Abstract

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:

• 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

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 ​and3,3, respectively) focus on the species content of the revised sections and include, respectively, 124 and 131 sequences.

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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 ​and3,3, 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).

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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.

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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).

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).

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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 D_intra 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 D_inter 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 D_intra max for a given pair of sister species clades (Table 3). The topology of /Bicolores strongly supports two distinct lineages within the section, one including C. cagei, C. evernius, C. plumulosus, C. refectus, C. sp1 and C. sp2, and another one including C. dolabratoides, C. dolabratus, C. glaphurus, C. hircinosmus, C. tortuosus and C. turgidipes (Fig. 2).

As sampled here, /Saturnini includes 6 species in Europe and North America, each represented by 1 to 44 sequences (Fig. 3, Table 2). Sequencing existing type material revealed a much higher rate of synonymy when compared to species in /Bicolores, with 17 binomials identified as later names for C. confirmatus, C. imbutus, C. lucorum or C. saturninus. A comparatively wider species concept has emerged in this section, as illustrated by the case of C. saturninus, which merged not less than 9 holotypes previously reported to belong in unrelated sections. The considerable rise in species polymorphism resulting from such finding has been dealt with at the infraspecific taxonomic level in the last release of the ADC (Bidaud et al. 2015). In order to stabilize the nomenclature and fix the concept of species widely accepted as genuine members of the Saturnini section – or interpreted by some authors in sect. Bicolores, we designated neotypes for C. saturninus, C. imbutus and C. lucorum (see Taxonomy). Our work also positioned C. stuntzii and a morphogenetic, widened concept of C. confirmatus in the revised section, and it unravelled C. cyprinus as an overlooked species in sect. Saturnini (Fig. 3, Table 2, ​,33).

Intraspecific phylogenetic distances were considerably larger in /Saturnini when compared to /Bicolores, with a D_intra 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 (D_inter 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 D_intra 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.

Cortinarius cinnamoviolaceus M.M. Moser, Nova Hedwigia 14: 514. 1967 — MycoBank MB#329008

• = Cortinarius basicyaneus Rob. Henry & Trescol ex Bidaud & Eyssart., Bull. Semestriel Féd. Assoc. Mycol. Méditerranéennes 25: 38. 2004.

• = Cortinarius contractus Rob. Henry, Doc. Mycol. 16, 61: 27. 1985.

• = Cortinarius cylindratus Rob. Henry, Bull. Soc. Mycol. France 99: 91. 1983.

• = Cortinarius subparevernius Rob. Henry, Bull. Soc. Mycol. France 85: 442. 1970.

• [= Cortinarius parevernius Rob. Henry, Fl. Anal. Champ. Sup.: 303. 1953, nom. inval. (no diagnosis, no type designated)].

Type. Austria, Tirol, near Hötting, in mixed forest, 18 Sept. 1948, M. Moser, IB 48/590, holotype. MycoBank MBT#372783. ITS (partial) sequence deposited in GenBank under KX964412.

Misapplied names

• – Cortinarius dolabratus Fr., Epicr. Syst. Mycol.: 311. 1838, sensu Bidaud et al. (2008).

• – Cortinarius imbutus Fr., Epicr. Syst. Mycol.: 306. 1838, sensu Brandrud et al. (1998).

• – Cortinarius evernius Fr., Epicr. Syst. Mycol.: 294. 1838, sensu auct.

Illustrations — Bidaud et al. 2008: pl. 639 (as C. dolabratus); Brandrud et al. 1998: pl. D60 (as C. imbutus).

Taxonomic descriptions — Bidaud et al. 2008: f. 817 (as C. dolabratus); Brandrud et al. 1998: pl. D60 (as C. imbutus).

Notes — This is C. evernius sensu Konrad & Maublanc (1930) and sensu Henry (1937), with smaller spores and raphanoid smell. Our phylogenetic analysis reveals a much wider range of chromatic variability for this species, making it compatible with both sect. Bicolores and Duracini. In addition, the /C. cinnamoviolaceus clade here delineated sheds new lights on the intricate links between these two sections and sect. Saturnini (Fig. 1). Indeed, as redefined here, the species falls outside the three sections but it merges:

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

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

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

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

5. 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).

Cortinarius sect. Bicolores (M.M. Moser) Melot, Doc. Mycol. 20, 77: 97. 1989, emend.

