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

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.

INTRODUCTION

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 morpho-anatomical and ecological traits, DNA phylogenies revealed characters that have been overemphasized in monographic studies but also uncovered significant taxonomic information that has been neglected by previous investigators (Bellanger et al. 2015, Loizides et al. 2016). The use of these modern tools a posteriori, to test the autonomy of previously defined 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.

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, Moënne-Loccoz & Reumaux 1990). In spite of their practical application, most of these lower level taxonomic divisions have proven to be artificial when placed under evolutionary scrutiny (Garnica et al. 2005). Subgenus Telamonia, however, breaks this rule as most of the numerous species known to date that produce dry-capped basidiomata lacking vivid colours – the 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 sect. Armillati, Brunnei, Bovini and Disjungendi and more are on their way to morphogenetic redefinition (Niskanen et al. 2009, 2011, 2013, Liimatainen et al. 2014a).

Here we deal with Cortinarius sect. Bicolores and Cortinarius sect. Saturnini, which encompass Cortinarius evernius, C. 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 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:

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. cagei	C. bicolor?	C. bicolor?
	C. periodolens ad. int.
C. dolabratoides sp. nov.
C. dolabratus	C. imbutoides
C. evernius	C. evernius, C. parvulior ad. int.	C. evernius	C. evernius, C. scutulatus	C. evernius
C. glaphurus	C. tubulosus, C. paranomalus (Sat.)
C. hircinosmus	C. livor		C. livor?
C. plumulosus	C. fundatus		C. bicolor?	C. bicolor?
C. refectus	C. refectus, C. testaceoviolaceus		C. bicolor?	C. bicolor?
C. sp1
C. sp2
C. tortuosus	C. tortuosus	C. tortuosus	C. plumbosus	C. tortuosus, C. plumbosus
C. turgidipes
C. cinnamoviolaceus	C. cinnamoviolaceus, C. parevernius,	C. imbutus	C. cinnamoviolaceus,	C. parevernius
	C. basicyaneus		C. parevernius
C. disjungendus	C. cyanosterix
C. mattiae	C. mattiae	C. mattiae	C. subviolascens
C. parevernioides	C. parevernioides
C. salicinus	C. salicinus, C. deceptivoides
	C. quadricolor
Sect. Saturnini	Sect. Saturnini	Sect. Firmiores + sect. Telamonia	Key 3.11.7.6.11	Sect. Bicolores
C. confirmatus	C. confirmatus
C. cyprinus	C. cyprinus
C. imbutus	C. imbutus		C. vilior	C. imbutus
C. lucorum	C. lucorum	C. lucorum	C. lucorum, C. umidicola
C. saturninus	C. saturninus	C. saturninus, C. subtorvus	C. saturninus,
			C. deceptivus,
			C. subtorvus
C. stuntzii
C. cypriacoides	C. cypriacoides		C. cypriacus	C. cypriacus
C. furiosus	C. furiosus
C. nefastus	C. nefastus
C. serratissimus*	C. saturninoides		C. serratissimus	C. saturninus
C. sciophylloides	C. sciophylloides
C. subbulliardioides*	C. illepidus
C. subfirmus	C. subfirmus
C. suboxytoneus	C. suboxytoneus, C. fuscocinctus
			C. sciophyllus	C. sciophyllus
			C. castaneus	C. castaneus
			C. calopus
			C. torvus
			C. impennis
				C. myrtillinus

Bold names indicate sequenced species. Dotted lines separate morphogenetic species included in /Bicolores and /Saturnini (upper parts) from those (morphological species, lower parts) phylogenetically unrelated to the two clades. (Sat.), Saturnini. Asterisk indicates unpublished data of nomenclatural significance.

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.

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.

Table 2

Specimens included in phylogenetic analyses.

