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

Balint Dima

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.

Persoonia 39, 2017: 175 – 200 www.ingentaconnect.com/content/nhn/pimj RESEARCH ARTICLE ISSN (Online) 1878-9080 https://doi.org/10.3767/persoonia.2017.39.08 Cortinarius section Bicolores and section Saturnini (Basidiomycota, Agaricales), a morphogenetic overview of European and north American species K. Liimatainen1, X. Carteret 2, B. Dima3,4, I. Kytövuori 5, A. Bidaud 6, P. Reumaux7, T. Niskanen1, J.F. Ammirati 8, J.-M. Bellanger 9 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. Cortinarius is the largest genus of ectomycorrhizal fungi worldwide, 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 species, 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 morphoanatomical 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 morphological species, has been instrumental in delineating objective boundaries to taxa, and when applied to type material, stabilizes 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. Jodrell Laboratory, Royal Botanic Gardens, Kew, Surrey, TW9 3AB, United Kingdom. 68, rue Alexis Maneyrol, F-92370 Chaville, France. Department of Plant Anatomy, Institute of Biology, Eötvös Loránd University, Pázmány Péter sétány 1/c, H-1117 Budapest, Hungary. Department of Biosciences, Plant Biology, P.O. Box 65, FI-00014 University of Helsinki, Finland. Botanical Museum, University of Helsinki, Helsinki, P.O. Box 7, FI-00014 Finland. 2436, route de Brailles, F-38510 Vézeronce-Curtin, France. 84, avenue de Wagram, F-75017 Paris, France. Department of Biology, University of Washington, Box 351800, Seattle, WA 98195-1800, USA. 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. Historically, mycologists have attempted to tackle the complexity of Cortinarius by organizing species in hierarchical infrageneric taxa deﬁned on supposedly stable sets of characteristics (Kühner & Romagnesi 1953, Moser 1967, Melot 1990, MoënneLoccoz & 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 morphological 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 InTRoduCTIon 1 2 3 4 5 6 7 8 9 © 2017 Naturalis Biodiversity Center & Westerdijk Fungal Biodiversity Institute You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. 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). 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: Here we deal with Cortinarius sect. Bicolores and Cortinarius sect. Saturnini, which encompass Cortinarius evernius, C. saturninus 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 supported 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 1. to circumscribe the phylogenetic boundaries of the two sections, through the analysis of a large internal transcribed spacer (ITS) rDNA sequence dataset built from pan-European and North American vouchered collections; 2. to stabilize the nomenclature and species limits of morphogenetic Bicolores and Saturnini, through sequencing type material and designating neotype or epitype when opportune; 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. Table 1 Cortinarius species classiﬁed in sections Bicolores and Saturnini by the main European authors. This study Bidaud et al. (1992, 2014, 2015) Brandrud et al. (1990, 1994, 1998), Niskanen et al. (2012) Moser (1967) Kühner & Romagnesi (1953) Sect. Bicolores Sect. Bicolores Sect. Bicolores Key 3.11.7.6.11 Sect. Bicolores Cortinarius cagei C. minicolor, C. periodolens ad. int. C. cagei C. bicolor ? C. bicolor ? C. evernius C. evernius, C. scutulatus C. evernius C. dolabratoides sp. nov. C. dolabratus C. imbutoides C. evernius C. evernius, C. parvulior ad. int. 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. plumbosus C. tortuosus, C. plumbosus C. cinnamoviolaceus, C. parevernius C. parevernius C. sp1 C. sp2 C. tortuosus C. tortuosus C. tortuosus C. turgidipes C. cinnamoviolaceus C. cinnamoviolaceus, C. parevernius, C. imbutus C. basicyaneus C. disjungendus C. cyanosterix C. mattiae C. mattiae C. parevernioides C. parevernioides C. salicinus C. salicinus, C. deceptivoides C. quadricolor Sect. Saturnini Sect. Saturnini C. confirmatus C. confirmatus C. mattiae C. subviolascens Sect. Firmiores + sect. Telamonia Key 3.11.7.6.11 Sect. Bicolores C. vilior C. imbutus C. cyprinus C. cyprinus C. imbutus 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. furiosus C. furiosus C. nefastus C. nefastus C. serratissimus* C. saturninoides C. sciophylloides C. sciophylloides C. subbulliardioides* C. illepidus C. subfirmus C. subfirmus C. suboxytoneus C. suboxytoneus, C. fuscocinctus C. cypriacus C. cypriacus C. serratissimus C. saturninus 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) phylogenetically unrelated to the two clades. (Sat.), Saturnini. Asterisk indicates unpublished data of nomenclatural significance. 177 K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini 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 European 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 spacers 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 mycologists, biased towards French authors. In an effort to stabilize nomenclature, 63 sequences were obtained from type collections, 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 contribute 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 species 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, respectively) 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 alignment 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 convergence 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 phylogram was computed from the remaining trees with Bayesian 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 material 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). Sequencing 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 species 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. evernius, 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(text continues on p. 185) 178 Table 2 Specimens included in phylogenetic analyses. Species C. = Cortinarius /Bicolores C. cagei Voucher/SH Voucher/SH annotation CFP 1260 cagei (neotype) C. evernius AB 04-09-186 RH 80814 AB 13-10-120 AB 04-09-169 AB 01-09-41 AB 98-09-94 AB 89-11-309 H:6033519 IK 02-033 IK 95-1576 IK 95-347 KS CO1576 KS CO1290 TN 12-200 TN 11-246 TN 09-196 TN 09-139 TN 03-1713 TN 02-1095 TN 02-959 XC 2013-103 SH188528.07FU (10 sequences) CFP 792 C. dolabratoides sp. nov. AB 00-09-83 PML 1727 AB 96-09-47 AB 91-08-42 PML 622 AB 09-07-44 PML 3469 T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos minicolor A. Bidaud minicolor A. Bidaud & R. Fillion minicolor R. Fillion periodolens ad int. A. Ferville basicyaneus A. Ferville basicyaneus R. Fillion cagei na sp. (holotype) I. Kytövuori marcellae cf. A. Bidaud & R. Fillion sp. I. Kytövuori sp. I. Kytövuori sp. I. Kytövuori ‘smell-of-viola’ I. Kytövuori ‘smell-of-viola’ I. Kytövuori dolabratus (epitype) T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos imbutoides (holotype) A. Bidaud phaeoruber (holotype) G. Chevassut saturninus cf. A. Bidaud armillariellus cf. A. Bidaud privignus sensu Quélet cf. A. Bidaud saturninus cf. A. Faurite orastriatus A. Bidaud dolabratus I. Kytövuori dolabratus I. Kytövuori dolabratus I. Kytövuori dolabratus I. Kytövuori imbutoides K. Soop imbutoides K. Soop dolabratus T. Niskanen dolabratus T. Niskanen dolabratus T. Niskanen dolabratus T. Niskanen dolabratus T. Niskanen dolabratus T. Niskanen dolabratus T. Niskanen privignus sensu Quélet P. Reumaux dolabratus na evernius (neotype) T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos evernius f. pseudoscutulatus (holotype) A. Bidaud evernius f. fragrans (holotype) D. Mazuir parvulior ad int. M. Martin evernius f. pseudoscutulatus A. Bidaud & C. Blanc evernius f. fragrans P. Moënne-Loccoz evernius var. insignis A. Bidaud & A. Faurite evernius var. evernius A. Bidaud Collection Country date Taxonomy Herbarium Accession* 1994 Sweden CFP: D48 (1998) S KX964295 2004 1992 1987 1993 1993 1988 na 2008 2007 2004 2008 2008 2004 2001 1990 France France France France France France Germany/Italy Finland France Finland Finland Finland Finland Finland Sweden AC 22: f1419 (2014) AC 22: f1419 (2014) AC 22: f1419 (2014) AC 22: f1417 (2014) this study this study na this study this study this study this study this study this study this study CFP: D52 (1998) ADC private ADC private ADC private ADC private ADC private ADC private na H ADC private H H H H H S KX964296 KX964297 KX964298 KX964299 KX964300 