. dolabratoides species, and major changes
relative to the current use of the other names are highlighted in
the notes. Because of its intricate taxonomic relationships with
C. imbutus and C. dolabratus, we also provide below a taxo-
nomic update of C. cinnamoviolaceus, even though the species
is not part of sect. Bicolores nor sect. Saturnini dealt with here.
A key to species treated in the present work is proposed at the
end of the article.
Cortinarius cinnamoviolaceus M.M. Moser, Nova Hedwigia
14: 514. 1967 — MycoBank MB#329008
= Cortinarius basicyaneus Rob. Henry & Trescol ex Bidaud & Eyssart.,
Bull. Semestriel Féd. Assoc. Mycol. Méditerranéennes 25: 38. 2004.
= Cortinarius contractus Rob. Henry, Doc. Mycol. 16, 61: 27. 1985.
= Cortinarius cylindratus Rob. Henry, Bull. Soc. Mycol. France 99: 91.
1983.
= Cortinarius subparevernius Rob. Henry, Bull. Soc. Mycol. France 85:
442. 1970.
[= Cortinarius parevernius Rob. Henry, Fl. Anal. Champ. Sup.: 303. 1953,
nom. inval. (no diagnosis, no type designated)].
Type.
AustriA, Tirol, near Hötting, in mixed forest, 18 Sept. 1948, M. Moser,
IB 48/590, holotype. MycoBank MBT#372783. ITS (partial) sequence de-
posited in GenBank under KX964412.
Misapplied names
– Cortinarius dolabratus Fr., Epicr. Syst. Mycol.: 311. 1838, sensu Bidaud
et al. (2008).
– Cortinarius imbutus Fr., Epicr. Syst. Mycol.: 306. 1838, sensu Brandrud
et al. (1998).
– Cortinarius evernius Fr., Epicr. Syst. Mycol.: 294. 1838, sensu auct.
Illustrations — Bidaud et al. 2008: pl. 639 (as C. dolabratus);
Brandrud et al. 1998: pl. D60 (as C. imbutus).
Taxonomic descriptions — Bidaud et al. 2008: f. 817 (as C. do-
labratus); Brandrud et al. 1998: pl. D60 (as C. imbutus).
Notes — This is C. evernius sensu Konrad & Maublanc
(1930) and sensu Henry (1937), with smaller spores and
raphanoid smell. Our phylogenetic analysis reveals a much
wider range of chromatic variability for this species, making it
compatible with both sect. Bicolores and Duracini. In addition,
the /C. cinnamoviolaceus clade here delineated sheds new
lights on the intricate links between these two sections and
sect. Saturnini (Fig. 1). Indeed, as redefined here, the species
falls outside the three sections but it merges:
i. typical Bicolores concepts – C. parevernius and C. cinna-
moviolaceus;
ii. typical Duracini concepts – C. subparevernius, C. cylin-
dratus and C. contractus;
iii. a species defined by its author as intermediate between
these two sections – C. basicyaneus;
iv. a Duracini concept hiding a phylogenetic Bicolores –
C. dolabratus; and
v. a Saturnini binomial interpreted by contemporary Nordic
authors as a Bicolores species – C. imbutus.
When displaying blue tinges, C. cinnamoviolaceus may be
confused with C. evernius but the spores of the latter are larger,
gills lack reddish hues and the smell is weak or indistinct. Cor-
tinarius mattiae may fruit in the same places and is similar in
appearance but the pileus is less dark coloured, not glabrous
and almost not hygrophanous, while lamellae display even
deeper red tinges. When blue pigments are absent, C. cinna-
moviolaceus looks like a Duracini with reddish lamellae and is
nearly identical to C. dolabratus, from which it can fortunately
be distinguished by larger spores (9.7 × 5.2 µm vs 8.6 × 4.9
µm, respectively) and stronger smell (Table 3).
Cortinarius sect. Bicolores (M.M. Moser) Melot, Doc. Mycol.
20, 77: 97. 1989, emend.
Type. Cortinarius cagei Melot, Doc. Mycol. 20, 80: 58. 1990.
Notes — As phylogenetically revised here, Cortinarius sect.
Bicolores has been redefined to a rather severe extent, with
well-known representative species excluded from the revised
section and half of its new content previously described out-
side Bicolores. The original diagnosis of the section should
be emended as follow: young basidiomata usually (but not
always) with violet tinges outside and/or in the context. Pileus
strongly hygrophanous, yellowish brown, chocolate brown to
reddish brown. Stipe cylindrical, often attenuate to rooting, usu-
ally with remnants of the white universal veil. Smell indistinct,
weakly raphanoid, of cedar-wood, rarely of geosmin (earth-like,
dusty). Spores amygdaloid to ellipsoid, sometimes fusiform,
(6.5–)7–12(13) × (4–) 4.3 –7(–7.2) µm (on average: 9.3 × 5.4
µm), verrucose. Widely distributed in the Northern Hemisphere,
fruiting solitary or gregarious, rarely cespitose, mostly under
coniferous trees.
In its current sampling, it includes 12 species, 10 of which have
been or can be assigned a Latin binomial.
Cortinarius cagei Melot, Doc. Mycol. 20, 80: 58. 1990 — Myco-
Bank MB#129526
≡ Cortinarius bicolor Cooke, Grevillea XVI: 45. 1873, nom. illeg.
– Cortinarius minicolor Rob. Henry, Bull. Soc. Mycol. France 104, 4: 300.
1989 ‘1988’, sensu Bidaud et al. (2014).
[= Cortinarius periodolens Carteret & Reumaux ad int., Atlas des Corti-
naires XXII: f. 1417. 2014, nom. inval. (no diagnosis, no type designated)].
Type. Sweden, Gotland, Lummelunda, Prästänget, under broadleaf trees,
1 Oct. 1994, T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos CFP1260,
S, neotype designated here. MycoBank MBT#373139. ITS sequence depo-
sited in GenBank under KX964295.
Illustrations — Bidaud et al. 2014: pl. 959 (as C. minicolor and
C. periodolens); Brandrud et al. 1998: pl. D48.
Taxonomic descriptions — Bidaud et al. 2014: f. 1419 (as C. mini-
color) but also f. 1417 (as C. periodolens); Niskanen et al. 2012:
864; Brandrud et al. 1998: pl. D48.
Notes — Historically, C. cagei was introduced to fix the nomen-
clatural issue associated with C. bicolor Cooke, an illegitimate
name because of an earlier use of the name for another, un-
related taxon. However, by omitting to designate a holotype or
other voucher specimen for his new name, Melot did not clarify
the taxonomic ambiguity of C. bicolor. Indeed, C. bicolor was
initially described as a species with medium-sized spores (10
× 5– 6 µm) fruiting under deciduous trees. However, five years
later, it was attributed much larger spores (12–14 × 6–7 µm),
and also a broader ecology – mixed woods. It is likely that
Cooke actually lumped together two phylogenetically distinct,
but morphologically very similar species, in his latest diagnosis,
making C. bicolor a nomen dubium. As such, the name may
just be discarded but the authors of the CFP proposed an
interpretation of C. cagei that fits very well the initial concept
of C. bicolor. Because:
i. the CFP plate D48 is well-known and widely recognized
as a good illustration of C. cagei;
ii. our work considerably extends our morphogenetic, bio-
geographical and ecological knowledge of this species;
and

191
K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini
iii. there is so far no convincing candidate for the second
C. bicolor, even though C. plumulosus has been postulated
to represent that one by the authors of the ADC (cf. notes
under C. plumulosus), we fix here the species in its primary
concept through designating the sequenced CFP1260
collection of plate D48 to neotypify C. cagei.
In these new morphogenetic boundaries, C. cagei is described
in the ADC under C. minicolor, an obvious lookalike that, how-
ever, fruits under coniferous trees. Unfortunately, the holotype of
C. minicolor could not be located in PC, preventing phylogenetic
placement of the species within /Bicolores. Cortinarius cagei
also includes C. periodolens, a Bicolores species described ad
interim in the ADC, as a C. obtusus with violaceous stipe and
strong iodine smell. Phylogenetically, C. cagei is well resolved
due to the absence of any intraspecific sequence polymorphism
and of a minimal distance to its sister species C. evernius of
3 substitutions plus 3 indels (Table 3). In the field, confusions
are possible with C. refectus and C. plumulosus but spore
shape ratios and host trees of the three species should prevent
misidentification (Table 3).
Cortinarius dolabratoides Kytöv., Carteret, Bidaud, Liimat.,
Niskanen, Bellanger, Dima, Reumaux & Ammirati, sp. nov.