Species C. = Cortinarius	Voucher/SH	Voucher/SH annotation	Leg.	Collection date	Country	Taxonomy	Herbarium	Accession*
/Bicolores
C. cagei	CFP 1260	cagei (neotype)	T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos	1994	Sweden	CFP: D48 (1998)	S	KX964295
	AB 04-09-266	minicolor	A. Bidaud	2004	France	AC 22: f1419 (2014)	ADC private	KX964296
	AB 92-10-256	minicolor	A. Bidaud & R. Fillion	1992	France	AC 22: f1419 (2014)	ADC private	KX964297
	PML 738	minicolor	R. Fillion	1987	France	AC 22: f1419 (2014)	ADC private	KX964298
	XC 2014-02	periodolens ad int.	A. Ferville	1993	France	AC 22: f1417 (2014)	ADC private	KX964299
	PML 3588	basicyaneus	A. Ferville	1993	France	this study	ADC private	KX964300
	PML 1057	basicyaneus	R. Fillion	1988	France	this study	ADC private	KX964301
	SH188634.07FU (2 sequences)	cagei	na	na	Germany/Italy	na	na	AY669676
C. dolabratoides sp. nov.	H:6033567	sp. (holotype)	I. Kytövuori	2008	Finland	this study	H	KX964302
	AB 07-08-48	marcellae cf.	A. Bidaud & R. Fillion	2007	France	this study	ADC private	KX964303
	H:6033615	sp.	I. Kytövuori	2004	Finland	this study	H	KX964304
	H:6033575	sp.	I. Kytövuori	2008	Finland	this study	H	KX964305
	H:6033570	sp.	I. Kytövuori	2008	Finland	this study	H	KX964306
	IK 04-051	‘smell-of-viola’	I. Kytövuori	2004	Finland	this study	H	KX964307
	IK 01-062	‘smell-of-viola’	I. Kytövuori	2001	Finland	this study	H	KX964308
C. dolabratus	CFP 990	dolabratus (epitype)	T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos	1990	Sweden	CFP: D52 (1998)	S	KX964309
	AB 04-09-186	imbutoides (holotype)	A. Bidaud	2004	France	AC 22: f1409 (2014)	PC	KX964310
	RH 80814	phaeoruber (holotype)	G. Chevassut	1980	France	DM 12(47): 52 (1982)	PC	KX964311
	AB 13-10-120	saturninus cf.	A. Bidaud	2013	France	this study	ADC private	KX964312
	AB 04-09-169	armillariellus cf.	A. Bidaud	2004	France	this study	ADC private	KX964313
	AB 01-09-41	privignus sensu Quélet cf.	A. Bidaud	2001	France	this study	ADC private	KX964314
	AB 98-09-94	saturninus cf.	A. Faurite	1998	Canada	this study	ADC private	KX964315
	AB 89-11-309	orastriatus	A. Bidaud	1989	France	this study	ADC private	KX964316
	H:6033519	dolabratus	I. Kytövuori	2001	Finland	this study	H	KX964317
	IK 02-033	dolabratus	I. Kytövuori	2002	Finland	this study	H	KX964318
	IK 95-1576	dolabratus	I. Kytövuori	1995	Finland	this study	H	KX964319
	IK 95-347	dolabratus	I. Kytövuori	1995	Finland	this study	H	KX964320
	KS CO1576	imbutoides	K. Soop	2005	Sweden	this study	K. Soop private	KX964321
	KS CO1290	imbutoides	K. Soop	2001	Sweden	this study	K. Soop private	KX964322
	TN 12-200	dolabratus	T. Niskanen	2012	USA	this study	H	KX964323
	TN 11-246	dolabratus	T. Niskanen	2011	USA	this study	H	KX964324
	TN 09-196	dolabratus	T. Niskanen	2009	USA	this study	H	KX964325
	TN 09-139	dolabratus	T. Niskanen	2009	USA	this study	H	KX964326
	TN 03-1713	dolabratus	T. Niskanen	2003	Slovakia	this study	H	KX964327
	TN 02-1095	dolabratus	T. Niskanen	2002	Finland	this study	H	KX964328
	TN 02-959	dolabratus	T. Niskanen	2002	Finland	this study	H	KX964329
	XC 2013-103	privignus sensu Quélet	P. Reumaux	1998	France	this study	ADC private	KX964330
	SH188528.07FU (10 sequences)	dolabratus	na	na	NA/FS/Slovakia	na	na	UDB018659
C. evernius	CFP 792	evernius (neotype)	T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos	1988	Sweden	CFP: A11 (1990)	S	KX964331
	AB 00-09-83	evernius f. pseudoscutulatus (holotype)	A. Bidaud	2000	France	AC 22: f1407 (2014)	PC	KX964332
	PML 1727	evernius f. fragrans (holotype)	D. Mazuir	1990	France	AC 22: f1406 (2014)	PC	KX964333
	AB 96-09-47	parvulior ad int.	M. Martin	1996	France	AC 22: f1418 (2014)	ADC private	KX964334
	AB 91-08-42	evernius f. pseudoscutulatus	A. Bidaud & C. Blanc	1991	France	AC 22: f1407 (2014)	ADC private	KX964335
	PML 622	evernius f. fragrans	P. Moënne-Loccoz	1987	France	AC 22: f1406 (2014)	ADC private	KX964336
	AB 09-07-44	evernius var. insignis	A. Bidaud & A. Faurite	2009	France	AC 22: f1405 (2014)	ADC private	KX964337
	PML 3469	evernius var. evernius	A. Bidaud	1993	France	AC 22: f1404 (2014)	ADC private	KX964338
	PML 212	evernius var. evernius	P. Moënne-Loccoz	1982	France	AC 22: f1404 (2014)	ADC private	KX964339
	AB 04-09-212	evernius	A. Bidaud	2004	France	this study	ADC private	KX964340
	IK 00-038	evernius	I. Kytövuori	2000	Finland	this study	H	KX964341
	IK 97-123	evernius	I. Kytövuori	1997	Finland	this study	H	KX964342
	PML 376	evernius	P. Moënne-Loccoz	1986	France	this study	ADC private	KX964343
	PML 230	evernius	P. Moënne-Loccoz	1984	France	this study	ADC private	KX964344
	TN 10-074	evernius	T. Niskanen	2010	Canada	this study	H	KX964345
	TN 10-055	evernius	T. Niskanen	2010	Canada	this study	H	KX964346
	TN 10-054	evernius	T. Niskanen	2010	Canada	this study	H	KX964347
	TN 07-328	evernius	T. Niskanen	2007	Canada	this study	H	KX964348
	TN 07-312	evernius	T. Niskanen	2007	Canada	this study	H	KX964349
	TN 07-223	evernius	T. Niskanen	2007	Canada	this study	H	KX964350
	TN 05-238	evernius	T. Niskanen	2005	Norway	this study	H	KX964351
	SH188514.07FU (11 sequences)	evernius	na	na	Canada/FS/Germany	na	na	AY669686
C. glaphurus	RH 71421	glaphurus (holotype)	G. Chevassut	1978	France	DM 12(47): 78 (1982)	PC	KX964352
	AB 03-11-87	tubulosus (holotype)	A. Bidaud & A. Faurite	2003	France	AC 22: f1414 (2014)	PC	KX964353
	AB 92-10-350	cedriosmus (holotype)	A. Bidaud	1992	France	AC 19: f1144 (2010)	PC	KX964354
	XC 2009-41	violaeolens (holotype)	A. & R. Bardet	1996	France	AC 19: f1112 (2010)	PC	KX964355
	GK1142	paranomalus (holotype)	G. Redeuilh	1987	France	K&R: 305 (1953, invalid), AC 4: f163 (1992)	GK	KX964356
	AB 08-11-445	tubulosus	M. Martin	2008	France	AC 22: f1414 (2014)	ADC private	KX964357
	AB 92-10-332	turibulosus	R. Fillion	1992	France	AC 19: f1108 (2010)	ADC private	KX964358
	AB 91-11-360	turibulosus	A. Bidaud	1991	France	AC 19: f1108 (2010)	ADC private	KX964359
	PML 2390	turibulosus	P. Moënne-Loccoz	1991	France	AC 19: f1108 (2010)	ADC private	KX964360
	PML 1067	turibulosus	P. Moënne-Loccoz	1988	France	AC 19: f1108 (2010)	ADC private	KX964361
	AB 14-11-138	minicolor cf.	P.-Y. Courio	2014	France	this study	ADC private	KX964362
	AB 03-10-56	sciophyllus cf.	A. Bidaud	2003	France	this study	ADC private	KX964363
	AB 99-11-345	livor cf.	M. Martin	1999	France	this study	ADC private	KX964364
	TN 12-221	sp.	T. Niskanen	2012	USA	this study	H	KX964365