KX964301 AY669676 KX964302 KX964303 KX964304 KX964305 KX964306 KX964307 KX964308 KX964309 2004 1980 2013 2004 2001 1998 1989 2001 2002 1995 1995 2005 2001 2012 2011 2009 2009 2003 2002 2002 1998 na 1988 France France France France France Canada France Finland Finland Finland Finland Sweden Sweden USA USA USA USA Slovakia Finland Finland France NA/FS/Slovakia Sweden AC 22: f1409 (2014) DM 12(47): 52 (1982) this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study this study na CFP: A11 (1990) PC PC ADC private ADC private ADC private ADC private ADC private H H H H K. Soop private K. Soop private H H H H H H H ADC private na S KX964310 KX964311 KX964312 KX964313 KX964314 KX964315 KX964316 KX964317 KX964318 KX964319 KX964320 KX964321 KX964322 KX964323 KX964324 KX964325 KX964326 KX964327 KX964328 KX964329 KX964330 UDB018659 KX964331 2000 1990 1996 1991 1987 2009 1993 France France France France France France France AC 22: f1407 (2014) AC 22: f1406 (2014) AC 22: f1418 (2014) AC 22: f1407 (2014) AC 22: f1406 (2014) AC 22: f1405 (2014) AC 22: f1404 (2014) PC PC ADC private ADC private ADC private ADC private ADC private KX964332 KX964333 KX964334 KX964335 KX964336 KX964337 KX964338 Persoonia – Volume 39, 2017 C. dolabratus AB 04-09-266 AB 92-10-256 PML 738 XC 2014-02 PML 3588 PML 1057 SH188634.07FU (2 sequences) H:6033567 AB 07-08-48 H:6033615 H:6033575 H:6033570 IK 04-051 IK 01-062 CFP 990 Leg. C. hircinosmus C. plumulosus C. refectus P. Moënne-Loccoz A. Bidaud I. Kytövuori I. Kytövuori P. Moënne-Loccoz P. Moënne-Loccoz T. Niskanen T. Niskanen T. Niskanen T. Niskanen T. Niskanen T. Niskanen T. Niskanen na G. Chevassut A. Bidaud & A. Faurite A. Bidaud A. & R. Bardet G. Redeuilh 1982 2004 2000 1997 1986 1984 2010 2010 2010 2007 2007 2007 2005 na 1978 2003 1992 1996 1987 France France Finland Finland France France Canada Canada Canada Canada Canada Canada Norway Canada / FS/Germany France France France France France AB 08-11-445 AB 92-10-332 AB 91-11-360 PML 2390 PML 1067 AB 14-11-138 AB 03-10-56 AB 99-11-345 TN 12-221 XC 2011-212 XC 2009-64 SH094444.07FU (3 sequences) SH094485.07FU (2 sequences) PML 334 AB 02-09-32 F44390 H:6033565 AB 97-10-341 AB 04-10-357 RH 3417 AB 10-09-183 AB 98-09-119 PML 657 PML 3308 IK 98-1612 TN 04-730 AB 96-09-73 AB 05-09-138 AB 04-10-321 AB 99-09-121 PML 2159 PML 769 PML 17 AB 92-10-293 AB 94-10-268 IK 96-1031 tubulosus turibulosus turibulosus turibulosus turibulosus minicolor cf. sciophyllus cf. livor cf. sp. laetior cf. paranomalus cf. turibulosus sp. hircinosmus (holotype) livor sp. hircinosmus scriptor imbutus cf. plumulosus (holotype) fundatus fundatus fundatus perscutulatus sp. sp. refectus (epitype) refectus refectus refectus refectus refectus refectus testaceoviolaceus scriptor refectus M. Martin R. Fillion A. Bidaud P. Moënne-Loccoz P. Moënne-Loccoz P.-Y. Courio A. Bidaud M. Martin T. Niskanen X. Carteret X. Carteret na na P. Moënne-Loccoz A. Bidaud K. Soop I. Kytövuori G. Chamonaz A. Bidaud R. Henry A. Bidaud & R. Fillion E. & A. Bidaud, A. Faurite P. Moënne-Loccoz A. Bidaud I. Kytövuori T. Niskanen A. Bidaud A. Bidaud A. Bidaud A. Bidaud A. Bidaud P. Moënne-Loccoz P. Moënne-Loccoz A. Bidaud A. Bidaud I. Kytövuori 2008 1992 1991 1991 1988 2014 2003 1999 2012 2011 2009 na na 1986 2002 na 2009 1997 2004 1972 2010 1998 1987 1992 1998 2004 1996 2005 2004 1999 1990 1987 1985 1992 1994 1996 France France France France France France France France USA France France NA/France Poland France France Sweden Finland France France France France Canada France France Finland Finland Germany France France France France France France France France Germany AC 22: f1404 (2014) this study this study this study this study this study this study this study this study this study this study this study this study na DM 12(47): 78 (1982) AC 22: f1414 (2014) AC 19: f1144 (2010) AC 19: f1112 (2010) K&R: 305 (1953, invalid), AC 4: f163 (1992) AC 22: f1414 (2014) AC 19: f1108 (2010) AC 19: f1108 (2010) AC 19: f1108 (2010) AC 19: f1108 (2010) this study this study this study this study this study this study na na AC 12: f575 (2002) AC 23: f1459 (2015) FN: 849 (2012) FN: 849 (2012) AC 19: f1109 (2010) this study SMF 93(3): 359 (1977) AC 22: f1411 (2014) AC 22: f1411 (2014) AC 22: f1411 (2014) this study this study this study AC 22: f1410 (2014) AC 22: f1410 (2014) AC 22: f1410 (2014) AC 22: f1410 (2014) AC 22: f1410 (2014) AC 22: f1410 (2014) AC 22: f1410 (2014) AC 22: f1402 (2014) AC 19: f1109 (2010) this study ADC private ADC private H H ADC private ADC private H H H H H H H na PC PC PC PC GK KX964339 KX964340 KX964341 KX964342 KX964343 KX964344 KX964345 KX964346 KX964347 KX964348 KX964349 KX964350 KX964351 AY669686 KX964352 KX964353 KX964354 KX964355 KX964356 ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private H ADC private ADC private na na PC ADC private S H ADC private ADC private PC ADC private ADC private ADC private ADC private H H PC ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private H KX964357 KX964358 KX964359 KX964360 KX964361 KX964362 KX964363 KX964364 KX964365 KX964366 KX964367 GQ159774 HQ115588 KX964368 KX964369 KX964370 KX964371 KX964372 KX964373 KX964374 KX964375 KX964376 KX964377 KX964378 KX964379 KX964380 KX964385 KX964382 KX964383 KX964384 KX964386 KX964387 KX964388 KX964389 KX964390 KX964381 179 evernius var. evernius evernius evernius evernius evernius evernius evernius evernius evernius evernius evernius evernius evernius evernius glaphurus (holotype) tubulosus (holotype) cedriosmus (holotype) violaeolens (holotype) paranomalus (holotype) K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini C. glaphurus PML 212 AB 04-09-212 IK 00-038 IK 97-123 PML 376 PML 230 TN 10-074 TN 10-055 TN 10-054 TN 07-328 TN 07-312 TN 07-223 TN 05-238 SH188514.07FU (11 sequences) RH 71421 AB 03-11-87 AB 92-10-350 XC 2009-41 GK1142 180 Table 2 (cont.) Voucher/SH Voucher/SH annotation Leg. Collection Country date Taxonomy Herbarium Accession* C. tortuosus IB 79/533 tortuosus (neotype) D. Lamoure 1979 Sweden IB KX964391 XC 2008-43 PAK 354 AB 01-09-19 AB 96-08-19 AB 95-09-34 PML 3551 PML 1225 PML 1214 PML 386 CFP 493 flabelloides (holotype) laetior (holotype) tortuosus tortuosus tortuosus tortuosus tortuosus tortuosus tortuosus tortuosus 2008 1879 2001 1996 1995 1993 1989 1989 1986 1986 France Finland France France France France France France France Norway PC H ADC private ADC private ADC private ADC private ADC private ADC private ADC private S KX964392 KX964393 KX964394 KX964395 KX964396 KX964397 KX964398 KX964399 KX964400 KX964401 AB 02-09-41 AB 96-10-124 IK 99-709 TN 10-087 TN 09-046 TN 07-307 TN 05-006 SH094369.07FU (7 sequences) AB 93-10-425 TN 12-217 UBCOGTR194 TN 05-033 saturninus cf. saturninus cf. tortuosus tortuosus tortuosus tortuosus tortuosus tortuosus turgidipes (holotype) sp. sp. (ectomycorrhiza) sp. M. Pèlerin P.A. Karsten A. Bidaud A. Bidaud C. Blanc A. Bidaud & R. Fillion P. Moënne-Loccoz P. Moënne-Loccoz P. Moënne-Loccoz T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos A. Bidaud C. Blanc I. Kytövuori T. Niskanen T. Niskanen T. Niskanen T. Niskanen na A. & E. Bidaud T. Niskanen na T. Niskanen Opera Botanica 100: 182 (1989) AC 19: f1136 (2010) BFNF 32: 387 (1879) AC 22: f1413 (2014) AC 22: f1413 (2014) AC 22: f1413 (2014) AC 22: f1413 (2014) AC 22: f1413 (2014) AC 22: f1413 (2014) AC 22: f1413 (2014) CFP: A06 (1990) 2002 1996 1999 2010 2009 2007 2005 na 1993 2012 na 2005 France France Finland Canada USA Canada Finland USA /U/Japan France USA Canada Finland this study this study this study this study this study this study this study na AC 17(1): f885 (2008) na na na ADC private ADC private H H H H H na PC H na H KX964402 KX964403 KX964404 KX964405 KX964406 KX964407 KX964408 AY669669 KX964409 KX964410 EU597034 KX964411 cinnamoviolaceus (holotype) M. Moser 1948 Austria IB KX964412 RH 70942 RH 4000 RH 526 RH 1240 RH 3258a78 AB 02-10-71 CFP 574 basicyaneus (holotype) cylindratus (holotype) subparevernius (holotype) contractus (holotype) parevernius (holotype) dolabratus imbutus 1976 1972 1956 1960 1955 2002 1987 France France France France France France Sweden PC PC PC PC PC ADC private S KX964413 KX964414 KX964415 KX964416 KX964417 KX964418 KX964419 AB 12-11-240 TN 05-198 TN 05-051 SH188640.07FU (2 sequences) RH 338 imbutus imbutus sensu Funga Nordica imbutus sensu Funga Nordica imbutus cyanosterix (holotype) M. Trimbach R. Henry R. Henry R. Henry R. Henry A. & M. Burat T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos A. Bidaud T. Niskanen T. Niskanen