— MycoBank MB#818596; Fig. 4
Etymology. The name refers to the close phylogenetic and morphological
affinities with C. dolabratus.
Type. FinlAnd, Koillismaa, Taivalkoski, Loukusa, the nature reserve of
Loukusanharju, dry Pinus forest on the esker, with some Picea and Betula,
some Picea-dominated depressions, 30 Aug. 2008, I. Kytövuori 08-465,
H:6033567 (holotype H; isotype K). ITS sequence deposited in GenBank
under KX964302.
Pileus 2–7 cm, conical when young, later expanding to plain
with a distinct button-like umbo, clay brown to purplish brown,
hygrophanous. Lamellae moderately distant, strongly emar-
ginate, at first bluish then brown. Stipe 5–12 cm cylindrical to
weakly clavate, sometimes slightly routing, white, with pale
lavender blue top. Veil white, as a thin coating or obscure
bands or patches on the stipe. Context whitish to purplish in
the pileus, watery whitish bluish in the stipe. Exsiccated pileus
dark blackish greyish brown, stipe much paler. Smell weakly
grass-like or stronger, of cedar wood. Macrochemistry (on the
context of the French collection only): Gaïac: ++; phénolaniline:
+++; FMP: +++; AgNO3: 0. Spores 7–8.3 – 9.5 × 3.5–4.6–5.0
µm, Q = 1.68–1.82 –1.96, (250 spores, 7 specimens), narrowly
fusoid (to almost cylindrical), with a low suprahilar depression,
often somewhat elongated at apex, fairly finely, densely verru-
cose, often prominently more strongly at the very apex, some-
what dark-coloured, faintly dextrinoid. Lamellar trama hyphae
pale olive brownish, smooth to very finely densely scabrous.
Basidia distinctly darker, olive brown (in MLZ). In damp to dryish
boreal or alpine Picea abies forests, sometimes in dry Pinus
sylvestris-dominated forests mixed by Picea abies.
Distribution — Fairly poorly known, but considered occa-
sional.
Other specimens examined (sequenced collections marked with an
asterisk, see Table 2 for GenBank accession numbers). FinlAnd, Varsinais-
Suomi, Kisko, Kaukuri, mesic Picea forest, 16 Aug. 2000, T. Niskanen &
I. Kytövuori, H:6033518; Etelä-Häme, Juupajoki, Hyytiälä, mesic Picea forest,
18 Aug. 2004, I. Kytövuori H:6033615*; Virrat, Monoskylä, Korpijärvi E, mesic
Picea forest, 15 Oct. 2001, I. Kytövuori 01-062*, H; Pohjois-Häme, Laukaa,
Äijälä, Heinäaho, mesic Picea forest, 10 Sept. 2004, I. Kytövuori 04-051*, H;
Kainuu, Paltamo, Kontiomäki, Tololanmäki W, Kylmänpuro, W sloping, mesic
Picea forest with some Pinus, Betula, Populus tremula and Salix, 14 Sept.
2008, I. Kytövuori 08-1771*, H:6033570; Koillismaa, Taivalkoski, Metsäkylä
SW, Katajavaara, N sloping, old, mesic Picea forest with damp depressions,
some Pinus, Betula and Populus tremula, 2 Sept. 2008, I. Kytövuori 08-788*,
H:6033575. – FrAnce, Haute-Savoie, Tanninges, cespitose under Picea abies
on a decalcified substrate, elev. 1500 m, 17 Aug. 2007, A. Bidaud & R. Fillion
AB 07-08-48*, personal herbarium of A. Bidaud.
Notes — Morphologically, C. dolabratoides is reminiscent of
its sister phylogenetic species C. dolabratus. Fortunately, the
two species can be distinguished microscopically, C. dolabra-
toides delivering the narrowest spores in the section (width =
3.5–4.6–5.0 µm, Av Q = 1.82, Table 3). By comparison, the
spores of C. dolabratus are distinctly wider (width = 4.4–4.9–
5.5 µm, Av Q = 1.76, Table 3) and strongly verrucose throughout
(Fig. 4b – c). Finnish collections consistently smelled of cedar
wood, but this criterion, as a diagnostic feature, may be used
with caution since the French material displayed only a weak
grass-like odour. At the molecular level, C. dolabratoides differs
from C. dolabratus by 3 substitutions only, but is not polymorphic
at the ITS locus across its pan-European distribution range,
making it well resolved within sect. Bicolores (Fig. 2, Table 3).
Fig. 4 Cortinarius dolabratoides sp. nov. — a. In situ photograph of the French collection A. Bidaud 07-08-48; b. sporogram of the holotype collection H:6033567;
c. sporogram of the C. dolabratus collection T. Niskanen 02-959 (for comparison purposes). — Scale bars: a = 5 cm; b–c = 10 µm.
cb
a

192 Persoonia – Volume 39, 2017
Cortinarius dolabratus Fr., Epicr. Syst. Mycol.: 311. 1838 —
MycoBank MB#216747; Fig. 5a
= Cortinarius imbutoides Bidaud & Carteret, Atlas des Cortinaires XXII:
1887. 2014.
= Cortinarius phaeoruber Chevassut & Rob. Henry, Doc. Mycol. 12, 47:
52. 1982.
Types. Plate ined. 181 directed/approved by Fries, S, neotype (iconotype)
designated here (Fig. 5a), MycoBank MBT#373156. Sweden, Jämtland,
Östansjö, Håsjö, under coniferous trees, 2 Sept. 1990, T.E. Brandrud, H. Lind-
ström, H. Marklund, S. Muskos CFP990, S, epitype designated here, Myco-
Bank MBT#373157. ITS sequence deposited in GenBank under KX964309.
Illustrations — Bidaud et al. 2014: pl. 951 (as C. imbutoides);
Brandrud et al. 1998: pl. D52.
Taxonomic descriptions — Bidaud et al. 2014: f. 1409 (as C. im-
butoides); Niskanen et al. 2012: 863; Brandrud et al. 1998:
pl. D52.
Notes — The original description of C. dolabratus is appar-
ently not a critical one and a plate later approved by Fries further
defined the species as a Duracini with reddish gills. Consistently,
the authors of the CFP and of the ADC delivered very similar
interpretations of C. dolabratus, both in good accordance with
the protologue and compatible with the unpublished plate.
However, sequencing the French and Scandinavian materials
of this species, unexpectedly, revealed that they are actually
phylogenetically distinct and unrelated to sect. Duracini (Fig.
1, 2). Homoplasy is reinforced by our finding that both species
encompass collections with or without blue pigments (Table 3).
The CFP version of C. dolabratus is part of /Bicolores and is
phylogenetically conspecific with C. imbutoides, a species with
obvious blue hues described as a typical Bicolores in the ADC.
Conversely, the version of C. dolabratus published in the ADC
falls, together with three other Duracini binomials, in the clade
of C. cinnamoviolaceus, of which it represents a collection lack-
ing blue colour (cf. above). The name is stabilized here in its
strict – and original – Nordic sense, through its neotypification
with the unpublished plate 181 and by epitypifying it with the
widely known and sequenced collection CFP990, illustrated on
plate D52 of the Scandinavian monograph. The intraspecific
polymorphism of C. dolabratus is the highest in the section
Fig. 5 Type material designated here. — a. Plate ined. 181 directed/approved by Fries, S, neotype (iconotype) of C. dolabratus; b. Atl. Tab. 377, f. 202 (1890),
lectotype (iconotype) of C. refectus; c. A. Bidaud 96-09-73, epitype of C. refectus; d. I. Kytövuori 97-1162, neotype of C. imbutus.
c
b
d
a

193
K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini
(3 substitutions, Table 3) but it should be considered with
respect to its wide biogeographical distribution and thorough
sampling (23 sequences analysed, Fig. 2). Its sister species,
C. dolabratoides, is distant by 3 substitutions (Table 3). Cor-
tinarius dolabratus and C. cinnamoviolaceus share similar
ecological niches and can both produce basidiomata with or
without blue hues. Fortunately, the distinction of the species
is usually fairly easy – the latter has a strong smell of radish,
its spores are, on average, larger than those of C. dolabratus,
and it is often also more robust than C. dolabratus (Table 3).
Cortinarius cinnamoviolaceus has so far been only found in
Europe whereas C. dolabratus displays a wide distribution
extending to western North America.
Cortinarius evernius (Fr.) Fr., Epicr. Syst. Mycol.: 294. 1838 —
MycoBank MB#233378
Basionym. ≡ Agaricus evernius Fr., Observ. Mycol. 2: 79. 1818: sanctioned
in Fr., Syst. Mycol. 1: 212. 1821.