	XC 2011-212	laetior cf.	X. Carteret	2011	France	this study	ADC private	KX964366
	XC 2009-64	paranomalus cf.	X. Carteret	2009	France	this study	ADC private	KX964367
	SH094444.07FU (3 sequences)	turibulosus	na	na	NA/France	na	na	GQ159774
	SH094485.07FU (2 sequences)	sp.	na	na	Poland	na	na	HQ115588
C. hircinosmus	PML 334	hircinosmus (holotype)	P. Moënne-Loccoz	1986	France	AC 12: f575 (2002)	PC	KX964368
	AB 02-09-32	livor	A. Bidaud	2002	France	AC 23: f1459 (2015)	ADC private	KX964369
	F44390	sp.	K. Soop	na	Sweden	FN: 849 (2012)	S	KX964370
	H:6033565	hircinosmus	I. Kytövuori	2009	Finland	FN: 849 (2012)	H	KX964371
	AB 97-10-341	scriptor	G. Chamonaz	1997	France	AC 19: f1109 (2010)	ADC private	KX964372
	AB 04-10-357	imbutus cf.	A. Bidaud	2004	France	this study	ADC private	KX964373
C. plumulosus	RH 3417	plumulosus (holotype)	R. Henry	1972	France	SMF 93(3): 359 (1977)	PC	KX964374
	AB 10-09-183	fundatus	A. Bidaud & R. Fillion	2010	France	AC 22: f1411 (2014)	ADC private	KX964375
	AB 98-09-119	fundatus	E. & A. Bidaud, A. Faurite	1998	Canada	AC 22: f1411 (2014)	ADC private	KX964376
	PML 657	fundatus	P. Moënne-Loccoz	1987	France	AC 22: f1411 (2014)	ADC private	KX964377
	PML 3308	perscutulatus	A. Bidaud	1992	France	this study	ADC private	KX964378
	IK 98-1612	sp.	I. Kytövuori	1998	Finland	this study	H	KX964379
	TN 04-730	sp.	T. Niskanen	2004	Finland	this study	H	KX964380
C. refectus	AB 96-09-73	refectus (epitype)	A. Bidaud	1996	Germany	AC 22: f1410 (2014)	PC	KX964385
	AB 05-09-138	refectus	A. Bidaud	2005	France	AC 22: f1410 (2014)	ADC private	KX964382
	AB 04-10-321	refectus	A. Bidaud	2004	France	AC 22: f1410 (2014)	ADC private	KX964383
	AB 99-09-121	refectus	A. Bidaud	1999	France	AC 22: f1410 (2014)	ADC private	KX964384
	PML 2159	refectus	A. Bidaud	1990	France	AC 22: f1410 (2014)	ADC private	KX964386
	PML 769	refectus	P. Moënne-Loccoz	1987	France	AC 22: f1410 (2014)	ADC private	KX964387
	PML 17	refectus	P. Moënne-Loccoz	1985	France	AC 22: f1410 (2014)	ADC private	KX964388
	AB 92-10-293	testaceoviolaceus	A. Bidaud	1992	France	AC 22: f1402 (2014)	ADC private	KX964389
	AB 94-10-268	scriptor	A. Bidaud	1994	France	AC 19: f1109 (2010)	ADC private	KX964390
	IK 96-1031	refectus	I. Kytövuori	1996	Germany	this study	H	KX964381
C. tortuosus	IB 79/533	tortuosus (neotype)	D. Lamoure	1979	Sweden	Opera Botanica 100: 182 (1989)	IB	KX964391
	XC 2008-43	flabelloides (holotype)	M. Pèlerin	2008	France	AC 19: f1136 (2010)	PC	KX964392
	PAK 354	laetior (holotype)	P.A. Karsten	1879	Finland	BFNF 32: 387 (1879)	H	KX964393
	AB 01-09-19	tortuosus	A. Bidaud	2001	France	AC 22: f1413 (2014)	ADC private	KX964394
	AB 96-08-19	tortuosus	A. Bidaud	1996	France	AC 22: f1413 (2014)	ADC private	KX964395
	AB 95-09-34	tortuosus	C. Blanc	1995	France	AC 22: f1413 (2014)	ADC private	KX964396
	PML 3551	tortuosus	A. Bidaud & R. Fillion	1993	France	AC 22: f1413 (2014)	ADC private	KX964397
	PML 1225	tortuosus	P. Moënne-Loccoz	1989	France	AC 22: f1413 (2014)	ADC private	KX964398
	PML 1214	tortuosus	P. Moënne-Loccoz	1989	France	AC 22: f1413 (2014)	ADC private	KX964399
	PML 386	tortuosus	P. Moënne-Loccoz	1986	France	AC 22: f1413 (2014)	ADC private	KX964400
	CFP 493	tortuosus	T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos	1986	Norway	CFP: A06 (1990)	S	KX964401
	AB 02-09-41	saturninus cf.	A. Bidaud	2002	France	this study	ADC private	KX964402
	AB 96-10-124	saturninus cf.	C. Blanc	1996	France	this study	ADC private	KX964403
	IK 99-709	tortuosus	I. Kytövuori	1999	Finland	this study	H	KX964404
	TN 10-087	tortuosus	T. Niskanen	2010	Canada	this study	H	KX964405
	TN 09-046	tortuosus	T. Niskanen	2009	USA	this study	H	KX964406
	TN 07-307	tortuosus	T. Niskanen	2007	Canada	this study	H	KX964407
	TN 05-006	tortuosus	T. Niskanen	2005	Finland	this study	H	KX964408
	SH094369.07FU (7 sequences)	tortuosus	na	na	USA/U/Japan	na	na	AY669669
C. turgidipes	AB 93-10-425	turgidipes (holotype)	A. & E. Bidaud	1993	France	AC 17(1): f885 (2008)	PC	KX964409
C. sp1	TN 12-217	sp.	T. Niskanen	2012	USA	na	H	KX964410
	UBCOGTR194	sp. (ectomycorrhiza)	na	na	Canada	na	na	EU597034
C. sp2	TN 05-033	sp.	T. Niskanen	2005	Finland	na	H	KX964411
Other (morphological) Bicolores
C. cinnamoviolaceus	IB 48/590	cinnamoviolaceus (holotype)	M. Moser	1948	Austria	Nova Hedwigia XIV(2-4): 514 (1967)	IB	KX964412
	RH 70942	basicyaneus (holotype)	M. Trimbach	1976	France	FAMM 25: 38 (2004)	PC	KX964413
	RH 4000	cylindratus (holotype)	R. Henry	1972	France	SMF 99(1): 91 (1983)	PC	KX964414
	RH 526	subparevernius (holotype)	R. Henry	1956	France	SMF 85(4): 442 (1969)	PC	KX964415
	RH 1240	contractus (holotype)	R. Henry	1960	France	SMF 85(4): 387 (1969)	PC	KX964416
	RH 3258a78	parevernius (holotype)	R. Henry	1955	France	K&R: 303 (1953, invalid)	PC	KX964417
	AB 02-10-71	dolabratus	A. & M. Burat	2002	France	AC 17(1): f817 (2008)	ADC private	KX964418
	CFP 574	imbutus	T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos	1987	Sweden	CFP: D60 (1998)	S	KX964419
	AB 12-11-240	imbutus	A. Bidaud	2012	France	this study	ADC private	KX964420
	TN 05-198	imbutus sensu Funga Nordica	T. Niskanen	2005	Finland	this study	H	KX964421
	TN 05-051	imbutus sensu Funga Nordica	T. Niskanen	2005	Finland	this study	H	KX964422
	SH188640.07FU (2 sequences)	imbutus	na	na	Sweden/Italy	na	na	UDB001160
C. cyanosterix (= C. disjungendus)	RH 338	cyanosterix (holotype)	R. Henry	1952	France	SMF 71(3): 259, 261 (1956)	PC	KX964423
C. mattiae	KS CO1936	mattiae (isotype)	K. Soop	2009	Sweden	JEC 13(12): 3 (2010)	S	KX964424
	AB 13-08-35	mattiae	A. Bidaud, F. Armada & R. Fillion	2013	France	AC 22: f1415 (2014)	ADC private	KX964425
	AB 99-09-77	subviolascens	A. Bidaud	1999	France	AC 12: f565 (2002)	ADC private	KX964426
	PML 650	subviolascens	P. Moënne-Loccoz	1987	France	AC 12: f565 (2002)	ADC private	KX964427
	CFP 1204	mattiae	T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos	1993	Sweden	CFP: D30 (1998)	S	KX964428
	AB 06-09-153	licinipes/poecilopus aff.	A. Bidaud, F. Armada & R. Fillion	2006	France	this study	ADC private	KX964429
	H:6029375	mattiae	T. Niskanen	2004	Finland	this study	H	KX964430
	H:6000560	mattiae	I. Kytövuori	2007	Finland	this study	H	KX964431
	IK 01-039	mattiae	I. Kytövuori	2001	Sweden	this study	H	KX964432
	IK 98-1127	mattiae	I. Kytövuori	1998	Finland	this study	H	KX964433
	PML 3989	umbrinoconnatus forma	A. Bidaud	1993	France	this study	ADC private	KX964434