na R. Henry Nova Hedwigia XIV(2-4): 514 (1967) FAMM 25: 38 (2004) SMF 99(1): 91 (1983) SMF 85(4): 442 (1969) SMF 85(4): 387 (1969) K&R: 303 (1953, invalid) AC 17(1): f817 (2008) CFP: D60 (1998) 2012 2005 2005 na 1952 France Finland Finland Sweden /Italy France this study this study this study na SMF 71(3): 259, 261 (1956) ADC private H H na PC KX964420 KX964421 KX964422 UDB001160 KX964423 KS CO1936 AB 13-08-35 AB 99-09-77 PML 650 CFP 1204 mattiae (isotype) mattiae subviolascens subviolascens mattiae 2009 2013 1999 1987 1993 Sweden France France France Sweden JEC 13(12): 3 (2010) AC 22: f1415 (2014) AC 12: f565 (2002) AC 12: f565 (2002) CFP: D30 (1998) S ADC private ADC private ADC private S KX964424 KX964425 KX964426 KX964427 KX964428 AB 06-09-153 H:6029375 licinipes /poecilopus aff. mattiae 2006 2004 France Finland this study this study ADC private H KX964429 KX964430 C. turgidipes C. sp1 C. sp2 other (morphological) Bicolores C. cinnamoviolaceus IB 48/590 C. cyanosterix (= C. disjungendus) C. mattiae K. Soop A. Bidaud, F. Armada & R. Fillion A. Bidaud P. Moënne-Loccoz T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos A. Bidaud, F. Armada & R. Fillion T. Niskanen Persoonia – Volume 39, 2017 Species C. = Cortinarius C. salicinus /saturnini C. confirmatus C. cyprinus C. imbutus I. Kytövuori I. Kytövuori I. Kytövuori A. Bidaud A. Bidaud na C. Gérard na C. Hugouvieux 2007 2001 1998 1993 1991 na 2002 na 2005 Finland Sweden Finland France France Canada France NA/U France this study this study this study this study this study na AC 22: f1408 (2014) na AC 22: f1416 (2014) H H H ADC private ADC private na PC na PC KX964431 KX964432 KX964433 KX964434 KX964435 FJ039684 KX964436 KF617653 KX964437 RH 3195 JVG 990125-31 MES 3541 RH 84/159 XC 2013-160 AB 13-10-97 XC 2011-199 XC 95-10-04-06 AB 09-11-452 AB 00-10-193 AB 11-11-324 PML 4722 XC 2012-171 AB 09-11-514 AB 05-11-423 AB 02-11-201 XC 2013-156 AB 03-11-78 AB 92-11-422 AB 09-11-450 FR2016052 FR2012405 FR2012089 FR2012076 XC 2006-204 XC 2005-249 SH094374.07FU (6 sequences) XC 2012-26 AB 11-11-251 AB 11-10-192 AB 06-09-144 PML 344 PML 81 XC 2013-15 XC 2007-103 AB 04-09-167 JMB 2014111802 PAM 13092901 PML 425 XC 2007-95 TEB 348-10 TAAM 128765 IK 97-1162 PML 4557 RH 3123 confirmatus (holotype) assiduus var. plesiocistus (isotype) assiduus (holotype) bulbosovolvatus (isotype) confirmatus ‘asp. subcylindratus’ confirmatus ‘asp. kuehneri’ confirmatus ‘asp. spurcatocephalus’ confirmatus ‘asp. spurcatocephalus’ confirmatus ‘asp. rubricosissimus’ confirmatus ‘asp. rubricosissimus’ confirmatus ‘asp. paracohabitans’ confirmatus ‘asp. imbutus’ confirmatus ‘asp. imbutus’ confirmatus ‘asp. assiduus’ confirmatus ‘asp. assiduus’ confirmatus ‘asp. assiduus’ confirmatus ‘asp. assiduus’ confirmatus ‘asp. confirmatus’ cistoadelphus ad int. cohabitans cf. assiduus assiduus assiduus assiduus bresadolae cf. saturninus cf. sp. cyprinus (holotype) cyprinus cyprinus cyprinus cyprinus cyprinus cyprinus cyprinus sciophyllus cf. circumvelatus cf. circumvelatus myrtillinus mutabilis cf. saturninus aff. sp. imbutus (neotype) laccatus (holotype) betulaecomes (holotype) R. Henry X. Llimona & J. Vila R. Mahiques M. Contu & L. Curreli na A. Bidaud X. Carteret X. Carteret A. Bidaud A. Bidaud F. Armada, A. Bidaud & J. Pardo P. Reumaux A. Lantz A. Bidaud A. & E. Bidaud F. Lopez F. Valade A. Faurite A. Bidaud A. Bidaud J.-M. Ourcival P.-A. Moreau F. Richard E. Taschen na X. Carteret na G. Redeuilh A. Bidaud A. Bidaud A. Bidaud P. Moënne-Loccoz P. Moënne-Loccoz P. Reumaux X. Carteret A. Bidaud P.-A. Moreau P.-A. Moreau P. Moënne-Loccoz na T.E. Brandrud A. Kollom I. Kytövuori P. Reumaux R. Henry 1970 1999 1999 1984 2013 2013 2011 1995 2009 2000 2011 1990 2012 2009 2005 2002 2013 2003 1992 2009 2016 2012 2011 2011 2006 2005 na 1993 2011 2011 2006 1986 1981 2013 2007 2004 2014 2013 1986 2007 na 2008 1997 1978 1976 France Spain Spain Italy France France France France France France France France France France France France France France France France France France France France France France U/Iran France France France France France France France France France France France France France Norway Estonia Finland France France SMF 99(1): 67 (1983) Mycotaxon 101: 140 (2007) FMDS 162: 42 (2001) DM 26 (61): 32 (1985) AC 23: f1441 (2015) AC 23: f1440 (2015) AC 23: f1439 (2015) AC 23: f1439 (2015) AC 23: f1438 (2015) AC 23: f1438 (2015) AC 23: f1437 (2015) AC 23: f1436 (2015) AC 23: f1436 (2015) AC 23: f1435 (2015) AC 23: f1435 (2015) AC 23: f1435 (2015) AC 23: f1435 (2015) AC 23: f1434 (2015) FAMM 6: 41 (1994) this study this study this study this study this study this study this study na AC 23: f1443 (2015) AC 23: f1443 (2015) AC 23: f1443 (2015) AC 23: f1443 (2015) AC 23: f1443 (2015) AC 23: f1443 (2015) AC 23: f1443 (2015) AC 23: f1443 (2015) this study this study this study this study this study this study na this study SMF 98(4): 348 (1982) SMF 93(3): 347 (1977) PC J. Vila private MES PC PC ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private CEFE private CEFE private CEFE private CEFE private ADC private ADC private na PC ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private CEFE private CEFE private ADC private ADC private na na H PC PC KX964438 AM713178 KX964439 KX964440 KX964441 KX964442 KX964443 KX964444 KX964445 KX964446 KX964447 KX964448 KX964449 KX964450 KX964451 KX964452 KX964453 KX964454 KX964455 KX964456 KX964457 KX964458 KX964459 KX964460 KX964461 KX964462 HQ204652 KX964463 KX964464 KX964465 KX964466 KX964467 KX964468 KX964469 KX964470 KX964471 KX964472 KX964473 KX964474 KX964475 KX964476 UDB016164 KX964498 KX964478 KX964479 181 mattiae mattiae mattiae umbrinoconnatus forma oxytoneus sp. parevernioides (holotype) malachius salicinus (holotype) K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini C. parevernioides H:6000560 IK 01-039 IK 98-1127 PML 3989 PML 2298 SH009438.07FU (1 sequence) AB 02-09-50 SH188502.07FU (15 sequences) XC 2014-03 Species C. = Cortinarius C. lucorum Voucher/SH Voucher/SH annotation Leg. Collection Country date XC 2013-13 XC 2014-77 XC 2014-61 XC 2007-104 AB 10-10-237 AB 09-11-471 AB 04-09-228 AB 98-10-358 PML 375 XC 2002-122 XC 2002-108 XC 2002-107 XC 2002-106 AB 08-10-307 AB 02-10-106 AB 02-09-58 AB 00-09-127 IK 98-2242 IK 94-1236 JMB 2008092703 RH 71030 TN 11-257 TN 11-252 TN 11-151 TN 11-150 TN 05-167 XC 2012-96 XC 2002-109 SH188563.07FU (6 sequences) CFP 490 imbutus ‘asp. laetior’ imbutus ‘asp. saturnalis’ imbutus ‘asp. saturnalis’ imbutus ‘asp. vilior’ imbutus ‘asp. imbutus’ imbutus ‘asp. imbutus’ imbutus ‘asp. imbutus’ imbutus ‘asp. imbutus’ imbutus ‘asp. imbutus’ imbutus ‘asp. imbutus’ imbutus ‘asp. imbutus’ imbutus ‘asp. imbutus’ imbutus ‘asp. imbutus’ cohabitans cohabitans cohabitans cohabitans cf. sp. sp. salicis cf. betulaecomes sp. sp. sp. sp. sp. laetior forma renidentoides cf. saturninus lucorum (neotype) 1998 1978 1986 2007 2010 2009 2004 1998 1986 2002 2002 2002 2002 2008 2002 2002 2000 1998 1994 2008 1976 2011 2011 2011 2011 2005 2012 2002 na 1986 RH 71502 incarnatolilascens (holotype) P. Reumaux P. Reumaux P. Reumaux X. Carteret A. Bidaud A. Bidaud & R. Fillion A. Bidaud & A. Faurite A. Bidaud P. Reumaux X. Carteret X. Carteret X. Carteret X. Carteret J. Garin M. Renard A. Bidaud A. Bidaud I. Kytövuori I. Kytövuori J.-M. Bellanger R. Henry T. Niskanen T. Niskanen T. Niskanen T. Niskanen T. Niskanen X. Carteret X. Carteret na T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos R. Henry PML 4142 montis-dei (holotype) PML 34 Taxonomy Herbarium Accession* France AC 23: f1447 (2015) France AC 23: f1446 (2015) France AC 23: f1446 (2015) France AC 23: f1445 (2015) France AC 23: f1444 (2015) France AC 23: f1444 (2015) France AC 23: f1444 (2015) France AC 23: f1444 (2015) France AC 23: f1444 (2015) France AC 23: f1444 (2015) France AC 23: f1444 (2015) France AC 23: f1444 (2015) France AC 23: f1444 (2015) France this study France this study France this study France this study Sweden this study Finland this study France this study France this study (Rob. Henry, ined.) USA this study USA this study USA this study USA this study Finland this study France this study France this study Canada /Estonia / China na Norway CFP: C10 (1994) ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private H H CEFE private PC H H H H H ADC private ADC private na S KX964480 KX964481 KX964482 KX964483 KX964484 KX964485 KX964486 KX964487 KX964488 KX964489 KX964490 KX964491 KX964492 KX964493 KX964494 KX964495 KX964496 KX964497 KX964477 KX964499 KX964500 KX964501 KX964502 KX964503 KX964504 KX964505 KX964506 KX964507 UDB018346 KX964585 1979 France PC KX964508 P. Reumaux 1980 France PC KX964509 circumvelatus (holotype) P. Reumaux 1976 France PC KX964510 10433 umidicola (syntype) C.H. Kauffman 1903 USA MICH KX964511 PML 4143 PAM 14090808 IK 89-748 KS CO513 TN 10-002 TN 03-1169 SH188495.07FU (21 sequences) CFP 514 lucorum ‘asp. montis-dei’ lucorum ‘asp. circumvelatus’ lucorum diabolicus lucorum lucorum lucorum saturninus (neotype) 1980 2014 1989 na 2010 2003 na 1986 France France Finland Sweden Canada Sweden NA / FS Sweden ADC private ADC private H na H H na S KX964512 KX964513 KX964514 KX964515 KX964516 KX964517 UDB019872 KX964584 PML 4578 urbicus var. sporanotandus (holotype) P. Reumaux P.