≡ Hydrocybe evernia (Fr.) M.M. Moser, Kleine Kryptogamenflora von
Mitteleuropa II: 161. 1953.
≡ Telamonia evernia (Fr.) Ricken, Die Blätterpilze. 1915.
= Cortinarius evernius f. fragrans M.M. Moser ex Bidaud & Carteret, Atlas
des Cortinaires XXII: 1887. 2014.
= Cortinarius evernius f. pseudoscutulatus Rob. Henry ex Bidaud &
Reumaux, Atlas des Cortinaires XXII: 1887. 2014.
[= Cortinarius evernius var. insignis Fr., Atlas des Cortinaires XXII: f. 1405.
2014, nom. inval. (no diagnosis, no type designated)].
[= Cortinarius parvulior Bidaud ad int., Atlas des Cortinaires XXII: f. 1418.
2014, nom. inval. (no diagnosis, no type designated)].
Type. Sweden, Ångermanland, Specksta, Härnösand, under coniferous
trees, 22 Sept. 1988, T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos
CFP792, S, neotype designated in Cortinarius Flora Photographica I (French
version), pl. A11 (1990), MycoBank MBT#372785. ITS sequence deposited
in GenBank under KX964331.
Illustrations — Bidaud et al. 2014: pl. 946–949 but also pl. 959
(as C. parvulior); Brandrud et al. 1990: pl. A11.
Taxonomic descriptions — Bidaud et al. 2014: f. 1404–1407
but also f. 1418 (as C. parvulior); Niskanen et al. 2012: 863;
Brandrud et al. 1990: pl. A11.
Notes — All contemporary authors seem to interpret this
widespread Friesian species the same way and, not consid-
ering infraspecific taxa and species described ad interim, no
later synonym of C. evernius has been introduced – however,
older authors like Konrad and Henry misapplied the name
to C. cinnamoviolaceus (see above). Phylogenetically, the
species displays very low intraspecific polymorphism despite
its wide biogeographical distribution (1 substitution plus one
length polymorphism out of 22 available sequences) and is
separated from its sister species C. cagei by 3 substitutions
plus 3 length polymorphisms (Table 3). In Europe, the species
may be confused only with C. cinnamoviolaceus, but the latter
strongly smells of radish, has smaller spores and displays a
much broader ecological range.
Cortinarius glaphurus Chevassut & Rob. Henry, Doc. Mycol.
12, 47: 78. 1982 — MycoBank MB#109708
= Cortinarius tubulosus Bidaud, Atlas des Cortinaires XXII: 1888. 2014.
= Cortinarius cedriosmus Bidaud, Atlas des Cortinaires XIX: 1510. 2010.
= Cortinarius violaeolens Carteret & Reumaux, Atlas des Cortinaires
XIX: 1509. 2010.
= Cortinarius paranomalus Rob. Henry, Atlas des Cortinaires IV: 105.
1992.
– Cortinarius turibulosus (Jul. Schäff. & E. Horak) Bon & G. Garnier,
Doc. Mycol. 21, 83: 10. 1991, sensu auct.
Type. FrAnce, Hérault, La Salvetat-sur-Agout, Lac de la Raviège, under
Picea, cespitous, 29 Oct. 1978, R. Henry 71421, PC, holotype, MycoBank
MBT#70172. ITS sequence deposited in GenBank under KX964352.
Illustrations — Bidaud et al. 2014: pl. 957 (as C. tubulosus);
2010: pl. 795 (as C. turibulosus), pl. 796 (as C. violaeolens) and
pl. 807 (as C. cedriosmus); 1992: pl. 83 (as C. paranomalus).
Taxonomic descriptions — Bidaud et al. 2014: f. 1414 (as
C. tubulosus) and 2010: f. 1108 (as C. turibulosus); Kühner
& Romagnesi 1953: 305 (as C. paranomalus); Chevassut &
Henry 1982: 78.
Notes — As redefined here, the concept of C. glaphurus
should be substantially widen so as to include those of C. ce-
driosmus, C. paranomalus, C. tubulosus and C. violaeolens,
as well as C. turibulosus sensu Bidaud et al. (2010). The proto-
logue should then be edited as follows: pileus diameter up to 55
mm, pileus dark chocolate-brown to reddish brown, not glabrous
and hygrophanous. Stipe not always straight nor isodiametric
but often (always?) hollow, with or without blue pigments and
with variable amounts of veil remnants that may form a mem-
branous ring. Often cespitose. Odour weakly raphanoid or of
cedar wood or viola. Associated with coniferous trees as well
as broad-leaved trees (Pinus, Quercus and Fagus confirmed
as hosts by ectomycorrhizal sequences). Phylogenetically, the
species is a bit polymorphic but is still well separated from its sis-
ter species C. tortuosus (Table 3). When collected under Picea
abies on calcareous soils and weakly smelling of cedar wood,
C. glaphurus may be difficult to distinguish from C. hircinosmus,
but the latter produces slightly smaller spores (Table 3). When
collected in hygrophilic and acidic soils under coniferous trees,
the species may be confused with C. tortuosus, but the latter
displays obvious blue tinges on the stipe, blood-red hues in the
gills, and is never cespitose.
Cortinarius hircinosmus Moënne-Locc., Atlas des Cortinaires
XII: 692. 2002 — MycoBank MB#489854
– Cortinarius livor Fr., Epicr. Syst. Mycol.: 306. 1838, sensu Bidaud et al.
(2015).
– Cortinarius scriptor Kühner, Doc. Mycol. 20, 77: 92. 1989, sensu Bidaud
et al. (2010) p.p.
Type. FrAnce, Haute-Savoie, Les Puisots, in Picea forest, elev. 700 m, 15
Sept. 1986, P. Moënne-Loccoz 334, PC, holotype, MycoBank MBT#101337.
ITS sequence deposited in GenBank under KX964368.
Illustrations — Bidaud et al. 2015: pl. 991 (as C. livor); 2002:
pl. 389.
Taxonomic descriptions — Bidaud et al. 2015: f. 1459 (as
C. livor); 2002: f. 575; Niskanen et al. 2012: 850.
Notes — This species has been initially described in subsect.
Hircini because of the strong smell of C. hircinus and C. cam-
phoratus of the holotype specimens. However, five additional
collections from France and Scandinavia, lacking such odour,
were later identified in the same clade. As revised here and
at least in France, C. hircinosmus fruits under Picea abies on
calcareous soils and includes the French concept of C. livor
and pro parte, that of C. scriptor. The original binomial is obvi-
ously unfortunate for an odourless or weakly smelling species,
so, provided additional collections confirm the strong smell of
some populations, infraspecific taxa may be introduced to more
adequately reflect the organoleptic diversity of the species.
Phylogenetically, the species is well resolved (Table 3). In the
field, C. hircinosmus may be confused with C. glaphurus (as
redefined here), but the latter displays a much broader ecologi-
cal niche, typically smells of cedar wood and has slightly larger
spores (9.3 × 5.2 µm vs 9 × 5 µm, on average).

194 Persoonia – Volume 39, 2017
Cortinarius plumulosus Rob. Henry, Bull. Soc. Mycol. France
93, 3: 362. 1977 — MycoBank MB#312090
– Cortinarius fundatus Britzelm., Ber. Naturhist. Vereins Augsburg 28:
127. 1885, sensu Bidaud et al. (2014).
Type. FrAnce, Vosges, Hennezel, in Abies forests, gregarious, 1972,
R. Henry 3417, PC, holotype, MycoBank MBT#155523. ITS sequence
deposited in GenBank under KX964374.
Illustrations — Bidaud et al. 2014: pl. 954 (as C. fundatus).
Taxonomic descriptions — Bidaud et al. 2014: f. 1411 (as C. fun-
datus); Henry 1977: 359.
Notes — This conifer-associated species has been treated in
the ADC as C. fundatus, and suspected by French authors, on
the basis of frequent macrospores up to 12 µm long observed
in some collections, to represent the second C. bicolor of
Cooke – the one with large spores and possible fruiting under
coniferous trees (cf. notes under C. cagei). Phylogenetically,
C. plumulosus is well separated from its closest neighbour C. ever-
nius (7 substitutions plus 3 indels, Table 3). Morphologically,
the species resembles C. refectus and C. cagei but the former
produces ovoid spores (Av Q = 1.5), the latter fruits under de-
ciduous trees and the cap of C. plumulosus is typically covered
by small flakes that are not found on that of its two lookalikes.
Cortinarius refectus Britzelm., Ber. Naturhist. Vereins Augs-
burg 28: 127. 1885 — MycoBank MB#560269; Fig. 5b – c
≡ Cortinarius reflectus Britzelm., Ber. Naturhist. Vereins Augsburg 28:
127. 1885.