Type. Cortinarius cagei Melot, Doc. Mycol. 20, 80: 58. 1990.

Notes — As phylogenetically revised here, Cortinarius sect. Bicolores has been redefined to a rather severe extent, with well-known representative species excluded from the revised section and half of its new content previously described outside Bicolores. The original diagnosis of the section should be emended as follow: young basidiomata usually (but not always) with violet tinges outside and/or in the context. Pileus strongly hygrophanous, yellowish brown, chocolate brown to reddish brown. Stipe cylindrical, often attenuate to rooting, usually with remnants of the white universal veil. Smell indistinct, weakly raphanoid, of cedar-wood, rarely of geosmin (earth-like, dusty). Spores amygdaloid to ellipsoid, sometimes fusiform, (6.5–)7–12(13) × (4–)4.3–7(–7.2) μm (on average: 9.3 × 5.4 μm), verrucose. Widely distributed in the Northern Hemisphere, fruiting solitary or gregarious, rarely cespitose, mostly under coniferous trees.

In its current sampling, it includes 12 species, 10 of which have been or can be assigned a Latin binomial.

Cortinarius cagei Melot, Doc. Mycol. 20, 80: 58. 1990 — MycoBank MB#129526

• ≡ Cortinarius bicolor Cooke, Grevillea XVI: 45. 1873, nom. illeg.

• – Cortinarius minicolor Rob. Henry, Bull. Soc. Mycol. France 104, 4: 300. 1989 ‘1988’, sensu Bidaud et al. (2014).

• [= Cortinarius periodolens Carteret & Reumaux ad int., Atlas des Cortinaires XXII: f. 1417. 2014, nom. inval. (no diagnosis, no type designated)].

Type. Sweden, Gotland, Lummelunda, Prästänget, under broadleaf trees, 1 Oct. 1994, T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos CFP1260, S, neotype designated here. MycoBank MBT#373139. ITS sequence deposited in GenBank under KX964295.

Illustrations — Bidaud et al. 2014: pl. 959 (as C. minicolor and C. periodolens); Brandrud et al. 1998: pl. D48.

Taxonomic descriptions — Bidaud et al. 2014: f. 1419 (as C. minicolor) but also f. 1417 (as C. periodolens); Niskanen et al. 2012: 864; Brandrud et al. 1998: pl. D48.

Notes — Historically, C. cagei was introduced to fix the nomenclatural issue associated with C. bicolor Cooke, an illegitimate name because of an earlier use of the name for another, unrelated taxon. However, by omitting to designate a holotype or other voucher specimen for his new name, Melot did not clarify the taxonomic ambiguity of C. bicolor. Indeed, C. bicolor was initially described as a species with medium-sized spores (10 × 5–6 μm) fruiting under deciduous trees. However, five years later, it was attributed much larger spores (12–14 × 6–7 μm), and also a broader ecology – mixed woods. It is likely that Cooke actually lumped together two phylogenetically distinct, but morphologically very similar species, in his latest diagnosis, making C. bicolor a nomen dubium. As such, the name may just be discarded but the authors of the CFP proposed an interpretation of C. cagei that fits very well the initial concept of C. bicolor. Because:

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

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

3. there is so far no convincing candidate for the second C. bicolor, even though C. plumulosus has been postulated to represent that one by the authors of the ADC (cf. notes under C. plumulosus), we fix here the species in its primary concept through designating the sequenced CFP1260 collection of plate D48 to neotypify C. cagei.

In these new morphogenetic boundaries, C. cagei is described in the ADC under C. minicolor, an obvious lookalike that, however, fruits under coniferous trees. Unfortunately, the holotype of C. minicolor could not be located in PC, preventing phylogenetic placement of the species within /Bicolores. Cortinarius cagei also includes C. periodolens, a Bicolores species described ad interim in the ADC, as a C. obtusus with violaceous stipe and strong iodine smell. Phylogenetically, C. cagei is well resolved due to the absence of any intraspecific sequence polymorphism and of a minimal distance to its sister species C. evernius of 3 substitutions plus 3 indels (Table 3). In the field, confusions are possible with C. refectus and C. plumulosus but spore shape ratios and host trees of the three species should prevent misidentification (Table 3).

Cortinarius dolabratoides Kytöv., Carteret, Bidaud, Liimat., Niskanen, Bellanger, Dima, Reumaux & Ammirati, sp. nov. — MycoBank MB#818596; Fig. 4

Etymology. The name refers to the close phylogenetic and morphological affinities with C. dolabratus.