-A. Moreau I. Kytövuori K. Soop T. Niskanen T. Niskanen na T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos A. Bidaud 1996 France PC KX964518 AC 23: f1431 (2015), SMF 97(3): 170 (1981) AC 23: f1430 (2015), SMF 96(3): 357 (1980) AC 23: f1429 (2015), SMF 96(3): 355 (1980) Bull. Torrey Bot. Club 32(6): 322 (1905) AC 23: f1430 (2015) AC 23: f1429 (2015) this study this study this study this study na CFP: C09 (1994) AC 23: f1455 (2015), AC 12: f560 (2002) Persoonia – Volume 39, 2017 C. saturninus 182 Table 2 (cont.) marginatosplendens (isotype) P. Reumaux 1978 France XC 2007-14 fulvorimosus (holotype) A. & R. Bardet 1992 France RH 3758 RH 81181 RH 71682 PR 258 RH 2623 RH 476 AB 02-10-179 AB 95-11-144 XC 2001-107 AB 14-11-160 (= AB 14-11-161) denseconnatus (holotype) gramineus (holotype) rastetteri (holotype) dissidens (holotype) salicis (holotype) umbrinoconnatus (holotype) saturninus ‘asp. urbicoides’ saturninus ‘asp. urbicoides’ saturninus ‘asp. urbicoides’ saturninus ‘asp. salicis’ 1973 1981 1980 1978 1968 1955 2002 1995 2001 2014 XC 2014-109 XC 2011-205 XC 2008-55 XC 2007-108 AB 14-09-47 AB 04-10-344 AB 98-10-381 XC 2014-63 XC 2014-116 XC 2014-114 XC 2007-97 AB 97-09-187, PML 5347 PML 3967 CFP 408 saturninus ‘asp. salicis’ saturninus ‘asp. salicis’ saturninus ‘asp. salicis’ saturninus ‘asp. salicis’ saturninus ‘asp. dionisiae’ saturninus ‘asp. deceptivus’ saturninus ‘asp. deceptivus’ saturninus ‘asp. cohabitans’ saturninus ‘asp. saturninus’ saturninus ‘asp. saturninus’ saturninus ‘asp. saturninus’ urbicus salicis var. salicis subtorvus AB 05-10-273 H:6029320 IK 94-631 JMB 2009101002 KH14 O50591 PML 75 TN 09-208 XC 2016-12 XC 2008-61 XC 2007-90 XC 2006-194 XC 2002-167 XC 2001-104 XC 96-10-26-09 SH094324.07FU (13 sequences) Rehner 394 deceptivus sensu Moser saturninus saturninus cohabitans subtorvus subtorvus urbicus saturninus euprivignus aff. salicis mutabilis cf. salicis holophaeus sensu Henry mutabilis subprivignus saturninus stuntzii (holotype) na R. Henry V. Rastetter P. Reumaux R. Henry R. Henry A. Faurite R. Fillion X. Carteret & P. Reumaux A. Bidaud, J. Cavet, R. Fillion & G. Raffini X. Carteret X. Carteret X. Carteret X. Carteret E. Bidaud A. Bidaud Dr. Misermont M. Pèlerin na L. Tarahu na E. & A. Bidaud M. Citérin T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos R. Fillion I. Kytövuori I. Kytövuori J.-M. Bellanger na na P. Moënne-Loccoz T. Niskanen P. Reumaux X. Carteret X. Carteret X. Carteret M. Pèlerin X. Carteret X. Carteret na S.A. Rehner other (morphological) saturnini C. cypriacoides PML 1269 G KX964519 PC KX964520 PC PC PC PC PC PC ADC private ADC private ADC private ADC private KX964521 KX964522 KX964523 KX964524 KX964525 KX964526 KX964527 KX964528 KX964529 KX964530 2014 2011 2008 2007 2014 2004 1998 1996 2014 2014 2007 1997 1994 1986 France France France France France France France France France France France France France Sweden AC 23: f1454 (2015) AC 23: f1454 (2015) AC 23: f1454 (2015) AC 23: f1454 (2015) AC 23: f1451 (2015) AC 23: f1450 (2015) AC 23: f1450 (2015) AC 23: f1449 (2015) AC 23: f1448 (2015) AC 23: f1448 (2015) AC 23: f1448 (2015) AC 12: f560 (2002) AC 12: f559 (2002) CFP: A04 (1990) ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private S KX964531 KX964532 KX964533 KX964534 KX964535 KX964536 KX964537 KX964538 KX964539 KX964540 KX964541 KX964542 KX964543 KX964544 2005 1998 1994 2009 2011 2011 1984 2009 1977 2008 2007 2006 2002 2001 1996 na 1981 France Finland Finland France Norway (Svalbard) Norway (Svalbard) France USA France France France France France France France USA/U USA this study this study this study this study na na this study this study this study this study this study this study this study this study this study na Mycologia 80(6): 903 (1988) ADC private H H CEFE private na na ADC private H ADC private ADC private ADC private ADC private ADC private ADC private ADC private na WTU KX964545 KX964546 KX964547 KX964548 GU234058 GU234013 KX964549 KX964550 KX964551 KX964552 KX964553 KX964554 KX964555 KX964556 KX964557 UDB017613 KX964558 AC 23: f1423 (2015), AC 2: f81 (1990) AC 23: f1423 (2015) AC 23: f1424 (2015), AC 9: f419 (1999) AC 23: f1458 (2015) na PC KX964559 ADC private ADC private KX964560 KX964561 PC na KX964562 KM576363 R. Fillion 1989 France PML 3984 PML 3979 cypriacoides ‘asp. cypriacoides’ (holotype) cypriacoides ‘asp. cypriacoides’ cypriacoides ‘asp. lucorum’ C. Guyot A. Bidaud 1989 1992 France France XC 2014-64c LM5411 furiosus (holotype) sp. (Quercus ectomycorrhiza) D. Brion na 2012 na France Austria 183 C. furiosus France France France France France France France France France France AC 23: f1453 (2015), SMF 96(3): 356 (1980) AC 23: f1452 (2015), AC 17: f869 (2008) SMF 99(1): 65 (1983) SMF 99(1): 64 (1983) SMF 97(3): 177 (1981) SMF 96(3): 370 (1980) SMF 93(3): 364 (1977) SMF 73(1): 53 (1957) AC 23: f1457 (2015) AC 23: f1457 (2015) AC 23: f1457 (2015) AC 23: f1454 (2015) K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini C. stuntzii PML 2215 184 Table 2 (cont.) Voucher/SH Voucher/SH annotation Leg. Collection Country date Taxonomy Herbarium Accession* C. illepidus sensu ADC (= C. subbulliardioides) AB 11-11-331 illepidus A. Bidaud & C. Gérard 2011 France AC 23: f1422 (2015) ADC private KX964563 AB 11-11-330 XC 2014-60 JB 604808 RH 931 AB 12-10-93 AB 00-10-148 RH 3451 XC 2014-119 XC 2014-64b XC 2013-144 XC 2010-56 XC 2010-29 SH188624.07FU (3 sequences) AB 99-10-254 AB 91-10-291 PML 5446 PML 2381 SH188568.07FU (6 sequences) AB 08-10-363 AB 01-09-56 MFT60 illepidus nefastus (holotype) ortovernus (holotype) oxytoneus (holotype) saturninoides saturninoides oxytoneus saturninoides saturninoides saturninoides saturninoides saturninoides lucorum sciophylloides (holotype) sciophylloides sciophylloides raphanodiabolicus valgus subfirmus (holotype) suboxytoneus (holotype) sp. (Fagus ectomycorrhiza) A. Bidaud & C. Gérard D. Brion J. Ballará R. Henry A. Bidaud & M. Renard A. Bidaud R. Henry R. Chalange D. Brion F. Valade X. Carteret X. Carteret na A. Bidaud J. Garin J. Cavet P. Reumaux na A. Bidaud & G. Raffini A. Bidaud na 2011 2012 2008 1957 2012 2000 1972 2014 2012 2013 2010 2010 na 1999 1991 1999 1991 na 2008 2001 na France France Spain France France France France France France France France France USA/Estonia /Italy France France France France Canada/U France France Germany AC 23: f1422 (2015) AC 23: f1426 (2015) JEC 12(11): 56 (2009) SMF 97(3): 277 (1981) AC 23: f1421 (2015) AC 23: f1421 (2015) SMF 97(3): 277 (1981) AC 23: f1421 (2015) AC 23: f1421 (2015) AC 23: f1421 (2015) AC 23: f1421 (2015) AC 23: f1421 (2015) na AC 23: f1425 (2015) AC 23: f1425 (2015) AC 23: f1425 (2015) na na AC 23: f1433 (2015) AC 23: f1442 (2015) na ADC private PC J. Ballara private PC ADC private ADC private ADC private ADC private ADC private ADC private ADC private ADC private na PC ADC private ADC private ADC private na PC PC na KX964564 KX964565 KX964566 KX964567 KX964568 KX964569 KX964570 KX964571 KX964572 KX964573 KX964574 KX964575 UDB016052 KX964576 KX964577 KX964578 KX964579 UDB002444 KX964580 KX964581 FJ403502 other telamonia C. alboviolaceus s.lat. C. anisatus SH188487.07FU (26 sequences) CFP 1200 alboviolaceus anisatus (holotype) na 1993 NA/U Sweden na CFP: E25 (2014) na S AF325596 DQ117931 C. anisochrous C. athabascus C. biformis C. bovinus C. brunneifolius IK 01-030 DBB27618, UC1860905 SH188479.07FU (41 sequences) IK 04-038 TN 06-146 anisochrous (holotype) athabascus (holotype) biformis bovinus (neotype) brunneifolius (holotype) na T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos T. Niskanen & I. Kytövuori D. Bojantchev na I. Kytövuori T. Niskanen 2001 2011 na 2004 2006 Estonia USA NA/U Finland Finland H, S, NY UC na H, S, NY H JX407297 JN133295 UDB002252 JX407276 EU259284 C. caesioarmeniacus