	PML 2298	oxytoneus	A. Bidaud	1991	France	this study	ADC private	KX964435
	SH009438.07FU (1 sequence)	sp.	na	na	Canada	na	na	FJ039684
C. parevernioides	AB 02-09-50	parevernioides (holotype)	C. Gérard	2002	France	AC 22: f1408 (2014)	PC	KX964436
	SH188502.07FU (15 sequences)	malachius	na	na	NA/U	na	na	KF617653
C. salicinus	XC 2014-03	salicinus (holotype)	C. Hugouvieux	2005	France	AC 22: f1416 (2014)	PC	KX964437
/Saturnini
C. confirmatus	RH 3195	confirmatus (holotype)	R. Henry	1970	France	SMF 99(1): 67 (1983)	PC	KX964438
	JVG 990125-31	assiduus var. plesiocistus (isotype)	X. Llimona & J. Vila	1999	Spain	Mycotaxon 101: 140 (2007)	J. Vila private	AM713178
	MES 3541	assiduus (holotype)	R. Mahiques	1999	Spain	FMDS 162: 42 (2001)	MES	KX964439
	RH 84/159	bulbosovolvatus (isotype)	M. Contu & L. Curreli	1984	Italy	DM 26 (61): 32 (1985)	PC	KX964440
	XC 2013-160	confirmatus ‘asp. subcylindratus’	na	2013	France	AC 23: f1441 (2015)	PC	KX964441
	AB 13-10-97	confirmatus ‘asp. kuehneri’	A. Bidaud	2013	France	AC 23: f1440 (2015)	ADC private	KX964442
	XC 2011-199	confirmatus ‘asp. spurcatocephalus’	X. Carteret	2011	France	AC 23: f1439 (2015)	ADC private	KX964443
	XC 95-10-04-06	confirmatus ‘asp. spurcatocephalus’	X. Carteret	1995	France	AC 23: f1439 (2015)	ADC private	KX964444
	AB 09-11-452	confirmatus ‘asp. rubricosissimus’	A. Bidaud	2009	France	AC 23: f1438 (2015)	ADC private	KX964445
	AB 00-10-193	confirmatus ‘asp. rubricosissimus’	A. Bidaud	2000	France	AC 23: f1438 (2015)	ADC private	KX964446
	AB 11-11-324	confirmatus ‘asp. paracohabitans’	F. Armada, A. Bidaud & J. Pardo	2011	France	AC 23: f1437 (2015)	ADC private	KX964447
	PML 4722	confirmatus ‘asp. imbutus’	P. Reumaux	1990	France	AC 23: f1436 (2015)	ADC private	KX964448
	XC 2012-171	confirmatus ‘asp. imbutus’	A. Lantz	2012	France	AC 23: f1436 (2015)	ADC private	KX964449
	AB 09-11-514	confirmatus ‘asp. assiduus’	A. Bidaud	2009	France	AC 23: f1435 (2015)	ADC private	KX964450
	AB 05-11-423	confirmatus ‘asp. assiduus’	A. & E. Bidaud	2005	France	AC 23: f1435 (2015)	ADC private	KX964451
	AB 02-11-201	confirmatus ‘asp. assiduus’	F. Lopez	2002	France	AC 23: f1435 (2015)	ADC private	KX964452
	XC 2013-156	confirmatus ‘asp. assiduus’	F. Valade	2013	France	AC 23: f1435 (2015)	ADC private	KX964453
	AB 03-11-78	confirmatus ‘asp. confirmatus’	A. Faurite	2003	France	AC 23: f1434 (2015)	ADC private	KX964454
	AB 92-11-422	cistoadelphus ad int.	A. Bidaud	1992	France	FAMM 6: 41 (1994)	ADC private	KX964455
	AB 09-11-450	cohabitans cf.	A. Bidaud	2009	France	this study	ADC private	KX964456
	FR2016052	assiduus	J.-M. Ourcival	2016	France	this study	CEFE private	KX964457
	FR2012405	assiduus	P.-A. Moreau	2012	France	this study	CEFE private	KX964458
	FR2012089	assiduus	F. Richard	2011	France	this study	CEFE private	KX964459
	FR2012076	assiduus	E. Taschen	2011	France	this study	CEFE private	KX964460
	XC 2006-204	bresadolae cf.	na	2006	France	this study	ADC private	KX964461
	XC 2005-249	saturninus cf.	X. Carteret	2005	France	this study	ADC private	KX964462
	SH094374.07FU (6 sequences)	sp.	na	na	U/Iran	na	na	HQ204652
C. cyprinus	XC 2012-26	cyprinus (holotype)	G. Redeuilh	1993	France	AC 23: f1443 (2015)	PC	KX964463
	AB 11-11-251	cyprinus	A. Bidaud	2011	France	AC 23: f1443 (2015)	ADC private	KX964464
	AB 11-10-192	cyprinus	A. Bidaud	2011	France	AC 23: f1443 (2015)	ADC private	KX964465
	AB 06-09-144	cyprinus	A. Bidaud	2006	France	AC 23: f1443 (2015)	ADC private	KX964466
	PML 344	cyprinus	P. Moënne-Loccoz	1986	France	AC 23: f1443 (2015)	ADC private	KX964467
	PML 81	cyprinus	P. Moënne-Loccoz	1981	France	AC 23: f1443 (2015)	ADC private	KX964468
	XC 2013-15	cyprinus	P. Reumaux	2013	France	AC 23: f1443 (2015)	ADC private	KX964469
	XC 2007-103	cyprinus	X. Carteret	2007	France	AC 23: f1443 (2015)	ADC private	KX964470
	AB 04-09-167	sciophyllus cf.	A. Bidaud	2004	France	this study	ADC private	KX964471
	JMB 2014111802	circumvelatus cf.	P.-A. Moreau	2014	France	this study	CEFE private	KX964472
	PAM 13092901	circumvelatus	P.-A. Moreau	2013	France	this study	CEFE private	KX964473
	PML 425	myrtillinus	P. Moënne-Loccoz	1986	France	this study	ADC private	KX964474
	XC 2007-95	mutabilis cf.	na	2007	France	this study	ADC private	KX964475
	TEB 348-10	saturninus aff.	T.E. Brandrud	na	Norway	this study	na	KX964476
	TAAM 128765	sp.	A. Kollom	2008	Estonia	na	na	UDB016164
C. imbutus	IK 97-1162	imbutus (neotype)	I. Kytövuori	1997	Finland	this study	H	KX964498
	PML 4557	laccatus (holotype)	P. Reumaux	1978	France	SMF 98(4): 348 (1982)	PC	KX964478
	RH 3123	betulaecomes (holotype)	R. Henry	1976	France	SMF 93(3): 347 (1977)	PC	KX964479
	XC 2013-13	imbutus ‘asp. laetior’	P. Reumaux	1998	France	AC 23: f1447 (2015)	ADC private	KX964480
	XC 2014-77	imbutus ‘asp. saturnalis’	P. Reumaux	1978	France	AC 23: f1446 (2015)	ADC private	KX964481
	XC 2014-61	imbutus ‘asp. saturnalis’	P. Reumaux	1986	France	AC 23: f1446 (2015)	ADC private	KX964482
	XC 2007-104	imbutus ‘asp. vilior’	X. Carteret	2007	France	AC 23: f1445 (2015)	ADC private	KX964483
	AB 10-10-237	imbutus ‘asp. imbutus’	A. Bidaud	2010	France	AC 23: f1444 (2015)	ADC private	KX964484
	AB 09-11-471	imbutus ‘asp. imbutus’	A. Bidaud & R. Fillion	2009	France	AC 23: f1444 (2015)	ADC private	KX964485
	AB 04-09-228	imbutus ‘asp. imbutus’	A. Bidaud & A. Faurite	2004	France	AC 23: f1444 (2015)	ADC private	KX964486
	AB 98-10-358	imbutus ‘asp. imbutus’	A. Bidaud	1998	France	AC 23: f1444 (2015)	ADC private	KX964487
	PML 375	imbutus ‘asp. imbutus’	P. Reumaux	1986	France	AC 23: f1444 (2015)	ADC private	KX964488
	XC 2002-122	imbutus ‘asp. imbutus’	X. Carteret	2002	France	AC 23: f1444 (2015)	ADC private	KX964489
	XC 2002-108	imbutus ‘asp. imbutus’	X. Carteret	2002	France	AC 23: f1444 (2015)	ADC private	KX964490
	XC 2002-107	imbutus ‘asp. imbutus’	X. Carteret	2002	France	AC 23: f1444 (2015)	ADC private	KX964491
	XC 2002-106	imbutus ‘asp. imbutus’	X. Carteret	2002	France	AC 23: f1444 (2015)	ADC private	KX964492
	AB 08-10-307	cohabitans	J. Garin	2008	France	this study	ADC private	KX964493
	AB 02-10-106	cohabitans	M. Renard	2002	France	this study	ADC private	KX964494
	AB 02-09-58	cohabitans	A. Bidaud	2002	France	this study	ADC private	KX964495
	AB 00-09-127	cohabitans cf.	A. Bidaud	2000	France	this study	ADC private	KX964496
	IK 98-2242	sp.	I. Kytövuori	1998	Sweden	this study	H	KX964497