– Cortinarius scriptor Kühner, Doc. Mycol. 20, 77: 92. 1989, sensu Bidaud
et al. (2010) p.p.
Misapplied name
– Cortinarius testaceoviolaceus Rob. Henry, Bull. Soc. Mycol. France 73, 1:
51. 1957, sensu Bidaud et al. (2014).
Type. Atl. Tab. 377, f. 202 (1890), lectotype (iconotype) designated here
(Fig. 5b), MycoBank MBT#373158. GermAny, Lombach, in Picea and Abies
forest, on calcareous soil, elev. 600 m, 24 Sept. 1996, A. Bidaud 96-09-73,
epitype designated here (Fig. 5c), MycoBank MBT#373159. ITS sequence
deposited in GenBank under KX964385.
Illustrations — This study: Fig. 5c; Bidaud et al. 2014, pl. 952,
953 but also pl. 945 (as C. testaceoviolaceus).
Taxonomic descriptions — Bidaud et al. 2014: f. 1410 but also
2010: f. 1109 (as C. scriptor).
Notes — No original material was kept by Britzelmayr to as-
sign C. refectus a molecular signature. The diagnosis is not very
elaborate but the atypical reported ovoid spores (8–9 × 5–6
µm, Av Q = 1.5) prompted the authors of the ADC to resurrect
this old binomial as their best candidate to the original – i.e., the
one with short spores (cf. notes under C. cagei ) – C. bicolor.
Although the latter hypothesis cannot be supported here for
ecological reasons, the French interpretation of C. refectus
does not contradict the protologue and it is compatible with
the original plate – although spore drawings on that plate do
not really support the protologue. We thus stabilize here the
name by lectotypifying it with plate n° 202, and epitypifying it
with the sequenced AB 96-09-73 collection from Germany. As
delineated here, C. refectus includes the ADC interpretations
of C. scriptor (p.p.) and C. testaceoviolaceus. The latter name
is, however, misapplied because the holotype of C. testaceo-
violaceus falls outside Telamonia (in subg. Myxacium, data
not shown). Phylogenetically, C. refectus is well resolved but
in the field, it could easily be confused with C. plumulosus and
C. cagei until spores examination and host trees are carefully
considered (Table 3).
Cortinarius tortuosus (Fr.) Fr., Epicr. Syst. Mycol.: 305. 1838
— MycoBank MB#165676
Basionym. ≡ Agaricus tortuosus Fr., Syst. Mycol. 1: 235. 1821.
≡ Hydrocybe tortuosa (Fr.) Wünsche, Die Pilze. Eine Anleitung zur
Kenntniss derselben: 121. 1877.
= Cortinarius flabelloides Carteret, Atlas des Cortinaires XIX: 1510. 2010.
= Cortinarius laetior P. Karst., Bidrag Kannedom Finlands Natur Folk 32:
387. 1879.
Type. Sweden, Smoland, Femsjö, Södra Färgen, Gatebäck, among Sphagnum
in spruce forest, 11 Sept. 1979, D. Lamoure, IB 79/533, neotype designated
in Opera Botanica 100: 182. 1989, MycoBank MBT#372784. ITS sequence
deposited in GenBank under KX964391.
Illustrations — Bidaud et al. 2014: pl. 955–956 but also 2010:
pl. 804 (as C. flabelloides); Brandrud et al. 1990: pl. A06.
Taxonomic descriptions — Bidaud et al. 2014: f. 1413 but also
2010: f. 1136 (as C. flabelloides); Niskanen et al. 2012: 863;
Brandrud et al. 1990: pl. A06.
Notes — This Friesian name has been interpreted in rather
similar ways by past and modern mycologists – with the no-
table exception of J. Favre, who referred to this species as
C. plumbosus – so that C. tortuosus taxonomy is not a proble-
matic issue. The species can be diagnosed by its narrow
ecological niche (hygrophilous and acidic soils, with conifer
trees) and the special purple-red tinges of the gills that tend to
darken upon bruising. The odour is usually reported as null or
weak of cedar wood but the conspecificity with C. flabelloides,
revealed in this work, indicates that basidiomata can also smell
of geosmin (i.e., of earth or dust, as C. variecolor for instance).
Phylogenetically, the species is remarkably stable at the ITS
locus and is well separated from its sister species C. glaphurus
(Table 3).
Cortinarius turgidipes Rob. Henry ex Rob. Henry, Atlas des
Cortinaires XVII, 1: 1179. 2008 — MycoBank MB#533088
Type. FrAnce, Creuse, Lavaud, under Picea, on granitic soil, subcespi-
tose, 19 Oct. 1993, A. & E. Bidaud, AB 93-10-425, PC, holotype, MycoBank
MBT#372786. ITS sequence deposited in GenBank under KX964409.
Illustration — Bidaud et al. 2008, pl. 672.
Taxonomic description — Bidaud et al. 2008: f. 885.
Notes — More collections of this species, originally described
in sect. Damasceni by its authors, are required to better as-
sess its morphogenetic variability as well as to define its eco-
logical niche. In its current sampling – limited to the holotype,
C. turgidipes is closest to C. dolabratus, from which it differs by
3 substitutions and 4 indels at the ITS locus (Table 3).
Cortinarius sect. Saturnini Rob. Henry ex Möenne-Locc. &
Reumaux, Atlas des Cortinaires I: 21 (1990), emend.
Type. Cortinarius saturninus (Fr.) Fr., Epicr. Syst. Mycol.: 306. 1838.
≡ Cortinarius subsect. Saturnini Bidaud, Moënne-Locc. & Reumaux, Doc.
Mycol. 24, 95: 41. 1994.
≡ Cortinarius sect. Firmiores (Fr.) Henn., in Engler & Prantl, Naturl.
Pfanzenf. I, 181: 246. 1900, p.p.
Notes — As revised here, sect. Saturnini is widely distributed
in the Northern Hemisphere and includes 6 species. They are
medium-sized, rarely stout Telamonia species, pale ochra-
ceous, brown to reddish brown, lilac-violet, hygrophanous, with
or without blue tinges in young lamellae and the upper part of the
stipe, with various amounts of veil remnants on the stipe and on
the pileus margin where it often forms a continuous covering or
discontinuous patches. Smell indistinct or weak. Spores broadly
or narrowly ellipsoid, (6–)6.5 –11(–14.4) × (3–) 4 –7(– 8.5) µm
(on average: 8.6 × 4.9 µm), verrucose. Gregarious to densely

195
K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini
cespitose, rarely solitary, typically fruiting under hygrophilous
deciduous trees (Salix, Populus, Betula) but also under Quercus
and Cistus in the Mediterranean area, rarely under coniferous
trees.
Cortinarius saturninus (Fr.) Fr., Epicr. Syst. Mycol.: 306. 1838
— MycoBank MB#177635
Basionym. ≡ Agaricus saturninus Fr., Syst. Mycol. 1: 219. 1821.
= Cortinarius fulvorimosus Carteret & Reumaux, Atlas des Cortinaires
XVII, 1: 1178. 2008.
= Cortinarius cohabitans var. urbicoides Bidaud & Fillion, Bull. Soc. Mycol.
France 119, 1–2: 70. 2004.
= Cortinarius urbicus var. sporanotandus Bidaud & Fillion, Atlas des
Cortinaires XII: 695. 2002.
= Cortinarius denseconnatus Rob. Henry, Bull. Soc. Mycol. France 99,
1: 65. 1983.
= Cortinarius gramineus Rob. Henry, Bull. Soc. Mycol. France 99, 1: 64.
1983.
= Cortinarius rastetteri Rob. Henry, Bull. Soc. Mycol. France 97, 3: 177.
1981.
= Cortinarius dissidens Reumaux, Bull. Soc. Mycol. France 96, 3: 356.
1980.
= Cortinarius marginatosplendens Reumaux, Bull. Soc. Mycol. France
96, 3: 356. 1980.
= Cortinarius salicis Rob. Henry, Bull. Soc. Mycol. France 93, 3: 364.
1977.
= Cortinarius umbrinoconnatus Rob. Henry, Bull. Soc. Mycol. France 73,
1: 53. 1957.
[= Cortinarius dionisiae Bidaud ad int., Atlas des Cortinaires XXIII: f. 1451.
2015, nom. inval. (no diagnosis, no type designated)].
– Cortinarius subtorvus Lamoure, Schweiz. Z. Pilzk. 47, 9: 169. 1969,
sensu auct.
– Cortinarius bresadolae Schulzer, Hedwigia 24, 4: 138. 1885, sensu
Lamoure (1978).