C. claroplaniusculus C. decipiens H:7000901 RH 2334 PML 366 caesioarmeniacus (holotype) claroplaniusculus (holotype) decipiens f. decipiens (neotype) K. Liimatainen & T. Niskanen R. Henry P. Moënne-Loccoz 2007 1967 1986 Canada France France H PC G KP137498 KP013184 FN428988 C. disjungendus C. duracinus C. duracinus s.lat. C. duracinus s.lat. C. fuscescens C. fuscobovinaster C. gallurae PAK 4370 PML 349 SH188648.07FU (2 sequences) SH094372.07FU (6 sequences) H:6001898 IK 09-537 CONS 00076 disjungendus (lectotype) duracinus (neotype) sp. rigens fuscescens (holotype) fuscobovinaster (holotype) gallurae (holotype) P.A. Karsten P. Moënne-Loccoz na na K. Liimatainen & T. Niskanen I. Kytövuori D. & M. Antonini, G. Consiglio < 1893 1986 na na 2008 2009 2002 Finland France Denmark/Germany NA /Italy Finland Norway Italy H G na na H H, S, NY CONS KP013190 KX964582 AJ889943 JF907880 KP165546 JX407316 FN428979 C. murinascens C. neofurvolaesus IK 08-958 CFP 1438 murinascens (holotype) neofurvolaesus (holotype) I. Kytövuori T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos 2008 1999 Finland Sweden Mycologia 105(4): 988 (2013) Mycotaxon 123: 382 (2013) na Mycologia 105(4): 981 (2013) Mycol. Progress 7(4): 241 (2008) IF 198: 1 (2014) SMF 99(1): 65 (1983) AC 11(1): f507 (2001), AC 2: f52 (1990) ASFFF 9(1): 6 (1893) AC 2: f76 (1990) na na IF 201: 2 (2014) Mycologia 105(4): 990 (2013) Il genereCortinarius in Italia 3: C101 (2005) IF 201: 3 (2014) CFP: E24 (2014) H S KP165570 DQ139999 C. nefastus C. ortovernus C. oxytoneus C. saturninoides sensu ADC (= C. serratissimus) C. sciophylloides C. subfirmus C. suboxytoneus Persoonia – Volume 39, 2017 Species C. = Cortinarius FJ717605 UDB002227 DQ663239 KF732554 KF732420 WTU na G H IB USA U France Finland USA 2007 na 1957 2008 1997 J.F. Ammirati na R. Kühner I. Kytövuori M. Moser sordidemaculatus (holotype) bovinus cf. aprinus subserratissimus (holotype) subturibulosus tacitus (holotype) torvus urbicus anomalovelatus (holotype) anomalus caesiocinctus (holotype) flavipallens (holotype) sannio (holotype) RH 1122 IB 86/172 TF-01-034 IK 11-017 SH188545.07FU (7 sequences) AB 05-09-72 SH009362.07FU (10 sequences) SH188612.07FU (3 sequences) JFA13109 SH196665.07FU (12 sequences) Sa57-13 IK 08-1729, H:6032393 MM 97/352, IB:1997/0352 C. sordidemaculatus C. sp. C. sp. C. subserratissimus C. subturibulosus C. tacitus C. torvus C. urbicus outgroup C. anomalovelatus C. lepidopus sensu auct. C. caesiocinctus C. flavipallens C. sannio 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. DQ139984 DQ139983 AJ889942 KP165552 FJ928484 KX964583 UDB001345 UDB000743 PC IB C H na PC na na France SMF 97(3): 196 (1981) Austria na Denmark na Sweden IF 201: 4 (2014) France /Spain /Portugal na France AC 22: f1400 (2014) NA /FS/Germany na Canada/FS na < 1981 1986 na 2011 na 2005 na na na na R. Henry T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos R. Henry M. Moser T. Frøslev I. Kytövuori na A. Bidaud na na niveotraganus olididisjungendus (holotype) orasericeus (holotype) quarciticus (holotype) SH188538.07FU (8 sequences) TN 07-191, H:7000854 RH 70239 CFP 765 C. niveotraganus C. olididisjungendus C. orasericeus C. quarciticus na IF 186: 2 (2014) SMF 99(1): 69 (1983) CFP: C59 (1994) FS Canada France Sweden na 2007 1975 1988 IF 93: 1 (2014) na DM 20(77): 92 (1989) Persoonia 33 : 125 (2014) Mycotaxon 72 : 315 (1999) KM273103 KM273091 KP013203 UDB000748 na H PC S K. Liimatainen et al.: Cortinarius section Bicolores and section Saturnini 185 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. hircinosmus, 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 comparatively 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 resulting 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 substitutions 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 allows 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. subparevernius, 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. Duracini and C. sciophylloides in sect. Brunneotincti (Fig. 1, Table 1). (text continues on p. 190) 186 Persoonia – Volume 39, 2017 100/0.88 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 /Saturnini (Fig. 3) /Bicolores (Fig. 2) 99/0.92 Cortinarius tacitus AB05-09-72 HOLOTYPE France C. subbulliardioides Cortinarius illepidus AB11-11-331 France (Sat) Cortinarius illepidus AB11-11-330 France Cortinarius subfirmus AB08-10-363 HOLOTYPE France (Sat) Cortinarius neofurvolaesus CFP1438 HOLOTYPE Sweden Cortinarius anisatus CFP1200 HOLOTYPE Sweden Cortinarius sordidemaculatus SH188519.07FU NA-FS-France (9 seq) Cortinarius fuscobovinaster IK09-537 HOLOTYPE Norway Cortinarius bovinus cf. IB86/172 Austria Cortinarius aprinus TF01-034 Denmark Cortinarius bovinus IK04-038 NEOTYPE Finland Cortinarius anisochrous IK01-030 HOLOTYPE Estonia Cortinarius cypriacoides asp. cypriacoides PML3984 France 60/Cortinarius cypriacoides asp. lucorum PML3979 France (Sat) Cortinarius cypriacoides asp. cypriacoides PML1269 HOLOTYPE France 84/0.7 61/97/0.42 63/99/0.75 100/0.77 C. cinnamoviolaceus (Bic) Cortinarius cinnamoviolaceus AB12-11-240 France Cortinarius imbutus CFP574 Sweden Cortinarius parevernius RH3258a78 HOLOTYPE France Cortinarius cinnamoviolaceus IB48/590 HOLOTYPE Austria (short) 67/Cortinarius subparevernius RH526 HOLOTYPE France (short) Cortinarius cylindratus RH4000 HOLOTYPE France Cortinarius dolabratus AB02-10-71 France Cortinarius basicyaneus RH70942 HOLOTYPE France Cortinarius contractus RH1240 HOLOTYPE France Cortinarius cinnamoviolaceus TN05-198 Finland Cortinarius cinnamoviolaceus TN05-051 Finland Cortinarius imbutus SH188640.07FU Sweden-Italy (2 seq) Cortinarius mattiae AB06-09-153 France Cortinarius subviolascens PML650 France Cortinarius subviolascens AB99-09-77 France Cortinarius mattiae PML3989 France Cortinarius mattiae PML2298 France Cortinarius mattiae IK98-001 Finland Cortinarius mattiae AB13-08-35 France Cortinarius mattiae H6000560 Finland Cortinarius mattiae CFP1204 Sweden Cortinarius mattiae IK01-039 Sweden 54/Cortinarius sp. SH009438.07FU Canada (1 seq) Cortinarius mattiae H6029375 Finland Cortinarius mattiae KS-CO1936 ISOTYPE Sweden 56/Cortinarius athabascus DBB27618 HOLOTYPE USA Cortinarius lucorum SH188624.07FU USA-Estonia-Italy (3 seq) Cortinarius saturninoides XC2010-56 France Cortinarius saturninoides XC2013-144 France Cortinarius saturninoides XC2014-119 France Cortinarius oxytoneus RH3451 France Cortinarius saturninoides AB00-10-148 France Cortinarius saturninoides AB12-10-93 France Cortinarius saturninoides XC2014-64b France Cortinarius saturninoides XC2010-29 France Cortinarius subserratissimus IK11-017 HOLOTYPE Sweden Cortinarius disjungendus PAK4370 LECTOTYPE Finland (Bic) 75/0.65 Cortinarius cyanosterix RH338 HOLOTYPE France Cortinarius orasericeus RH70239 HOLOTYPE France Cortinarius claroplaniusculus RH2334 HOLOTYPE France Cortinarius olididisjungendus H7000854 HOLOTYPE Canada 94/Cortinarius oxytoneus RH931 HOLOTYPE France (Sat) Cortinarius rigens SH094372.07FU NA-Italy (6 seq) Cortinarius sp. SH188648.07FU Denmark-Germany (2 seq) Cortinarius duracinus PML349 NEOTYPE France Cortinarius parevernioides AB02-09-50 HOLOTYPE France (Bic) Cortinarius malachius SH188502.07FU NA-U (15 seq) 67/ 0.65 Cortinarius suboxytoneus AB01-09-56 HOLOTYPE France (Sat) Cortinarius sp. (ecto. Fagus) MFT60 Germany Cortinarius alboviolaceus SH188487.07FU NA-U (26 seq) 51/Cortinarius quarciticus CFP765 HOLOTYPE Sweden Cortinarius biformis SH188479.07FU NA-U (41 seq) Cortinarius caesioarmeniacus H7000901 HOLOTYPE Canada Cortinarius murinascens IK08-958 HOLOTYPE Finland Cortinarius urbicus SH188612.07FU Canada-FS (3 seq) 71/55/Cortinarius niveotraganus SH188538.07FU FS (8 seq) Cortinarius torvus SH009362.07FU NA-FS-Germany (10 seq) Cortinarius sciophylloides PML5446 France Cortinarius sciophylloides PML2381 France Cortinarius valgus SH188568.07FU Canada-U (6 seq) (Sat) Cortinarius sciophylloides AB99-10-254 HOLOTYPE France 91/Cortinarius sciophylloides AB91-10-291 France (Sat) Cortinarius ortovernus JB604808 HOLOTYPE Spain Cortinarius subturibulosus SH188545.07FU France-Spain-Portugal (7 seq) (Bic) Cortinarius salicinus XC2014-03 HOLOTYPE France Cortinarius gallurae CONS00076 HOLOTYPE Italy 99/Cortinarius decipiens PML366 NEOTYPE France Cortinarius nefastus XC2014-60 HOLOTYPE France (Sat) Cortinarius brunneifolius TN-06146 HOLOTYPE Finland Cortinarius sp. (ecto. Quercus) LM5411 Austria (Sat) 65/ Cortinarius furiosus XC201464c HOLOTYPE France 0.80 Cortinarius fuscescens H6001898 HOLOTYPE Finland Cortinarius anomalovelatus