	IK 94-1236	sp.	I. Kytövuori	1994	Finland	this study	H	KX964477
	JMB 2008092703	salicis cf.	J.-M. Bellanger	2008	France	this study	CEFE private	KX964499
	RH 71030	betulaecomes	R. Henry	1976	France	this study (Rob. Henry, ined.)	PC	KX964500
	TN 11-257	sp.	T. Niskanen	2011	USA	this study	H	KX964501
	TN 11-252	sp.	T. Niskanen	2011	USA	this study	H	KX964502
	TN 11-151	sp.	T. Niskanen	2011	USA	this study	H	KX964503
	TN 11-150	sp.	T. Niskanen	2011	USA	this study	H	KX964504
	TN 05-167	sp.	T. Niskanen	2005	Finland	this study	H	KX964505
	XC 2012-96	laetior forma	X. Carteret	2012	France	this study	ADC private	KX964506
	XC 2002-109	renidentoides cf.	X. Carteret	2002	France	this study	ADC private	KX964507
	SH188563.07FU (6 sequences)	saturninus	na	na	Canada/Estonia/China	na	na	UDB018346
C. lucorum	CFP 490	lucorum (neotype)	T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos	1986	Norway	CFP: C10 (1994)	S	KX964585
	RH 71502	incarnatolilascens (holotype)	R. Henry	1979	France	AC 23: f1431 (2015), SMF 97(3): 170 (1981)	PC	KX964508
	PML 4142	montis-dei (holotype)	P. Reumaux	1980	France	AC 23: f1430 (2015), SMF 96(3): 357 (1980)	PC	KX964509
	PML 34	circumvelatus (holotype)	P. Reumaux	1976	France	AC 23: f1429 (2015), SMF 96(3): 355 (1980)	PC	KX964510
	10433	umidicola (syntype)	C.H. Kauffman	1903	USA	Bull. Torrey Bot. Club 32(6): 322 (1905)	MICH	KX964511
	PML 4143	lucorum ‘asp. montis-dei’	P. Reumaux	1980	France	AC 23: f1430 (2015)	ADC private	KX964512
	PAM 14090808	lucorum ‘asp. circumvelatus’	P.-A. Moreau	2014	France	AC 23: f1429 (2015)	ADC private	KX964513
	IK 89-748	lucorum	I. Kytövuori	1989	Finland	this study	H	KX964514
	KS CO513	diabolicus	K. Soop	na	Sweden	this study	na	KX964515
	TN 10-002	lucorum	T. Niskanen	2010	Canada	this study	H	KX964516
	TN 03-1169	lucorum	T. Niskanen	2003	Sweden	this study	H	KX964517
	SH188495.07FU (21 sequences)	lucorum	na	na	NA/FS	na	na	UDB019872
C. saturninus	CFP 514	saturninus (neotype)	T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos	1986	Sweden	CFP: C09 (1994)	S	KX964584
	PML 4578	urbicus var. sporanotandus (holotype)	A. Bidaud	1996	France	AC 23: f1455 (2015), AC 12: f560 (2002)	PC	KX964518
	PML 2215	marginatosplendens (isotype)	P. Reumaux	1978	France	AC 23: f1453 (2015), SMF 96(3): 356 (1980)	G	KX964519
	XC 2007-14	fulvorimosus (holotype)	A. & R. Bardet	1992	France	AC 23: f1452 (2015), AC 17: f869 (2008)	PC	KX964520
	RH 3758	denseconnatus (holotype)	na	1973	France	SMF 99(1): 65 (1983)	PC	KX964521
	RH 81181	gramineus (holotype)	R. Henry	1981	France	SMF 99(1): 64 (1983)	PC	KX964522
	RH 71682	rastetteri (holotype)	V. Rastetter	1980	France	SMF 97(3): 177 (1981)	PC	KX964523
	PR 258	dissidens (holotype)	P. Reumaux	1978	France	SMF 96(3): 370 (1980)	PC	KX964524
	RH 2623	salicis (holotype)	R. Henry	1968	France	SMF 93(3): 364 (1977)	PC	KX964525
	RH 476	umbrinoconnatus (holotype)	R. Henry	1955	France	SMF 73(1): 53 (1957)	PC	KX964526
	AB 02-10-179	saturninus ‘asp. urbicoides’	A. Faurite	2002	France	AC 23: f1457 (2015)	ADC private	KX964527
	AB 95-11-144	saturninus ‘asp. urbicoides’	R. Fillion	1995	France	AC 23: f1457 (2015)	ADC private	KX964528
	XC 2001-107	saturninus ‘asp. urbicoides’	X. Carteret & P. Reumaux	2001	France	AC 23: f1457 (2015)	ADC private	KX964529
	AB 14-11-160 (= AB 14-11-161)	saturninus ‘asp. salicis’	A. Bidaud, J. Cavet, R. Fillion & G. Raffini	2014	France	AC 23: f1454 (2015)	ADC private	KX964530
	XC 2014-109	saturninus ‘asp. salicis’	X. Carteret	2014	France	AC 23: f1454 (2015)	ADC private	KX964531
	XC 2011-205	saturninus ‘asp. salicis’	X. Carteret	2011	France	AC 23: f1454 (2015)	ADC private	KX964532
	XC 2008-55	saturninus ‘asp. salicis’	X. Carteret	2008	France	AC 23: f1454 (2015)	ADC private	KX964533
	XC 2007-108	saturninus ‘asp. salicis’	X. Carteret	2007	France	AC 23: f1454 (2015)	ADC private	KX964534
	AB 14-09-47	saturninus ‘asp. dionisiae’	E. Bidaud	2014	France	AC 23: f1451 (2015)	ADC private	KX964535
	AB 04-10-344	saturninus ‘asp. deceptivus’	A. Bidaud	2004	France	AC 23: f1450 (2015)	ADC private	KX964536
	AB 98-10-381	saturninus ‘asp. deceptivus’	Dr. Misermont	1998	France	AC 23: f1450 (2015)	ADC private	KX964537
	XC 2014-63	saturninus ‘asp. cohabitans’	M. Pèlerin	1996	France	AC 23: f1449 (2015)	ADC private	KX964538
	XC 2014-116	saturninus ‘asp. saturninus’	na	2014	France	AC 23: f1448 (2015)	ADC private	KX964539
	XC 2014-114	saturninus ‘asp. saturninus’	L. Tarahu	2014	France	AC 23: f1448 (2015)	ADC private	KX964540
	XC 2007-97	saturninus ‘asp. saturninus’	na	2007	France	AC 23: f1448 (2015)	ADC private	KX964541
	AB 97-09-187, PML 5347	urbicus	E. & A. Bidaud	1997	France	AC 12: f560 (2002)	ADC private	KX964542
	PML 3967	salicis var. salicis	M. Citérin	1994	France	AC 12: f559 (2002)	ADC private	KX964543
	CFP 408	subtorvus	T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos	1986	Sweden	CFP: A04 (1990)	S	KX964544
	AB 05-10-273	deceptivus sensu Moser	R. Fillion	2005	France	this study	ADC private	KX964545
	H:6029320	saturninus	I. Kytövuori	1998	Finland	this study	H	KX964546
	IK 94-631	saturninus	I. Kytövuori	1994	Finland	this study	H	KX964547
	JMB 2009101002	cohabitans	J.-M. Bellanger	2009	France	this study	CEFE private	KX964548
	KH14	subtorvus	na	2011	Norway (Svalbard)	na	na	GU234058
	O50591	subtorvus	na	2011	Norway (Svalbard)	na	na	GU234013
	PML 75	urbicus	P. Moënne-Loccoz	1984	France	this study	ADC private	KX964549
	TN 09-208	saturninus	T. Niskanen	2009	USA	this study	H	KX964550
	XC 2016-12	euprivignus aff.	P. Reumaux	1977	France	this study	ADC private	KX964551
	XC 2008-61	salicis	X. Carteret	2008	France	this study	ADC private	KX964552
	XC 2007-90	mutabilis cf.	X. Carteret	2007	France	this study	ADC private	KX964553
	XC 2006-194	salicis	X. Carteret	2006	France	this study	ADC private	KX964554
	XC 2002-167	holophaeus sensu Henry	M. Pèlerin	2002	France	this study	ADC private	KX964555
	XC 2001-104	mutabilis	X. Carteret	2001	France	this study	ADC private	KX964556
	XC 96-10-26-09	subprivignus	X. Carteret	1996	France	this study	ADC private	KX964557
	SH094324.07FU (13 sequences)	saturninus	na	na	USA/U	na	na	UDB017613
C. stuntzii	Rehner 394	stuntzii (holotype)	S.A. Rehner	1981	USA	Mycologia 80(6): 903 (1988)	WTU	KX964558
Other (morphological) Saturnini
C. cypriacoides	PML 1269	cypriacoides ‘asp. cypriacoides’ (holotype)	R. Fillion	1989	France	AC 23: f1423 (2015), AC 2: f81 (1990)	PC	KX964559