– Cortinarius cohabitans P. Karst., Bidrag Kannedom Finlands Natur Folk
32: 388. 1879, sensu auct.
– Cortinarius urbicus (Fr.) Fr., Epicr. Syst. Mycol.: 293. 1838, sensu Bidaud
et al. (2002) p.p.
Type. Sweden, Västergötland, Eggby, Drottningkullen, deciduous forest on
calcareous ground (Corylus, Tilia, Quercus), 17 Sept. 1986, T.E. Brandrud,
H. Lindström, H. Marklund, S. Muskos CFP514, S, neotype designated here,
MBT#373160. ITS sequence deposited in GenBank under KX964584.
Illustrations — Bidaud et al. 2015: pl. 983–989; Brandrud et al.
1994: pl. C09, but also 1990: pl. A04 (as C. subtorvus).
Taxonomic descriptions — Bidaud et al. 2015: f. 1448–1457;
Niskanen et al. 2012: 847–848; Brandrud et al. 1994: pl. C09,
but also 1990: pl. A04 (as C. subtorvus).
Notes — All contemporary and past authors agree on the
fact that C. saturninus is a collective species, that Fries him-
self contributed to confuse through multiple diagnoses across
his successive monographs, which, in addition, do not fit the
plates he later directed. The French mycologist Robert Henry
devoted decades of his life trying to sort out this complex,
adding to the literature many new names and interpretations
(for review, see Bidaud et al. 2015). The simplest way to clarify
this issue would undoubtedly be to consider C. saturninus as
a nomen dubium and readily discard it. However, the wide
use of the name that pertained throughout modern literature
and the general consensus about the species illustrated on
the plate C09 of the CFP, prompted us to fix C. saturninus in
its current, Nordic concept, through the neotypification of the
name with the CFP514 collection. Our phylogenetic analysis
reveals a tremendously polymorphic species, with no less than
9 holotypes previously thought to be unrelated to sect. Saturnini,
falling as later synonyms of C. saturninus. Cortinarius subtorvus
and C. cohabi tans, usually considered as akin to C. saturni-
nus, are most likely two additional synonyms, although their
respective type material could not be sequenced to ascertain
conspecificity. This work also establishes that C. oxytoneus,
considered by Henry as the most typical form of C. saturninus,
is evolutionarily unrelated to sect. Saturnini (sect. Duracini; Fig.
1). As revised here, C. saturninus displays highly apparent ITS
sequence polymorphism (Dintra max = 7 substitutions + 3 indels;
Table 3) but the latter is essentially driven by two Norwegian
(Svalbard) sequences for which no trace file is available. In
addition, the one substitution segregating a subclade within
the lineage (see * in Fig. 3) could demonstrably be attributed
to intra-individual polymorphism. Thus, the unbiased Dintra max
in C. saturninus is actually of 4 nt changes, a value that stems
from three French collections (PML 75 in one hand and AB
04-10-344 and XC 2002-167 in the other) which may deserve
taxonomic autonomy – at the infraspecific rank – when more
thoroughly sampled (Table 3, Fig. 3). Although its suspected
association with Salix is here demonstrated by the presence in
the clade of several ectomycorrhizal sequences isolated from
willow roots (within SH094324.07FU, Table 2, Fig. 3), C. sa-
turninus may also be associated with other deciduous, but also
coniferous trees. Morphologically, the species displays unprece-
dented levels of variability that represent a serious issue for
field diagnosis. Practically, one should consider C. saturninus
as a possible hit – and check the numerous aspects of this
species in the last release of the ADC for instance (Bidaud
et al. 2015) – whenever collecting a cespitose or gregarious
medium-size Telamonia: i) under Salix spp. or other hygro-
philous deciduous trees (and Dryas octopetala in the alpine
zone), with or without blue hues at the stipe apex and with veil
remnants ranging from none to white patches or covering at the
cap margin, to copious and web-like covering the whole young
fruit body; or ii) under coniferous trees and in this case with a
ring and with short (L < 10 µm), ellipsoid spores. Highest risks
of confusion are with other members of the revised sect. Sa-
turnini (see notes under C. confirmatus, C. cyprinus and C. im-
butus), and, for blue-lacking and densely veiled basidiomata
collected under Salix spp. (referred to as C. saturninus ‘aspect’
salicis, ‘aspect’ urbicoides and ‘aspect’ sporanotandus in the
ADC), with C. urbicus. The latter species displays more whit-
ish hues on the fresh pileus and is typically less hygrophanous
than C. saturninus, with no ‘Kuehneromyces-like’ dehydration.
Cortinarius confirmatus Rob. Henry, Bull. Soc. Mycol. France
99, 1: 67. 1983 — MycoBank MB#818598 (var. confirmatus);
MycoBank MB#818597 (var. plesiocistus)
= Cortinarius assiduus var. plesiocistus A. Ortega et al., Mycotaxon 101:
140. 2007.
= Cortinarius assiduus Mahiques, A. Ortega & Bidaud, Bull. Féd. Mycol.
Dauphiné-Savoie 162: 42. 2001.
= Cortinarius bulbosovolvatus Rob. Henry & Contu, Doc. Mycol. XVI, 61:
32. 1985.
[= Cortinarius kuehneri Bidaud ad int., Atlas des Cortinaires XXIII: f. 1440.
2015, nom. inval. (no diagnosis, no type designated)].
[= Cortinarius spurcatocephalus Carteret ad int., Atlas des Cortinaires
XXIII: f. 1439. 2015, nom. inval. (no diagnosis, no type designated)].
[= Cortinarius paracohabitans Bidaud ad int., Atlas des Cortinaires XXIII:
f. 1437. 2015, nom. inval. (no diagnosis, no type designated)].
[= Cortinarius subcylindratus Carteret ad int., Bull. Soc. Mycol. France
128(3– 4): 280. 2014, nom. inval. (no diagnosis, no type designated)].
[= Cortinarius cistoadelphus Bidaud ad int., Bull. Féd. Assoc. Mycol. Mé-
diterranéennes 6: 41 (1994), nom. inval. (no diagnosis, no type designated)].
– Cortinarius cypriacus Fr., Epicr. Syst. Mycol.: 307. 1838, sensu Consiglio
(1999) non Moënne-Loccoz & Reumaux (1989).
Type. FrAnce, unknown locality and collection date, under Quercus ilex,
R. Henry 3195, PC, holotype, MycoBank MBT#69663. ITS sequence depo-
sited in GenBank under KX964438.
New combination. Cortinarius confirmatus var. plesiocistus (A. Ortega,
Vila & Bidaud) Carteret, Bidaud, Reumaux & Bellanger, comb. nov.
Basionym. Cortinarius assiduus var. plesiocistus A. Ortega, Vila & Bidaud
in Ortega et al., Mycotaxon 101: 140. 2007. ITS sequence deposited in
GenBank under AM713178.

196 Persoonia – Volume 39, 2017
Illustrations — Bidaud et al. 2015: pl. 970–973; Ortega et al.
2007: pl. 2; Mahiques et al. 2001.
Taxonomic descriptions — Bidaud et al. 2008: f. 1434–1441;
Ortega et al. 2007: 140; Mahiques et al. 2001: 42; Henry 1983:
67.
Notes — In its original concept, C. confirmatus is a cespitose
species without blue tinges, fruiting in Mediterranean Quercus
ilex woodlands, included by Henry in his sect. Damasceni. As
phylogenetically redefined here, the species concept is dramati-
cally widened both morphologically and ecologically, so as to
encompass 7 former morphologically delimited species and one
variety, caespitose or not, with or without blue hues, and occur-
ring in the Mediterranean area under Quercus spp. or Cistus
spp., but also in temperate continental forests, under various
deciduous trees as well as under Picea abies. The presence of
two ectomycorrhizal sequences from Northern Iran in the clade
considerably extends the known geographic distribution of the
species, that may occur across a broad Eurasiatic belt. The
clade displays the highest sequence variability within the section
(Dintra max = 6 nt changes, Table 3) and its topology delineates
3 supported subclades that may, in principle, deserve their own
taxonomic autonomy, as well as C. cistoadelphus Bidaud ad
int. (Fig. 3). The infraspecific rank should be favoured for such
distal lineages because:
i. electing these subclades at the species level would leave 8
basal sequences unresolved, in paraphyletic relationships
with the 3 recognized species;
ii. two of the resulting species would be totally cryptic, as none
of the morphological, ecological or geographical features
identified in the inclusive clade segregate into the two
relevant subclades; and
iii. the third subclade, which overlaps with the cisticolous
C. as siduus var. plesiocistus and C. bulbosovolvatus, has
already been assigned a varietal rank, on morphogenetic
bases (Ortega et al. 2007).