JFA13109 HOLOTYPE USA Cortinarius lepidopus s. auct. SH196665.07FU U (12 seq) Cortinarius sannio IB97/352 HOLOTYPE USA Outgroup Cortinarius flavipallens IK08-1729 HOLOTYPE Finland Cortinarius caesiocinctus Sa5713 HOLOTYPE France 73/- Sect. Bovini C. mattiae (Bic) C. serratissimus (Sat) 98/0.66 Sect. Sciophylli Sect. Disjungendi Sect. Duracini Sect. Firmiores Sect. Urbici Sect. Telamonia Sect. Brunneotincti Sect. Hydrocybe 0.02 substitution per site Fig. 1 Sections Bicolores and Saturnini within subg. Telamonia. — Bayesian 50 % majority-rule consensus tree inferred from the analysis of 348 ITS sequences (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 85/- 67/- Cortinarius tortuosus PML3551 France Cortinarius tortuosus IB79/533 NEOTYPE Sweden (short) Cortinarius tortuosus SH094369.07FU USA-U-Japan (7 seq) Cortinarius tortuosus AB01-09-19 France Cortinarius tortuosus AB96-10-124 France Cortinarius tortuosus PML1214 France Cortinarius tortuosus TN09-046 USA Cortinarius tortuosus CFP493 Norway Cortinarius tortuosus IK99-709 Finland Cortinarius tortuosus TN07-307 Canada Cortinarius tortuosus TN10-087 Canada Cortinarius tortuosus AB02-09-41 France Cortinarius tortuosus PML1225 France Cortinarius tortuosus TN05-006 Finland Cortinarius laetior PAK354 HOLOTYPE Finland (short) Cortinarius tortuosus AB96-08-19 France Cortinarius flabelloides XC2008-43 HOLOTYPE France Cortinarius tortuosus PML386 France Cortinarius tortuosus AB95-09-34 France Cortinarius turibulosus SH094444.07FU NA-France (3 seq) 95/Cortinarius glaphurus TN12-221 USA Cortinarius glaphurus RH71421 HOLOTYPE France Cortinarius glaphurus AB03-10-56 France Cortinarius turibulosus PML1067 France (short) Cortinarius glaphurus XC2011-212 France Cortinarius tubulosus AB03-11-87 HOLOTYPE France Cortinarius tubulosus AB08-11-445 France Cortinarius cedriosmus AB92-10-350 HOLOTYPE France Cortinarius turibulosus AB92-10-332 France (short) Cortinarius sp. SH094485.07FU Poland (2 seq) Cortinarius glaphurus AB99-11-345 France Cortinarius glaphurus AB14-11-138 France Cortinarius turibulosus AB91-11-360 France (short) Cortinarius turibulosus PML2390 France Cortinarius violaeolens XC2009-41 HOLOTYPE France Cortinarius glaphurus XC2009-64 France Cortinarius paranomalus GK1142 HOLOTYPE France Cortinarius tortuosus Cortinarius glaphurus Cortinarius hircinosmus AB04-10-357 France Cortinarius scriptor AB97-10-341 France Cortinarius livor AB02-09-32 France Cortinarius hircinosmus H6033565 Finland Cortinarius hircinosmus F44390 Sweden Cortinarius hircinosmus PML334 HOLOTYPE France 65/0 80/- 82/- 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 Cortinarius dolabratus AB89-11-309 France Cortinarius dolabratus AB13-10-120 France Cortinarius dolabratus AB01-09-41 France Cortinarius dolabratus AB04-09-169 France Cortinarius dolabratus CFP990 EPITYPE Sweden Cortinarius dolabratus TN02-1095 Finland Cortinarius imbutoides KS-CO1576 Sweden Cortinarius phaeoruber RH80814 HOLOTYPE France Cortinarius dolabratus TN03-1713 Slovakia Cortinarius dolabratus SH188528.07FU NA-FS-Slovakia (10 seq) 75/Cortinarius dolabratus TN09-196 USA Cortinarius dolabratus AB98-09-94 Canada Cortinarius dolabratus Cortinarius dolabratus TN02-959 Finland Cortinarius dolabratus TN09-139 USA Cortinarius dolabratus H6033519 Finland Cortinarius dolabratus XC2013-103 France Cortinarius dolabratus IK95-347 Finland Cortinarius dolabratus IK95-1576 Finland Cortinarius dolabratus TN12-200 USA Cortinarius imbutoides AB04-09-186 HOLOTYPE France Cortinarius imbutoides KS-CO1290 Sweden Cortinarius dolabratus TN11-246 USA Cortinarius dolabratus IK02-033 Finland Cortinarius dolabratoides H6033615 Finland Cortinarius dolabratoides AB07-08-48 France Cortinarius dolabratoides H6033567 HOLOTYPE Finland Cortinarius dolabratoides Cortinarius dolabratoides IK04-051 Finland Cortinarius dolabratoides H6033575 Finland Cortinarius dolabratoides H6033570 Finland Cortinarius dolabratoides IK01-062 Finland Cortinarius turgidipes AB93-10-425 HOLOTYPE France Cortinarius turgidipes Cortinarius refectus AB05-09-138 France Cortinarius testaceoviolaceus AB92-10-293 France Cortinarius refectus PML2159 France Cortinarius refectus AB04-10-321 France Cortinarius refectus AB96-09-73 EPITYPE Germany Cortinarius refectus PML769 France Cortinarius scriptor AB94-10-268 France Cortinarius refectus IK96-1031 Germany Cortinarius refectus PML17 France 79/Cortinarius refectus AB99-09-121 France Cortinarius plumulosus RH3417 HOLOTYPE France Cortinarius fundatus PML657 France Cortinarius plumulosus PML3308 France Cortinarius plumulosus TN04-730 Finland (short) Cortinarius fundatus AB98-09-119 Canada Cortinarius fundatus AB10-09-183 France Cortinarius plumulosus IK98-1612 Finland Cortinarius minicolor AB92-10-256 France Cortinarius cagei PML1057 France Cortinarius cagei SH188634.07FU Germany-Italy (2 seq) Cortinarius cagei CFP1260 Sweden NEOTYPE 91/0.94 Cortinarius cagei PML3588 France Cortinarius minicolor PML738 France Cortinarius minicolor AB04-09-266 France Cortinarius periodolens ad int. XC2014-02 France Cortinarius sp. UBCOGTR194 Canada Cortinarius sp. TN12-217 USA Cortinarius evernius AB04-09-212 France Cortinarius evernius PML376 France Cortinarius evernius SH188514.07FU Canada-FS-Germany (11 seq) Cortinarius evernius TN05-238 Norway Cortinarius evernius f. pseudoscutulatus AB00-09-83 HOLOTYPE France Cortinarius evernius PML230 France Cortinarius evernius TN07-328 Canada Cortinarius evernius f. pseudoscutulatus AB91-08-42 France 66/Cortinarius evernius TN07-223 Canada Cortinarius parvulior ad int. AB96-09-47 France Cortinarius evernius f. fragrans PML1727 HOLOTYPE France Cortinarius evernius IK97-123 Finland Cortinarius evernius TN07-312 Canada Cortinarius evernius CFP792 NEOTYPE Sweden Cortinarius evernius IK00-038 Finland Cortinarius evernius PML622 France Cortinarius evernius TN10-054 Canada Cortinarius evernius var. insignis AB09-07-44 France Cortinarius evernius TN10-055 Canada Cortinarius evernius PML212 France Cortinarius evernius var. evernius PML3469 France Cortinarius evernius TN10-074 Canada Cortinarius sp. TN05-033 Finland Cortinarius hircinosmus Cortinarius refectus Cortinarius plumulosus Cortinarius cagei Cortinarius sp1 Cortinarius evernius Cortinarius sp2 0.005 substitution per site Fig. 2 The morphogenetic Bicolores section. — Bayesian 50 % majority-rule consensus tree inferred from the analysis of the ITS sequence of 124 (153 represented, 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 Cortinarius imbutus IK97-1162 NEOTYPE Finland Cortinarius imbutus IK98-2242 Sweden Cortinarius imbutus AB08-10-307 France Cortinarius imbutus asp. imbutus AB09-11-471 France Cortinarius imbutus asp. imbutus AB98-10-358 France Cortinarius imbutus asp. imbutus AB10-10-237 France Cortinarius imbutus asp. imbutus PML375 France Cortinarius imbutus asp. laetior XC2013-13 France Cortinarius imbutus asp. saturnalis XC2014-77 France Cortinarius betulaecomes RH3123 HOLOTYPE France Cortinarius imbutus AB02-09-58 France Cortinarius imbutus asp. imbutus XC2002-107 France Cortinarius imbutus AB02-10-106 France Cortinarius imbutus asp. imbutus AB04-09-228 France Cortinarius imbutus asp. imbutus XC2002-106 France Cortinarius imbutus XC2012-96 France Cortinarius saturninus SH188563.07FU Canad stonia-China (6 seq) Cortinarius imbutus AB00-09-127 France Cortinarius betulaecomes R 30 France Cortinarius imbutus asp. imbutus XC2002-108 France Cortinarius imbutus TN11-257 USA Cortinarius imbutus XC2002-109 France Cortinarius imbutus TN05-167 Finland Cortinarius imbutus TN11-151 USA Cortinarius imbutus asp. imbutus XC2002-122 France Cortinarius laccatus PML4557 HOLOTYPE France Cortinarius imbutus JMB2008092703 France Cortinarius imbutus TN11-252 USA Cortinarius imbutus asp. vilior XC2007-104 France Cortinarius imbutus asp. saturnalis XC2014-61 France Cortinarius imbutus TN11-150 USA Cortinarius imbutus IK94-1236 Finland Cortinarius confirmatus asp. assiduus AB02-11-201 France Cortinarius confirmatus asp. subcylindratus XC2013-160 France Cortinarius confirmatus asp. assiduus AB09-11-514 France Cortinarius confirmatus FR2405 France Cortinarius sp. SH094374.07FU U-Iran (6 seq) Cortinarius confirmatus asp. assiduus AB05-11-423 France Cortinarius confirmatus FR6052 France Cortinarius confirmatus RH3195 HOLOTYPE France Cortinarius confirmatus asp. assiduus XC2013-156 France Cortinarius confirmatus FR2076 France Cortinarius confirmatus XC2006-204 France Cortinarius confirmatus asp. imbutus XC2012-171 France Cortinarius confirmatus XC2005-249 France Cortinarius