	PML 3984	cypriacoides ‘asp. cypriacoides’	C. Guyot	1989	France	AC 23: f1423 (2015)	ADC private	KX964560
	PML 3979	cypriacoides ‘asp. lucorum’	A. Bidaud	1992	France	AC 23: f1424 (2015), AC 9: f419 (1999)	ADC private	KX964561
C. furiosus	XC 2014-64c	furiosus (holotype)	D. Brion	2012	France	AC 23: f1458 (2015)	PC	KX964562
	LM5411	sp. (Quercus ectomycorrhiza)	na	na	Austria	na	na	KM576363
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	illepidus	A. Bidaud & C. Gérard	2011	France	AC 23: f1422 (2015)	ADC private	KX964564
C. nefastus	XC 2014-60	nefastus (holotype)	D. Brion	2012	France	AC 23: f1426 (2015)	PC	KX964565
C. ortovernus	JB 604808	ortovernus (holotype)	J. Ballará	2008	Spain	JEC 12(11): 56 (2009)	J. Ballara private	KX964566
C. oxytoneus	RH 931	oxytoneus (holotype)	R. Henry	1957	France	SMF 97(3): 277 (1981)	PC	KX964567
C. saturninoides sensu	AB 12-10-93	saturninoides	A. Bidaud & M. Renard	2012	France	AC 23: f1421 (2015)	ADC private	KX964568
ADC (= C. serratissimus)	AB 00-10-148	saturninoides	A. Bidaud	2000	France	AC 23: f1421 (2015)	ADC private	KX964569
	RH 3451	oxytoneus	R. Henry	1972	France	SMF 97(3): 277 (1981)	ADC private	KX964570
	XC 2014-119	saturninoides	R. Chalange	2014	France	AC 23: f1421 (2015)	ADC private	KX964571
	XC 2014-64b	saturninoides	D. Brion	2012	France	AC 23: f1421 (2015)	ADC private	KX964572
	XC 2013-144	saturninoides	F. Valade	2013	France	AC 23: f1421 (2015)	ADC private	KX964573
	XC 2010-56	saturninoides	X. Carteret	2010	France	AC 23: f1421 (2015)	ADC private	KX964574
	XC 2010-29	saturninoides	X. Carteret	2010	France	AC 23: f1421 (2015)	ADC private	KX964575
	SH188624.07FU (3 sequences)	lucorum	na	na	USA/Estonia/Italy	na	na	UDB016052
C. sciophylloides	AB 99-10-254	sciophylloides (holotype)	A. Bidaud	1999	France	AC 23: f1425 (2015)	PC	KX964576
	AB 91-10-291	sciophylloides	J. Garin	1991	France	AC 23: f1425 (2015)	ADC private	KX964577
	PML 5446	sciophylloides	J. Cavet	1999	France	AC 23: f1425 (2015)	ADC private	KX964578
	PML 2381	raphanodiabolicus	P. Reumaux	1991	France	na	ADC private	KX964579
	SH188568.07FU (6 sequences)	valgus	na	na	Canada/U	na	na	UDB002444
C. subfirmus	AB 08-10-363	subfirmus (holotype)	A. Bidaud & G. Raffini	2008	France	AC 23: f1433 (2015)	PC	KX964580
C. suboxytoneus	AB 01-09-56	suboxytoneus (holotype)	A. Bidaud	2001	France	AC 23: f1442 (2015)	PC	KX964581
	MFT60	sp. (Fagus ectomycorrhiza)	na	na	Germany	na	na	FJ403502
OtherTelamonia
C. alboviolaceus s.lat.	SH188487.07FU (26 sequences)	alboviolaceus	na	na	NA/U	na	na	AF325596
C. anisatus	CFP 1200	anisatus (holotype)	T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos	1993	Sweden	CFP: E25 (2014)	S	DQ117931
C. anisochrous	IK 01-030	anisochrous (holotype)	T. Niskanen & I. Kytövuori	2001	Estonia	Mycologia 105(4): 988 (2013)	H, S, NY	JX407297
C. athabascus	DBB27618, UC1860905	athabascus (holotype)	D. Bojantchev	2011	USA	Mycotaxon 123: 382 (2013)	UC	JN133295
C. biformis	SH188479.07FU (41 sequences)	biformis	na	na	NA/U	na	na	UDB002252
C. bovinus	IK 04-038	bovinus (neotype)	I. Kytövuori	2004	Finland	Mycologia 105(4): 981 (2013)	H, S, NY	JX407276
C. brunneifolius	TN 06-146	brunneifolius (holotype)	T. Niskanen	2006	Finland	Mycol. Progress 7(4): 241 (2008)	H	EU259284
C. caesioarmeniacus	H:7000901	caesioarmeniacus (holotype)	K. Liimatainen & T. Niskanen	2007	Canada	IF 198: 1 (2014)	H	KP137498
C. claroplaniusculus	RH 2334	claroplaniusculus (holotype)	R. Henry	1967	France	SMF 99(1): 65 (1983)	PC	KP013184
C. decipiens	PML 366	decipiens f. decipiens (neotype)	P. Moënne-Loccoz	1986	France	AC 11(1): f507 (2001), AC 2: f52 (1990)	G	FN428988
C. disjungendus	PAK 4370	disjungendus (lectotype)	P.A. Karsten	< 1893	Finland	ASFFF 9(1): 6 (1893)	H	KP013190
C. duracinus	PML 349	duracinus (neotype)	P. Moënne-Loccoz	1986	France	AC 2: f76 (1990)	G	KX964582
C. duracinus s.lat.	SH188648.07FU (2 sequences)	sp.	na	na	Denmark/Germany	na	na	AJ889943
C. duracinus s.lat.	SH094372.07FU (6 sequences)	rigens	na	na	NA/Italy	na	na	JF907880
C. fuscescens	H:6001898	fuscescens (holotype)	K. Liimatainen & T. Niskanen	2008	Finland	IF 201: 2 (2014)	H	KP165546
C. fuscobovinaster	IK 09-537	fuscobovinaster (holotype)	I. Kytövuori	2009	Norway	Mycologia 105(4): 990 (2013)	H, S, NY	JX407316
C. gallurae	CONS 00076	gallurae (holotype)	D. & M. Antonini, G. Consiglio	2002	Italy	Il genereCortinarius in Italia 3: C101 (2005)	CONS	FN428979
C. murinascens	IK 08-958	murinascens (holotype)	I. Kytövuori	2008	Finland	IF 201: 3 (2014)	H	KP165570
C. neofurvolaesus	CFP 1438	neofurvolaesus (holotype)	T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos	1999	Sweden	CFP: E24 (2014)	S	DQ139999
C. niveotraganus	SH188538.07FU (8 sequences)	niveotraganus	na	na	FS	na	na	KM273103
C. olididisjungendus	TN 07-191, H:7000854	olididisjungendus (holotype)	na	2007	Canada	IF 186: 2 (2014)	H	KM273091
C. orasericeus	RH 70239	orasericeus (holotype)	R. Henry	1975	France	SMF 99(1): 69 (1983)	PC	KP013203
C. quarciticus	CFP 765	quarciticus (holotype)	T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos	1988	Sweden	CFP: C59 (1994)	S	UDB000748
C. sordidemaculatus	RH 1122	sordidemaculatus (holotype)	R. Henry	< 1981	France	SMF 97(3): 196 (1981)	PC	DQ139984
C. sp.	IB 86/172	bovinus cf.	M. Moser	1986	Austria	na	IB	DQ139983
C. sp.	TF-01-034	aprinus	T. Frøslev	na	Denmark	na	C	AJ889942
C. subserratissimus	IK 11-017	subserratissimus (holotype)	I. Kytövuori	2011	Sweden	IF 201: 4 (2014)	H	KP165552
C. subturibulosus	SH188545.07FU (7 sequences)	subturibulosus	na	na	France/Spain/Portugal	na	na	FJ928484
C. tacitus	AB 05-09-72	tacitus (holotype)	A. Bidaud	2005	France	AC 22: f1400 (2014)	PC	KX964583
C. torvus	SH009362.07FU (10 sequences)	torvus	na	na	NA/FS/Germany	na	na	UDB001345
C. urbicus	SH188612.07FU (3 sequences)	urbicus	na	na	Canada/FS	na	na	UDB000743
Outgroup
C. anomalovelatus	JFA13109	anomalovelatus (holotype)	J.F. Ammirati	2007	USA	IF 93: 1 (2014)	WTU	FJ717605
C. lepidopus sensu auct.	SH196665.07FU (12 sequences)	anomalus	na	na	U	na	na	UDB002227
C. caesiocinctus	Sa57-13	caesiocinctus (holotype)	R. Kühner	1957	France	DM 20(77): 92 (1989)	G	DQ663239
C. flavipallens	IK 08-1729, H:6032393	flavipallens (holotype)	I. Kytövuori	2008	Finland	Persoonia 33 : 125 (2014)	H	KF732554
C. sannio	MM 97/352, IB:1997/0352	sannio (holotype)	M. Moser	1997	USA	Mycotaxon 72 : 315 (1999)	IB	KF732420