Type. Finland, Koillismaa, Taivalkoski, Loukusa, the nature reserve of Loukusanharju, dry Pinus forest on the esker, with some Picea and Betula, some Picea-dominated depressions, 30 Aug. 2008, I. Kytövuori 08-465, H:6033567 (holotype H; isotype K). ITS sequence deposited in GenBank under KX964302.

Pileus 2–7 cm, conical when young, later expanding to plain with a distinct button-like umbo, clay brown to purplish brown, hygrophanous. Lamellae moderately distant, strongly emarginate, at first bluish then brown. Stipe 5–12 cm cylindrical to weakly clavate, sometimes slightly routing, white, with pale lavender blue top. Veil white, as a thin coating or obscure bands or patches on the stipe. Context whitish to purplish in the pileus, watery whitish bluish in the stipe. Exsiccated pileus dark blackish greyish brown, stipe much paler. Smell weakly grass-like or stronger, of cedar wood. Macrochemistry (on the context of the French collection only): Gaïac: ++; phénolaniline: +++; FMP: +++; AgNO3: 0. Spores 7–8.3–9.5 × 3.5–4.6–5.0 μm, Q = 1.68–1.82–1.96, (250 spores, 7 specimens), narrowly fusoid (to almost cylindrical), with a low suprahilar depression, often somewhat elongated at apex, fairly finely, densely verrucose, often prominently more strongly at the very apex, somewhat dark-coloured, faintly dextrinoid. Lamellar trama hyphae pale olive brownish, smooth to very finely densely scabrous. Basidia distinctly darker, olive brown (in MLZ). In damp to dryish boreal or alpine Picea abies forests, sometimes in dry Pinus sylvestris-dominated forests mixed by Picea abies.

Distribution — Fairly poorly known, but considered occasional.

Other specimens examined (sequenced collections marked with an asterisk, see Table 2 for GenBank accession numbers). Finland, 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).

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

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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).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.

Cortinarius evernius (Fr.) Fr., Epicr. Syst. Mycol.: 294. 1838 — MycoBank MB#233378

• Basionym. ≡ Agaricus evernius Fr., Observ. Mycol. 2: 79. 1818: sanctioned in Fr., Syst. Mycol. 1: 212. 1821.

• ≡ Hydrocybe evernia (Fr.) M.M. Moser, Kleine Kryptogamenflora von Mitteleuropa II: 161. 1953.

• ≡ Telamonia evernia (Fr.) Ricken, Die Blätterpilze. 1915.

• = Cortinarius evernius f. fragrans M.M. Moser ex Bidaud & Carteret, Atlas des Cortinaires XXII: 1887. 2014.

• = Cortinarius evernius f. pseudoscutulatus Rob. Henry ex Bidaud & Reumaux, Atlas des Cortinaires XXII: 1887. 2014.

• [= Cortinarius evernius var. insignis Fr., Atlas des Cortinaires XXII: f. 1405. 2014, nom. inval. (no diagnosis, no type designated)].

• [= Cortinarius parvulior Bidaud ad int., Atlas des Cortinaires XXII: f. 1418. 2014, nom. inval. (no diagnosis, no type designated)].

Type. Sweden, Ångermanland, Specksta, Härnösand, under coniferous trees, 22 Sept. 1988, T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos CFP792, S, neotype designated in Cortinarius Flora Photographica I (French version), pl. A11 (1990), MycoBank MBT#372785. ITS sequence deposited in GenBank under KX964331.

Illustrations — Bidaud et al. 2014: pl. 946–949 but also pl. 959 (as C. parvulior); Brandrud et al. 1990: pl. A11.

Taxonomic descriptions — Bidaud et al. 2014: f. 1404–1407 but also f. 1418 (as C. parvulior); Niskanen et al. 2012: 863; Brandrud et al. 1990: pl. A11.

Notes — All contemporary authors seem to interpret this widespread Friesian species the same way and, not considering infraspecific taxa and species described ad interim, no later synonym of C. evernius has been introduced – however, older authors like Konrad and Henry misapplied the name to C. cinnamoviolaceus (see above). Phylogenetically, the species displays very low intraspecific polymorphism despite its wide biogeographical distribution (1 substitution plus one length polymorphism out of 22 available sequences) and is separated from its sister species C. cagei by 3 substitutions plus 3 length polymorphisms (Table 3). In Europe, the species may be confused only with C. cinnamoviolaceus, but the latter strongly smells of radish, has smaller spores and displays a much broader ecological range.