confirmatus FR2089 France Cortinarius confirmatus asp. spurcatocephalus XC2011-199 France Cortinarius confirmatus asp. rubricosissimus AB09-11-452 France Cortinarius bulbosovolvatus RH84159 ISOTYPE France Cortinarius assiduus var. plesiocistus JVG990125-31 ISOTYPE Spain Cortinarius confirmatus asp. imbutus PML4722 France Cortinarius confirmatus asp. spurcatocephalus XC95-10-04-06 France Cortinarius assiduus MES3541 HOLOTYPE Spain Cortinarius confirmatus AB09-11-450 France Cortinarius cistoadelphus ad int. AB92-11-422 France Cortinarius confirmatus asp. paracohabitans AB11-11-324 France Cortinarius confirmatus asp. confirmatus AB03-11-78 France Cortinarius confirmatus asp. kuehneri AB13-10-97 France Cortinarius confirmatus asp. rubricosissimus AB00-10-193 France Cortinarius saturninus XC2006-194 France Cortinarius salicis RH2623 HOLOTYPE France 79/0.81 Cortinarius saturninus XC2008-61 France * Cortinarius saturninus asp. salicis XC2011-205 France Cortinarius fulvorimosus XC2007-14 HOLOTYPE France Cortinarius saturninus XC2002-167 France Cortinarius saturninus asp. deceptivus AB04-10-344 France Cortinarius saturninus asp. urbicoides XC2001-107 France Cortinarius saturninus asp. urbicoides AB95-11-144 France Cortinarius saturninus XC2001-104 France Cortinarius saturninus asp. cohabitans XC2014-63 France Cortinarius saturninus XC2007-90 France Cortinarius subtorvus CFP408 Sweden Cortinarius saturninus TN09-208 USA Cortinarius saturninus asp. salicis XC2007-108 France Cortinarius saturninus H6029320 Finland Cortinarius saturninus asp. urbicoides AB02-10-179 France Cortinarius urbicus AB97-09-187/PML5347 France Cortinarius saturninus XC2016-12 France Cortinarius urbicus PML75 France (short) Cortinarius umbrinoconnatus RH476 HOLOTYPE France (short) Cortinarius saturninus asp. salicis XC2014-109 France Cortinarius saturninus AB05-10-273 France Cortinarius saturninus asp. saturninus XC2007-97 France 51/Cortinarius salicis PML3967 France Cortinarius saturninus JMB2009101002 France Cortinarius saturninus asp. dionisiae AB14-09-47 France Cortinarius saturninus IK94-631 Finland Cortinarius subtorvus O50591 Norway (Svalbard) Cortinarius saturninus asp. saturninus XC2014-114 France Cortinarius marginatosplendens PML2215 ISOTYPE France Cortinarius saturninus asp. deceptivus AB98-10-381 France Cortinarius saturninus CFP514 NEOTYPE Sweden Cortinarius dissidens PR258 HOLOTYPE France Cortinarius saturninus XC96-10-26-09 France Cortinarius denseconnatus RH3758 HOLOTYPE France Cortinarius urbicus var. sporanotandus PML4578 HOLOTYPE France Cortinarius saturninus asp. salicis XC2008-55 France Cortinarius gramineus RH81181 HOLOTYPE France Cortinarius saturninus asp. salicis AB14-11-160 France Cortinarius saturninus asp. saturninus XC2014-116 France Cortinarius subtorvus KH14 Norway (Svalbard) Cortinarius rastetteri RH71682 HOLOTYPE France Cortinarius saturninus SH094324.07FU USA-U (13 seq) Cortinarius cyprinus PML425 France Cortinarius cyprinus XC2013-15 France Cortinarius cyprinus XC2007-95 France Cortinarius cyprinus XC2007-103 France Cortinarius cyprinus AB11-11-251 France Cortinarius cyprinus AB06-09-144 France Cortinarius cyprinus PAM13092901 France Cortinarius sp. TAAM12 stonia Cortinarius cyprinus PML81 France Cortinarius cyprinus JMB2014111802 France Cortinarius cyprinus AB04-09-167 France Cortinarius cyprinus PML344 France Cortinarius cyprinus XC2012-26 HOLOTYPE France Cortinarius cyprinus AB11-10-192 France Cortinarius cyprinus TB348-10 Norway Cortinarius stuntzii Rehner394 HOLOTYPE USA Cortinarius lucorum CFP490 NEOTYPE Norway Cortinarius montis-dei PML4142 HOLOTYPE France Cortinarius lucorum asp. circumvelatus PAM14090808 France Cortinarius incarnatolilascens RH71502 HOLOTYPE France 85/0.84 Cortinarius lucorum IK89-748 Finland Cortinarius lucorum KS-CO513 Sweden Cortinarius lucorum TN03-1169 Sweden 56/Cortinarius lucorum SH188495.07FU NA-FS (21 seq) Cortinarius lucorum asp. montis-dei PML4143 France Cortinarius circumvelatus PML34 HOLOTYPE France Cortinarius lucorum TN10-002 Canada Cortinarius umidicola 10433 SYNTYPE USA (short) 92/0.87 58/0.92 87/0.3 96/- 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 Cortinarius imbutus Cortinarius confirmatus Cortinarius saturninus Cortinarius cyprinus Cortinarius stuntzii Cortinarius lucorum 0.004 substitution per site Fig. 3 The morphogenetic Saturnini section. — Bayesian 50 % majority-rule consensus tree inferred from the analysis of the ITS sequence of 131 (173 represented, 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 2011205 (within the subclade) was fruiting from the same mycelium as XC 2007-108 and XC 2014-109 (outside the subclade). Species Blue hues a Odour(s) b Cortinarius cinnamoviolaceus + or – R, r, i L min Av L L max l min Av l l max Av Q Reported host c,d Dintra max / difference rate (incl. indels) a Dinter min / difference rate (incl. indels) c 8.25 9.65 11.07 4.66 5.21 6.00 1.86 Picea, Abies, Pinus, Tilia, Quercus, Betula, Populus na na 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 % 9.86 3 nts (to C. dolabratoides) / 0.5 % C. dolabratus + or – ce, co 7.42 8.62 4.41 4.90 5.51 1.76 Pinus, Picea, Betula, Fagus, Quercus 3 nts / 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 %) C. glaphurus + or – ce, r, CE, V 8.03 10.60 4.82 5.23 5.78 1.78 Pinus, Quercus, Fagus, Abies, Picea, Populus, Betula 2 nts + 4 indels / 1 % 9.32 3 nts + 3 indels (to C. cagei) / 1 % 4 nts + 2 indels (to C. tortuosus) / 1 % 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 % + or – 0, ca, r, g 6.91 8.26 9.79 4.27 4.79 5.61 1.73 Quercus, Cistus, Pinus, Betula, Populus, Picea 6 nts + 1 indel / 1.2 % 3 nts (to C. imbutus) / 0.5 % 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. sect. saturnini C. confirmatus 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, Carpinus, Picea 3 nts + 1 indel (0.7 %) 3 nts (to C. confirmatus) / 0.5 % C. lucorum + r, ca, 0 8.07 9.56 11.07 5.36 5.86 6.71 1.63 Populus, Betula, Carpinus, Quercus, Picea, Tsuga 2 nts + 1 indel (0.5 %) 16 nts + 3 indels (to C. confirmatus) / 3.1 % 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 % 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 + 2 indels (to C. cyprinus) / 0.8 % 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. 189 190 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 restricted 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 taxonomic 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 deposited 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. dolabratus); 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. cinnamoviolaceus; ii. typical Duracini concepts – C. subparevernius, C. cylindratus 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. Cortinarius 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. cinnamoviolaceus looks like a Duracini with reddish lamellae and is Persoonia – Volume 39, 2017 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 outside 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, usually 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 — MycoBank 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 Cortinaires 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 deposited 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. minicolor) 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 nomenclatural issue associated with C. bicolor Cooke, an illegitimate name because of an earlier use of the name for another, unrelated 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, biogeographical 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, however, 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 emarginate, 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 verrucose, often prominently more strongly at the very apex, somewhat 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 occasional. Other specimens examined (sequenced collections marked with an asterisk, see Table 2 for GenBank accession numbers). FinlAnd, VarsinaisSuomi, 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. dolabratoides 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). a b c 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. 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. Lindström, H. Marklund, S. Muskos CFP990, S, epitype designated here, MycoBank 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. imbutoides); Niskanen et al. 2012: 863; Brandrud et al. 1998: pl. D52. Notes — The original description of C. dolabratus is apparently 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 lacking 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 a b c d 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. 