CFP, Cortinarius, Flora Photographica; AC, Atlas des Cortinaires; FN, Funga Nordica; DM, Documents Mycologiques; SMF, Bulletin de la Société Mycologique de France; FMDS, Bulletin de la Fédération Mycologique Dauphiné-Savoie; JEC, Journal des Journées Européennes du Cortinaire; K&R, Flore analytique des Champignons supérieurs (Küehner & Romagnesi); IF, Index Fungorum; ASFFF, Acta Societatis pro Fauna et Flora Fennica; BFNF, Bidrag till kännedom av Finlands Natur och Folk; NA, North America (USA, Canada); FS, Fennoscandia (Denmark, Sweden, Norway, Finland, Estonia, Lithuania, Latvia); U, Europe.

* Sequences generated for the present work are highlighted in bold.

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.

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).

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.

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 2011-205 (within the subclade) was fruiting from the same mycelium as XC 2007-108 and XC 2014-109 (outside the subclade).

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).

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.

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 transcontinental 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).

Table 3

Morphogenetic features of C. cinnamoviolaceus and species in sect. Bicolores and sect. Saturnini.

Species	Blue huesa	Odour(s)b	L min	Av L	L max	l min	Av l	l max	Av Q	Reported hostc,d	Dintra max / difference rate (incl. indels)a	Dinter min / difference rate (incl. indels)c
Cortinarius cinnamoviolaceus	+ or –	R, r, i	8.25	9.65	11.07	4.66	5.21	6.00	1.86	Picea, Abies, Pinus, Tilia, Quercus, Betula, 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 %
C. dolabratus	+ or –	ce, co	7.42	8.62	9.86	4.41	4.90	5.51	1.76	Pinus, Picea, Betula, Fagus, Quercus	3 nts / 0.5 %	3 nts (to C. dolabratoides) / 0.5 %
C. evernius	+	0, ce, r, R	8.75	10.34	11.85	5.35	6.01	6.77	1.72	Picea, Abies	1 nt / 0.2 % (1 nt + 1 indel / 0.3 %)	3 nts + 3 indels (to C. cagei) / 1 %
C. glaphurus	+ or –	ce, r, CE, V	8.03	9.32	10.60	4.82	5.23	5.78	1.78	Pinus, Quercus, Fagus, Abies, Picea, Populus, Betula	2 nts + 4 indels / 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 %
Sect. Saturnini
C. confirmatus	+ or –	0, ca, r, g	6.91	8.26	9.79	4.27	4.79	5.61	1.73	Quercus, Cistus, Pinus, Betula, Populus, Picea	6 nts + 1 indel / 1.2 %	3 nts (to C. imbutus) / 0.5 %
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, Populus, Carpinus, Picea, Abies	4 nts + 1 indel / 0.8 % (7 nts + 3 indels / 1.6 %)	3 nts + 2 indels (to C. cyprinus) / 0.8 %
C. stuntzii	(+)	0	9.60	11.50	14.40	5.90	6.70	8.50	1.72	Salix	na	3 nts + 5 indels (to C. saturninus) / 1.3 %

nt = nucleotide change; indel = insertion or deletion; na = not applicable (single sequence) or not available.