Thus, in a conservative approach and following an integrative
method of species limits delineation, here we define C. con-
firmatus within the boundaries of its most inclusive clade and
introduce C. confirmatus var. plesiocistus (A. Ortega, Vila &
Bidaud) comb. nov. to accommodate the cisticolous popula-
tions. Future studies may unveil cryptic criteria to diagnose the
two other subclades. When collected under meridional oaks or
Cistus spp., C. confirmatus cannot be misidentified as one of
the other Saturnini members, as none of the latter have so far
been reported in the Mediterranean area. However, in more
continental locations, especially in mixed deciduous forests,
the species may co-occur with C. saturninus, C. imbutus and
C. cyprinus and the risk of confusing these taxa is high. In this
biome, C. confirmatus differs from its morphogenetic lookalikes
by one of the following combinations of features:
i. absence of veil remnants on the stipe and not fruiting
densely cespitose; or
ii. abundant veil remnants on the stipe and densely cespitose
under Populus alba (‘aspect’ paracohabitans); or
iii. reddish hues on the cap and densely cespitose under Betula
pendula (‘aspect’ rubricosissimus).
Cortinarius cyprinus Bidaud, Carteret & Reumaux, Atlas des
Cortinaires XXIII: 1981. 2015 — MycoBank MB#815172
[= Cortinarius saturninus var. bresadolae M.M. Moser, Kleine Kryptoga-
menflora von Mitteleuropa II: 162. 1953, nom inval. (ined.)].
– Cortinarius cypriacus Fr., Epicr. Syst. Mycol.: 307. 1838, sensu Moënne-
Loccoz & Reumaux (1989), non Consiglio (1999).
Type. FrAnce, Yvelines, Gambais, under deciduous trees, on calcare-
ous soil, 3 Oct. 1993, G. Redeuilh, XC 2012-26, PC, holotype, MycoBank
MBT#373189. ITS sequence deposited in GenBank under KX964463.
Illustration — Bidaud et al. 2015: pl. 973–976.
Taxonomic description — Bidaud et al. 2015, f. 1443.
Notes — This recently described species used to be called
C. saturninus var. bresadolae or C. cypriacus by French authors
but in the field, C. cohabitans (= C. saturninus) and C. circum-
velatus (= C. lucorum) are likely the first names that come to
the collectors’ mind, due to the crown-like veil remnants at the
pileus margin, violet hues in young lamellae and gregarious
fruiting under hygrophilous deciduous trees. However, mo-
lecular analysis of the large herbarium of the authors of the
ADC unveiled phylogenetic autonomy of a subset of collections
that differ from other Saturnini members by very reduced veil
remnants on the stipe that never form a ring, and occurrence
so far restricted to calcareous soils. As currently sampled, the
species seems rather widespread in France but it has been
rarely reported elsewhere, as it is represented by a single col-
lection from southern Norway and possibly an additional one
from Estonia (TAAM128765/ UDB016164). Phylogenetically,
C. cyprinus is sister to C. saturninus, from which it differs by
3 substitutions and 2 indels (Table 3). The ITS sequence of the
French collections and of the Norwegian collection are 100 %
identical, and they differ from the Estonian sequence by substi-
tutions. The lack of publically available trace file for UDB016164
prevents us from critically examining these polymorphisms and
the possible conspecificity of TAAM128765 with C. cyprinus.
Further taxon sampling and sequencing of Estonian Saturnini
collections will be necessary to clarify this issue and to better
estimate the intraspecific variability of the species at the ITS
locus.
Cortinarius imbutus Fr., Epicr. Syst. Mycol.: 306. 1838 — Myco-
Bank MB#233557; Fig. 5d
= Cortinarius laccatus Reumaux, Bull. Soc. Mycol. France 98, 4: 348.
1982.
= Cortinarius betulaecomes Rob. Henry, Bull. Soc. Mycol. France 93, 3:
347. 1977.
[= Cortinarius saturnalis Reumaux ad int., Atlas des Cortinaires XXIII:
f. 1446. 2015, nom. inval. (no diagnosis, no type designated)].
Type. FinlAnd, Perä-Pohjanmaa, Tornio, Arpela, Runteli, rich grass-herb
spruce forest with deciduous bushes and some pines, slightly paludified
depressions, calcareous ground, 10 Sept. 1997, I. Kytövuori 97-1162, H,
neotype designated here, MycoBank MBT#373161 (Fig. 5d). ITS sequence
deposited in GenBank under KX964498.
Illustrations — This study: Fig. 5d; Bidaud et al. 2015: pl.
976–982.
Taxonomic descriptions — Bidaud et al. 2015: f. 1445 –1447.
Notes — The two major contemporary interpretations of
C. imbutus are in marked contrast, as the CFP authors consider
the species in sect. Bicolores, while those of the ADC place it
in sect. Saturnini. The Friesian diagnosis of C. imbutus is, as
often with old names, not precise enough to support a single,
unequivocal interpretation. However, Fries described his spe-
cies between C. saturninus and C. cypriacus, indicating that
the original concept would be naturally placed in sect. Saturnini.
Our work reveals that the French version of C. imbutus is one
of the morphogenetic Saturnini, widely distributed across the
northern hemisphere, whereas the CFP one corresponds to a
blue-pigmented collection of C. cinnamoviolaceus (and is then
conspecific with the French C. dolabratus, see notes under this
species). We thus here stabilize the name in the revised sect.
Saturnini, by neotypifying it with the sequenced IK97-1162
collection from Finland. Phylogenetically, C. imbutus is rather
polymorphic at the ITS locus (Dintra max = 3 substitutions + 1
indel) and simultaneously very close from its closest species
C. confirmatus (Dinter min = 3 nt changes, Table 3). Morphologi-

197
K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini
cally, C. imbutus is quite variable, especially regarding the col-
our of the pileus and the intensity of blue tinges in basidio mata.
Typically, the species fruits under deciduous trees in hygrophil-
ous places but collections (referred to as C. imbutus ‘aspect’
laccatus in the ADC) have been reported in pure coniferous
forests. In the field, C. imbutus may easily be confused with
C. confirmatus, C. cyprinus and most notably C. saturninus,
which can occur in similar habitats. Combining the 3 following
criteria – not diagnostic on their own – should help identifying
C. imbutus from its evolutionary siblings:
i. the lilac-greyish, not violaceous, hues of young lamellae;
ii. elongated spores (Av Q > 1.8, Table 3); and
iii. copious veil remnants on the stipe.
Macrochemistry may be useful as well to distinguish C. imbutus
from C. confirmatus (gaïacol and silver nitrate), although the
reliability of these reactions is still questionable.
Cortinarius lucorum (Fr.) Berger, Cat. Herb. III: 89. 1846 —
MycoBank MB#818604
Basionym. ≡ Cortinarius impennis var. lucorum Fr., Epicr. Syst. Mycol.:
294. 1838.
≡ Hydrocybe lucorum (Fr.) M.M. Moser, Kleine Kryptogamenflora von
Mitteleuropa II: 162. 1953.
≡ Cortinarius lucorum (Fr.) Mussat: 101. 1901.
≡ Cortinarius impennis subsp. lucorum (Fr.) Sacc.: 951. 1887.
= Cortinarius incarnatolilascens Rob. Henry, Bull. Soc. Mycol. France 97,
3: 170. 1981.
= Cortinarius montis-dei Reumaux, Bull. Soc. Mycol. France 96: 357.
1980.
= Cortinarius circumvelatus Reumaux, Bull. Soc. Mycol. France 96: 355.
1980.
? = Cortinarius umidicola Kauffman, Bull. Torrey Bot. Club 32, 6: 322.
1905.
Type. NorwAy, Vestfold, Moss, Jelöy, under Populus tremula, 13 Sept.
1986, T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos CFP490, S,
neotype designated here, MycoBank MBT#373173. ITS sequence deposited
in GenBank under KX964585.
Illustrations — Bidaud et al. 2015: pl. 967–969; Brandrud et
al. 1994: pl. C10.
Taxonomic descriptions — Bidaud et al. 2015: f. 1428–1431;
Niskanen et al. 2012: 847; Brandrud et al. 1994: pl. C10;
Matheny & Ammirati 2006.