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). Cortinarius 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 considering 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. cedriosmus, C. paranomalus, C. tubulosus and C. violaeolens, as well as C. turibulosus sensu Bidaud et al. (2010). The protologue 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 membranous 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 sister 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. camphoratus 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 obviously 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 ecological 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 tortuosus (Fr.) Fr., Epicr. Syst. Mycol.: 305. 1838 — MycoBank MB#165676 – Cortinarius fundatus Britzelm., Ber. Naturhist. Vereins Augsburg 28: 127. 1885, sensu Bidaud et al. (2014). 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. 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. fundatus); 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. evernius (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 deciduous 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 Augsburg 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. 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 notable exception of J. Favre, who referred to this species as C. plumbosus – so that C. tortuosus taxonomy is not a problematic 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, subcespitose, 19 Oct. 1993, A. & E. Bidaud, AB 93-10-425, PC, holotype, MycoBank MBT#372786. ITS sequence deposited in GenBank under KX964409. Illustrations — This study: Fig. 5c; Bidaud et al. 2014, pl. 952, 953 but also pl. 945 (as C. testaceoviolaceus). Illustration — Bidaud et al. 2008, pl. 672. Taxonomic descriptions — Bidaud et al. 2014: f. 1410 but also 2010: f. 1109 (as C. scriptor). Notes — More collections of this species, originally described in sect. Damasceni by its authors, are required to better assess its morphogenetic variability as well as to deﬁne its ecological 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). Notes — No original material was kept by Britzelmayr to assign 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. testaceoviolaceus 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). Taxonomic description — Bidaud et al. 2008: f. 885. 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 ochraceous, 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 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 himself 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. cohabitans, usually considered as akin to C. saturninus, 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, 195 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. saturninus may also be associated with other deciduous, but also coniferous trees. Morphologically, the species displays unprecedented 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 hygrophilous 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. Saturnini (see notes under C. confirmatus, C. cyprinus and C. imbutus), 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 whitish 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. Méditerranéennes 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 deposited 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 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 dramatically 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 occurring 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. assiduus 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. confirmatus within the boundaries of its most inclusive clade and introduce C. confirmatus var. plesiocistus (A. Ortega, Vila & Bidaud) comb. nov. to accommodate the cisticolous populations. 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 Kryptogamenflora von Mitteleuropa II: 162. 1953, nom inval. (ined.)]. – Cortinarius cypriacus Fr., Epicr. Syst. Mycol.: 307. 1838, sensu MoënneLoccoz & Reumaux (1989), non Consiglio (1999). Type. FrAnce, Yvelines, Gambais, under deciduous trees, on calcareous soil, 3 Oct. 1993, G. Redeuilh, XC 2012-26, PC, holotype, MycoBank MBT#373189. ITS sequence deposited in GenBank under KX964463. Persoonia – Volume 39, 2017 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. circumvelatus (= 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, molecular 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 collection 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 substitutions. 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 — MycoBank 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 species 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 colour of the pileus and the intensity of blue tinges in basidiomata. Typically, the species fruits under deciduous trees in hygrophilous 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, sometimes 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. incarnatolilascens and C. montis-dei, on the basis of deviating macromorphological or ecological features while oddly, their initial – 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 contemporary 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 species (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 surprisingly, 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 topology 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 fruiting 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 Populusassociated 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 additional 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 deposited 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 placement 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 members 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 octopetala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 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 verrucose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 different . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 Persoonia – Volume 39, 2017 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 Cortinarius, 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 sections 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 species, 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 phylogenetic 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. cedriosmus, 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. circumvelatus, 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 all associated with different host trees within their respective clades. Spores and host plants are usually considered as reliable 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 support 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, making 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 Telamonia 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-imbutuscypriacus, but shown here to belong in sect. Bicolores (Fig. 2). Interestingly, the present work yields strong phylogenetic support to the prospective placement or overlap of the morphologically deﬁned sect. Bicolores and sect. Duracini. Natural relationships 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. cinnamoviolaceus 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. parevernius, 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 evolutionary 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 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 identiﬁcation and hierarchical organization of visible characteristics – 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 necessarily subjective nature of the selection process involved in this approach has led to highly artiﬁcial groupings at multiple taxonomic levels (i.e., Aphyllophorales, Clavariaceae, Clitocybe, 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 characteristics that delinate a lineage with high taxonomic value can now be distinguished from those, less valuable and taxonomically 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 several 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. 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Cortinarius section Bicolores and section Saturnini (Basidiomycota, Agaricales), a morphogenetic overview of European and North American species

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

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