Cortinarius glaphurus Chevassut & Rob. Henry, Doc. Mycol. 12, 47: 78. 1982 — MycoBank MB#109708

• = Cortinarius tubulosus Bidaud, Atlas des Cortinaires XXII: 1888. 2014.

• = Cortinarius cedriosmus Bidaud, Atlas des Cortinaires XIX: 1510. 2010.

• = Cortinarius violaeolens Carteret & Reumaux, Atlas des Cortinaires XIX: 1509. 2010.

• = Cortinarius paranomalus Rob. Henry, Atlas des Cortinaires IV: 105. 1992.

• – Cortinarius turibulosus (Jul. Schäff. & E. Horak) Bon & G. Garnier, Doc. Mycol. 21, 83: 10. 1991, sensu auct.

Type. France, Hérault, La Salvetat-sur-Agout, Lac de la Raviège, under Picea, cespitous, 29 Oct. 1978, R. Henry 71421, PC, holotype, MycoBank MBT#70172. ITS sequence deposited in GenBank under KX964352.

Illustrations — Bidaud et al. 2014: pl. 957 (as C. tubulosus); 2010: pl. 795 (as C. turibulosus), pl. 796 (as C. violaeolens) and pl. 807 (as C. cedriosmus); 1992: pl. 83 (as C. paranomalus).

Taxonomic descriptions — Bidaud et al. 2014: f. 1414 (as C. tubulosus) and 2010: f. 1108 (as C. turibulosus); Kühner & Romagnesi 1953: 305 (as C. paranomalus); Chevassut & Henry 1982: 78.

Notes — As redefined here, the concept of C. glaphurus should be substantially widen so as to include those of C. cedriosmus, C. paranomalus, C. tubulosus and C. violaeolens, as well as C. turibulosus sensu Bidaud et al. (2010). The protologue should then be edited as follows: pileus diameter up to 55 mm, pileus dark chocolate-brown to reddish brown, not glabrous and hygrophanous. Stipe not always straight nor isodiametric but often (always?) hollow, with or without blue pigments and with variable amounts of veil remnants that may form a membranous ring. Often cespitose. Odour weakly raphanoid or of cedar wood or viola. Associated with coniferous trees as well as broad-leaved trees (Pinus, Quercus and Fagus confirmed as hosts by ectomycorrhizal sequences). Phylogenetically, the species is a bit polymorphic but is still well separated from its sister species C. tortuosus (Table 3). When collected under Picea abies on calcareous soils and weakly smelling of cedar wood, C. glaphurus may be difficult to distinguish from C. hircinosmus, but the latter produces slightly smaller spores (Table 3). When collected in hygrophilic and acidic soils under coniferous trees, the species may be confused with C. tortuosus, but the latter displays obvious blue tinges on the stipe, blood-red hues in the gills, and is never cespitose.

Cortinarius hircinosmus Moënne-Locc., Atlas des Cortinaires XII: 692. 2002 — MycoBank MB#489854

• – Cortinarius livor Fr., Epicr. Syst. Mycol.: 306. 1838, sensu Bidaud et al. (2015).

• – Cortinarius scriptor Kühner, Doc. Mycol. 20, 77: 92. 1989, sensu Bidaud et al. (2010) p.p.

Type. France, Haute-Savoie, Les Puisots, in Picea forest, elev. 700 m, 15 Sept. 1986, P. Moënne-Loccoz 334, PC, holotype, MycoBank MBT#101337. ITS sequence deposited in GenBank under KX964368.

Illustrations — Bidaud et al. 2015: pl. 991 (as C. livor); 2002: pl. 389.

Taxonomic descriptions — Bidaud et al. 2015: f. 1459 (as C. livor); 2002: f. 575; Niskanen et al. 2012: 850.