Notes — In Nordic countries, this widespread species is
tightly associated with Populus spp. and it is well known, in
large part thanks to the plate C10 published in the CFP. North
American mycologists, following Kauffman’s footsteps, some-
times name this species C. umidicola, even though the latter
binomial has been originally applied to a mushroom fruiting in
conifer forests, e.g., Tsuga (Kauffman 1932). French authors
described it repeatedly, as C. circumvelatus, C. incarnatoli-
lascens and C. montis-dei, on the basis of deviating macro-
morphological or ecological features while oddly, their ini-
tial – pre-molecular – concept of C. lucorum does not belong to
/Saturnini (cf. C. cypriacoides in Fig. 1). Fries does not mention
violaceous tinges on the stipe nor the typical crown-like veil
in the protologue and he does not give much detail about the
lamellae. However, his concept does not contradict the contem-
porary one in use in Nordic countries, so in order to stabilize
C. lucorum, we here neotypify the name with the sequenced
Norwegian collection CFP490 of plate C10. Our phylogenetic
analysis slightly alters the morphological definition of the spe-
cies (see above) and provide information on its biogeography
and its extended ecological niche. Indeed, as revised here,
C. lucorum can be collected under Populus spp. – with proven
association through ectomycorrhizal sequences found in the
clade – on both continents, but it also fruits under other hygro-
philous deciduous trees, at least in France and, more surpris-
ingly, under Tsuga and Picea. Phylogenetically, the species is
well separated from the rest of Saturnini members, with a Dinter
min far exceeding Dintra max (Table 3). Interestingly, the topo-
logy of the clade segregates, by a 1 substitution each; i) North
American populations from European ones; and ii) European
populations fruiting under deciduous trees from the ones fruit-
ing under coniferous trees – referred to as C. lucorum ‘aspect’
incarnatolilascens in the ADC. Such finding, if confirmed by
further sampling, would support the autonomy of concerned
collections at an infraspecific rank. The identity of C. umidicola
with C. lucorum remains provisional because the sequence
we obtained from Kauffman’s syntype encompasses only the
ITS1 domain. Thus, although 100 % identical to the Populus-
associated Canadian collection TN10-002 along this part of
the ribosomal locus (the basal-most and unsupported branch
of the clade in Fig. 3 is artefactual and likely results from the
shorter sequence of C. umidicola), one cannot preclude ad-
ditional differences to take place in the ITS2 domain, splitting
the two species apart. When occurring under Populus spp.
or other hygrophilous broadleaved trees, and considering the
massive fruiting and typical crown-like veil, C. lucorum might
only be confused with C. cyprinus and C. saturninus, but these
species are usually less robust and their spores are much
smaller (Table 3).
Cortinarius stuntzii S.A. Rehner & Ammirati, Mycologia 80,
6: 903. 1988 — MycoBank MB#135248
Type. UsA, Washington, Grant County, Crab Creek, 5 Nov. 1981, S.A.
Rehner 394, WTU, holotype, MycoBank MBT#78780. ITS sequence depo-
sited in GenBank under KX964558.
Illustration — Rehner et al. 1988: f. 1.
Taxonomic description — Rehner et al. 1988: 904 – 906.
Notes — This stout species densely fruiting under Salix
exigua and S. rigida, so far known only from a small location
of North-western USA, was compared to C. umidicola and
C. subtorvus in the original publication, compatible with a place-
ment into sect. Saturnini. However, a positioning elsewhere
in subg. Telamonia, or even in subg. Sericeocybe – due to its
low hygrophaneity – has also been invoked. The present work
unambiguously establishes C. stuntzii as a genuine Saturnini,
phylogenetically most closely related to C. saturninus, from
which it differs by 3 substitutions and 5 indels (Table 3). Not
considering biogeography, so far restricted to the type locality,
the species is easily distinguished from other Saturnini mem-
bers by its unusually large spores, up to 14.4 µm long and 8.5
µm wide (on average: 11.5 × 6.7 µm, Table 3).
KEY TO SPECIES TREATED IN THE PRESENT STUDY
1. Alpine and arctic zone, under Salix spp. or Dryas octo-
petala .............................. C. saturninus
1. Mediterranean thermophilic area, under Quercus ilex or
Cistus spp. ......................... C. confirmatus
1. Continental zone ............................... 2
2. Coniferous trees ............................... 3
2. Deciduous trees ............................... 20
3. Acidic soils, in or near peatlands, Picea or Abies ...... 4
3. Dry to mesic acidic woodlands .................... 6
3. Basic to neutral, often calcareous woodlands ........ 13
4. Average spore length > 10 µm, blue tinges obvious, usually
odourless .............................C. evernius
4. Average spore length < 10 µm, usually smelling .......5

198 Persoonia – Volume 39, 2017
5. Average spore width > 5.2 µm, smell of cedar wood or
earthy.............................. C. tortuosus
5. Average spore width < 5.2 µm, smell of coconut.......
.................................. C. dolabratus
[with raphanoid smell, cf. C. cinnamoviolaceus]
6. Average spore width ≤ 5 µm .....................7
6. 5 µm < average spore width < 6 µm ...............8
6. Average spore width > 6 µm ............. C. refectus
7. Spores narrowly fusoid (Av Q > 1.8) and finely verrucose
................................C. dolabratoides
7. Spores elongated (1.7 < Av Q < 1.8) and strongly verru-
cose .............................. C. dolabratus
7. Spores ellipsoid (Av Q = 1.6) and strongly verrucose ...
.................................. C. saturninus
8. Tsuga, Pseudotsuga (North America) ...... C. lucorum
8. Picea, Abies, Pinus (Europe).....................9
9. Spores ovoid to ellipsoid (Av Q < 1.7)............. 10
9. More elongated spores (Av Q > 1.7) .............. 11
10. Average spore size < 9 × 5.5 µm, smooth pileus ......
...................................C. turgidipes
10. Average spore size > 9 × 5.5 µm, fibrillose pileus ......
.................................... C. lucorum
11. Average spore size < 9 × 5 µm ......... C. saturninus
11. Average spore size > 9 × 5 µm ..................12
12. Average spore width < 5.5 µm, smooth pileus ........
...................................C. glaphurus
12. Average spore width ≥ 5.5 µm, pileus covered with flakes
..................................C. plumulosus
13. Cespitose................................... 14
13. Not cespitose................................ 16
14. Strong veil remnants on the stipe ................15
14. Naked silky stipe .....................C. glaphurus
15. Average spore length < 8 µm .......... C. saturninus
15. Average spore length > 8 µm ......... C. confirmatus
16. Average spore length < 9 µm ............ C. imbutus
16. Average spore length ≥ 9 µm....................17
17. Average spore width > 6 µm ............. C. refectus
17. Average spore width < 6 µm ....................18
18. Average spore length > 9.5 µm, pileus covered with flakes
..................................C. plumulosus
18. Average spore length < 9.5 µm, smooth pileus...... 19
19. Smell of cedar wood ..................C. glaphurus
19. Smell weak or different ..............C. hircinosmus
20. Average spore width > 6 µm ....................21
20. 5 µm < average spore width < 6 µm ..............22
20. Average spore width ≤ 5 µm ....................24
21. Average spore length > 10.5 µm, Salix, USA . C. stuntzii
21. Average spore length < 10.5 µm, Fagaceae, Europe ...
.....................................C. refectus
22. Spores elongated (1.7 < Av Q < 1.8), smell of cedar wood
or Viola.............................C. glaphurus
22. Spores ovoid to ellipsoid (Av Q ≤ 1.7), smell null or differ-
ent ........................................23
23. Stout basidiomata, average spore size > 9.5 × 5.7 µm,
hygrophilous ......................... C. lucorum
23. Small to medium-size basidiomata, average spore size
≤ 9.5 × 5.7 µm .......................... C. cagei
24. Smell of cedar wood ................. C. dolabratus
24. Smell null or different.......................... 25
25. Spores ovoid (Av Q ≤ 1.6) ......................26
25. Spores ellipsoid (1.6 < Av Q < 1.7), orange hues on the
pileus ................................C. imbutus
25. Spores elongated to subcyndrical (1.7 ≤ Av Q ≤ 1.9) . 27
26. Naked silky stipe ................... C. confirmatus
26. Persistent veil remnants on the stipe .... C. saturninus
27. Densely cespitose . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
27. Gregarious or loosely cespitose .................29
28. Populus alba .......................C. confirmatus
28. Other deciduous trees, mostly Salix spp. ..C. saturninus
29. Persistent veil remnants on the stipe........C. imbutus
29. Naked silky stipe .............................30
30. Average spore length < 8.3 µm ............C. imbutus
30. 8.3 µm < average spore length < 8.6 µm...........31
30. Average spore length > 8.6 µm .........C. confirmatus
31. AgNO3: –..........................C. confirmatus
31. AgNO3: +............................ C. cyprinus
DISCUSSION
The present work significantly updates our knowledge of Cor-
tinarius, by revealing the number and the limits of species within
sections Bicolores and Saturnini. It also places phylogenetically
the morphological species described in these sections that do
not belong in /Bicolores or / Saturnini, illustrating the homoplasic
nature of morphological traits traditionally used to delineate
boundaries of these sections and their relations to other sec-
tions such as Bovini, Disjungendi, Duracini, Hydrocybe and
Sciophylli.