a Brackets mark uncertainty because of single collections (column ‘Blue hues’) or lack of available trace files for public sequences (column ‘Dintra max’).

b 0 = odourless; b = burnt keratin; ca = camphorated; ce = cedar wood; co = coconut; e = earth-like; g = grass-like; i = iodine; p = plum; r = radish. Upper/lower case relates to odour intensity. Bold indicates the most frequent odour.

c Bold indicates proven interaction (ectomycorrhizal sequences, column ‘Reported host’) or species with Dinter min > Dintra max (column Dinter min).

d Names are in the order of citation frequency.

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).

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.

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:

typical Bicolores concepts – C. parevernius and C. cinnamoviolaceus;

typical Duracini concepts – C. subparevernius, C. cylindratus and C. contractus;

a species defined by its author as intermediate between these two sections – C. basicyaneus;

a Duracini concept hiding a phylogenetic Bicolores – C. dolabratus; and

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 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).

(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.

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:

the CFP plate D48 is well-known and widely recognized as a good illustration of C. cagei;

our work considerably extends our morphogenetic, biogeographical and ecological knowledge of this species; and

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).

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, Varsinais-Suomi, Kisko, Kaukuri, mesic Picea forest, 16 Aug. 2000, T. Niskanen & I. Kytövuori, H:6033518; Etelä-Häme, Juupajoki, Hyytiälä, mesic Picea forest, 18 Aug. 2004, I. Kytövuori H:6033615*; Virrat, Monoskylä, Korpijärvi E, mesic Picea forest, 15 Oct. 2001, I. Kytövuori 01-062*, H; Pohjois-Häme, Laukaa, Äijälä, Heinäaho, mesic Picea forest, 10 Sept. 2004, I. Kytövuori 04-051*, H; Kainuu, Paltamo, Kontiomäki, Tololanmäki W, Kylmänpuro, W sloping, mesic Picea forest with some Pinus, Betula, Populus tremula and Salix, 14 Sept. 2008, I. Kytövuori 08-1771*, H:6033570; Koillismaa, Taivalkoski, Metsäkylä SW, Katajavaara, N sloping, old, mesic Picea forest with damp depressions, some Pinus, Betula and Populus tremula, 2 Sept. 2008, I. Kytövuori 08-788*, H:6033575. – France, Haute-Savoie, Tanninges, cespitose under Picea abies on a decalcified substrate, elev. 1500 m, 17 Aug. 2007, A. Bidaud & R. Fillion AB 07-08-48*, personal herbarium of A. Bidaud.

Notes — Morphologically, C. dolabratoides is reminiscent of its sister phylogenetic species C. dolabratus. Fortunately, the two species can be distinguished microscopically, C. 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).

Fr., Epicr. Syst. Mycol.: 311. 1838 — MycoBank MB#216747; Fig. 5a

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.

= 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 (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.

(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.

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.

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).

Rob. Henry, Bull. Soc. Mycol. France 93, 3: 362. 1977 — MycoBank MB#312090

– Cortinarius fundatus Britzelm., Ber. Naturhist. Vereins Augsburg 28: 127. 1885, sensu Bidaud et al. (2014).

Type. France, Vosges, Hennezel, in Abies forests, gregarious, 1972, R. Henry 3417, PC, holotype, MycoBank MBT#155523. ITS sequence deposited in GenBank under KX964374.

Illustrations — Bidaud et al. 2014: pl. 954 (as C. fundatus).

Taxonomic descriptions — Bidaud et al. 2014: f. 1411 (as C. 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.

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.

Illustrations — This study: Fig. 5c; Bidaud et al. 2014, pl. 952, 953 but also pl. 945 (as C. testaceoviolaceus).

Taxonomic descriptions — Bidaud et al. 2014: f. 1410 but also 2010: f. 1109 (as C. scriptor).

Notes — No original material was kept by Britzelmayr to 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).

(Fr.) Fr., Epicr. Syst. Mycol.: 305. 1838 — MycoBank MB#165676

Basionym. ≡ Agaricus tortuosus Fr., Syst. Mycol. 1: 235. 1821.

≡ Hydrocybe tortuosa (Fr.) Wünsche, Die Pilze. Eine Anleitung zur Kenntniss derselben: 121. 1877.

= Cortinarius flabelloides Carteret, Atlas des Cortinaires XIX: 1510. 2010.

= Cortinarius laetior P. Karst., Bidrag Kannedom Finlands Natur Folk 32: 387. 1879.

Type. Sweden, Smoland, Femsjö, Södra Färgen, Gatebäck, among Sphagnum in spruce forest, 11 Sept. 1979, D. Lamoure, IB 79/533, neotype designated in Opera Botanica 100: 182. 1989, MycoBank MBT#372784. ITS sequence deposited in GenBank under KX964391.

Illustrations — Bidaud et al. 2014: pl. 955–956 but also 2010: pl. 804 (as C. flabelloides); Brandrud et al. 1990: pl. A06.

Taxonomic descriptions — Bidaud et al. 2014: f. 1413 but also 2010: f. 1136 (as C. flabelloides); Niskanen et al. 2012: 863; Brandrud et al. 1990: pl. A06.

Notes — This Friesian name has been interpreted in rather similar ways by past and modern mycologists – with the 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).

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.

Illustration — Bidaud et al. 2008, pl. 672.

Taxonomic description — Bidaud et al. 2008: f. 885.

Notes — More collections of this species, originally described in sect. Damasceni by its authors, are required to better 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).

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 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.

(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, 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.

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. var. (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.

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:

electing these subclades at the species level would leave 8 basal sequences unresolved, in paraphyletic relationships with the 3 recognized species;

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

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:

absence of veil remnants on the stipe not fruiting densely cespitose; or

abundant veil remnants on the stipe densely cespitose under Populus alba (‘aspect’ paracohabitans); or

reddish hues on the cap densely cespitose under Betula pendula (‘aspect’ rubricosissimus).

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 Moënne-Loccoz & 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.

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.

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). Morphologically, 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:

the lilac-greyish, not violaceous, hues of young lamellae;

elongated spores (Av Q > 1.8, Table 3); and

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.

(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 macro-morphological 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 hygrophilous 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 Populus-associated Canadian collection TN10-002 along this part of the ribosomal locus (the basal-most and unsupported branch of the clade in Fig. 3 is artefactual and likely results from the shorter sequence of C. umidicola), one cannot preclude 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).

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).

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

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

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 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-imbutus-cypriacus, 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.