What do we learn about species?
The major advanced molecular tools bring to taxonomy the
ability to identify natural relationships between taxa, including
those previously regarded as unrelated, to reveal cryptic spe-
cies, and to correct species boundaries which were based on
the use of non-diagnostic morphological traits. Sequencing
numerous materials from sect. Bicolores and sect. Saturnini
as well as species falling outside these sections, we identified
10 morphogenetic species and 2 phylogenetic species in sect.
Bicolores, and 6 morphogenetic species in sect. Saturnini,
including C. cyprinus as a cryptic species. The sequencing of
type materials showed that 25 binomials are later synonyms
of the 15 revised names.
The limits of only two species – C. glaphurus and C. dolabratus
– in sect. Bicolores have been significantly altered after phylo-
genetic analysis, whereas all previously known species in sect.
Saturnini have been severely redefined following molecular
revision, except C. stuntzii, represented by only the holotype
collection. In most cases, several morphological species are
nested in single evolutionary units as a result of overreliance
in the past on often non-diagnostic morphological traits. The
presence of blue hues and the detection of a specific odour
are among the most misleading taxonomic features unveiled
in this work, as they have led to the erroneous autonomy of
C. assiduus, C. denseconnatus, C. gramineus, C. imbutoides,
C. phaeoruber, C. rastetteri, C. umbrinoconnatus, and C. ce-
driosmus, C. flabelloides, C. periodolens and C. violaeolens,
respectively. Pigments and volatiles of basidiomata, as the
products of the fungal secondary metabolism, are expected to
display some levels of variability in response to environmental
cues. Similarly, differences in the habit or abundance of veil
tissue on fruit bodies, that was used to segregate e.g., C. cir-
cumvelatus, C. fulvorimosus, C. parvulior or C. salicis from their
evolutionary lineages, might be explained by soil features or
weather conditions at, or preceding fruiting.
More surprising is our finding that spore size and ecology also
can be misleading, as illustrated by the lack of phylogenetic
autonomy of C. sporanotandus, which produces much smaller
spores than other C. saturninus collections, and C. deceptivus,
C. incarnatolilascens, C. laccatus or C. umidicola, which are

199
K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini
natural boundaries is instrumental in considering the revised
concept of C. dolabratus, here epitypified in the revised sect.
Bicolores despite the fact that all authors have initially placed
the species in sect. Duracini. The case of C. turgidipes also
illustrates this overlap of traditional sections, as the holotype of
this morphological Duracini nests within /Bicolores.
It should be concluded from these examples that the presence/
absence of blue pigments has been overemphasized in the
definition of all morphospecies cited above but also in that of
sections Bicolores, Saturnini and Duracini.
Strength and limits of integrative taxonomy
Higher Fungi systematics has been entirely built on the identifi-
cation and hierarchical organization of visible characteris-
tics – both macroscopic and microscopic, that were supposed
to be stable within a given taxon and which in combination
were supposed to be diagnostic of each species. The neces-
sarily subjective nature of the selection process involved in
this approach has led to highly artificial groupings at multiple
taxonomic levels (i.e., Aphyllophorales, Clavariaceae, Clito-
cybe, Gasteromycetes) and to divergences in the concept of
species that culminate in the genus Cortinarius. Unravelling
evolutionary history of Fungi through molecular phylogenies
had tremendously impacted taxonomy, in part because char-
acteristics that delinate a lineage with high taxonomic value
can now be distinguished from those, less valuable and taxo-
nomically overemphasized, which have appeared repeatedly
in distant branches of the fungal tree of life. However, if more
natural, the alternate organization of taxa that emerges from
these molecular analyses brings contemporary mycologists
the major challenge to uncover phylogenetically supported
sets of features that will be diagnostic of each morphogenetic
taxon. This process, especially in the species-richest genus
Cortinarius, is certainly the most time-consuming part of the
revision work and importantly, it heavily relies on the skills of
expert field taxonomists, not phylogeneticists.
Acknowledgements We are grateful to the curators of PC (Bart Buyck), IB
(Ursula Peintner, Regina Kuhnert) and S (Jens Klackenberg) for making sev-
eral reference collections available to us, as well as Karl Soop, Josep Ballarà
and Tor-Erik Brandrud for providing material from their personal herbaria. Part
of the molecular work (DNA extraction and PCR amplifications) was done at
the genetic markers in the ecology facility (SMGE) of the CEFE. The visit of
Bálint Dima in PC was financially supported by SYNTHESYS, the European
Union-funded Integrated Activities grant (application FR-TAF-4253). This work
was supported by the Ministry of Environment, Finland (YM38/5512/2009)
and the Swedish Taxonomy Initiative (dha 165/08 1.4).
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all associated with different host trees within their respective
clades. Spores and host plants are usually considered as reli-
able elements for taxonomic purposes because anatomy of the
reproductive structures and the complex molecular machinery
involved in mycorrhizal recognition are expected to have higher
selective pressure when compared to macroscopic features,
which are more prone to homoplasia. Part of our findings may
be explained by abnormal individuals or spectacular ecological
plasticity of species in sect. Saturnini, but the relatively high
levels of polymorphism revealed in C. confirmatus rather sup-
port on-going and cryptic speciation in this lineage. Thus, we
believe species limits delineated in the present work, especially
in the revised sect. Saturnini, are more conservative than what
short interspecific phylogenetic distances may suggest.
What do we learn about sections limits?
The segregation of sect. Saturnini within Cortinarius has been
intricately linked to that of separating subg. Hydrocybe from
subg. Telamonia, on the basis of the presence or absence of
veil remnants on the stipe (Moënne-Loccoz & Reumaux 1990).
However, such splitting is not phylogenetically supported, mak-
ing Hydrocybe an artificial grouping and stipe ornamentation a
confounding taxonomic criterion within Telamonia. As a result,
species previously described in sect. Saturnini are not expected
to form a single monophyletic lineage but are rather likely to
share evolutionary history with members of other sections in
the subgenus, especially the blue-coloured species from sect.
Bicolores. Consistently, only 5 out of the 14 species recently
described in sect. Saturnini in the ADC belong in that section.
The remaining morphological species are distributed across
Telamonia and illustrate the overlap of the original section with
sect. Sciophylli (C. saturninoides), defined to accommodate
very similar blue taxa, but more hygrophanous than genuine
Saturnini, and revised sect. Bovini (C. cypriacoides, C. illepidus
and C. subfirmus), so far not supposed to include blue Tela-
monia species. Species previously included in sect. Saturnini
also displayed obvious common features with sect. Duracini,
as assessed by the presence of C. oxytoneus, considered by
Henry as one of Fries’ C. saturninus, in sect. Duracini (Fig. 1).
Similar but somewhat reversed cases are the presence in the
revised sect. Saturnini of C. confirmatus, C. denseconnatus
and C. fulvorimosus, originally described in sect. Duracini. The
expected overlap of morphological characters in sect. Saturnini
and sect. Bicolores is best illustrated by the case of C. laetior
P. Karst., placed by its author in the trilogy saturninus-imbutus-
cypriacus, but shown here to belong in sect. Bicolores (Fig. 2).
Interestingly, the present work yields strong phylogenetic sup-
port to the prospective placement or overlap of the morphologi-
cally defined sect. Bicolores and sect. Duracini. Natural rela-
tionships or transitions between these two sections have long
been commented on by classical authors, on the basis of very
similar habits and the suspected weakness of the ‘blue colour’
criterion in Cortinarius systematics (Melot 1990, Frøslev et al.
2007). However, the issue was virtually impossible to address
in the absence of molecular data and the revision of C. cin-
namoviolaceus here sheds decisive light on this issue. Indeed,
although not part of /Bicolores and phylogenetically unrelated to
sect. Duracini, this species is built from concepts that typically
belong in traditional Bicolores (C. cinnamoviolaceus, C. parev-
ernius, and C. imbutus sensu CFP), in traditional Duracini (C.
contractus, C. cylindratus, C. subparevernius and C. dolabratus
sensu ADC), or somewhere in between the two sections (C.
basicyaneus). This unexpected assemblage within a single evo-
lutionary species somehow cracks the code of the secret dialog
between the two sections, revealing the totally artificial nature
of their main diagnostic feature, i.e., the presence/ absence of
blue pigments in fruit bodies. Knowing C. cinnamoviolaceus

200 Persoonia – Volume 39, 2017
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