Notes — This species has been initially described in subsect. Hircini because of the strong smell of C. hircinus and C. camphoratus of the holotype specimens. However, five additional collections from France and Scandinavia, lacking such odour, were later identified in the same clade. As revised here and at least in France, C. hircinosmus fruits under Picea abies on calcareous soils and includes the French concept of C. livor and pro parte, that of C. scriptor. The original binomial is obviously unfortunate for an odourless or weakly smelling species, so, provided additional collections confirm the strong smell of some populations, infraspecific taxa may be introduced to more adequately reflect the organoleptic diversity of the species. Phylogenetically, the species is well resolved (Table 3). In the field, C. hircinosmus may be confused with C. glaphurus (as redefined here), but the latter displays a much broader ecological niche, typically smells of cedar wood and has slightly larger spores (9.3 × 5.2 μm vs 9 × 5 μm, on average).

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

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

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

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

Taxonomic descriptions — Bidaud et al. 2014: f. 1411 (as C. fundatus); Henry 1977: 359.

Notes — This conifer-associated species has been treated in the ADC as C. fundatus, and suspected by French authors, on the basis of frequent macrospores up to 12 μm long observed in some collections, to represent the second C. bicolor of Cooke – the one with large spores and possible fruiting under coniferous trees (cf. notes under C. cagei). Phylogenetically, C. plumulosus is well separated from its closest neighbour C. evernius (7 substitutions plus 3 indels, Table 3). Morphologically, the species resembles C. refectus and C. cagei but the former produces ovoid spores (Av Q = 1.5), the latter fruits under deciduous trees and the cap of C. plumulosus is typically covered by small flakes that are not found on that of its two lookalikes.

Cortinarius refectus Britzelm., Ber. Naturhist. Vereins Augsburg 28: 127. 1885 — MycoBank MB#560269; Fig. 5b–c

• ≡ Cortinarius reflectus Britzelm., Ber. Naturhist. Vereins Augsburg 28: 127. 1885.

• – Cortinarius scriptor Kühner, Doc. Mycol. 20, 77: 92. 1989, sensu Bidaud et al. (2010) p.p.

Misapplied name

• – Cortinarius testaceoviolaceus Rob. Henry, Bull. Soc. Mycol. France 73, 1: 51. 1957, sensu Bidaud et al. (2014).

Type. Atl. Tab. 377, f. 202 (1890), lectotype (iconotype) designated here (Fig. 5b), MycoBank MBT#373158. Germany, Lombach, in Picea and Abies forest, on calcareous soil, elev. 600 m, 24 Sept. 1996, A. Bidaud 96-09-73, epitype designated here (Fig. 5c), MycoBank MBT#373159. ITS sequence deposited in GenBank under KX964385.

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).

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

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

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

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

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

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

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

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

Notes — This Friesian name has been interpreted in rather similar ways by past and modern mycologists – with the notable exception of J. Favre, who referred to this species as C. plumbosus – so that C. tortuosus taxonomy is not a problematic issue. The species can be diagnosed by its narrow ecological niche (hygrophilous and acidic soils, with conifer trees) and the special purple-red tinges of the gills that tend to darken upon bruising. The odour is usually reported as null or weak of cedar wood but the conspecificity with C. flabelloides, revealed in this work, indicates that basidiomata can also smell of geosmin (i.e., of earth or dust, as C. variecolor for instance). Phylogenetically, the species is remarkably stable at the ITS locus and is well separated from its sister species C. glaphurus (Table 3).

Cortinarius turgidipes Rob. Henry ex Rob. Henry, Atlas des Cortinaires XVII, 1: 1179. 2008 — MycoBank MB#533088

Type. France, Creuse, Lavaud, under Picea, on granitic soil, subcespitose, 19 Oct. 1993, A. & E. Bidaud, AB 93-10-425, PC, holotype, MycoBank MBT#372786. ITS sequence deposited in GenBank under KX964409.

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).

Cortinarius sect. Saturnini Rob. Henry ex Möenne-Locc. & Reumaux, Atlas des Cortinaires I: 21 (1990), emend.

Type. Cortinarius saturninus (Fr.) Fr., Epicr. Syst. Mycol.: 306. 1838.

• ≡ Cortinarius subsect. Saturnini Bidaud, Moënne-Locc. & Reumaux, Doc. Mycol. 24, 95: 41. 1994.

• ≡ Cortinarius sect. Firmiores (Fr.) Henn., in Engler & Prantl, Naturl. Pfanzenf. I, 181: 246. 1900, p.p.

Notes — As revised here, sect. Saturnini is widely distributed in the Northern Hemisphere and includes 6 species. They are medium-sized, rarely stout Telamonia species, pale ochraceous, brown to reddish brown, lilac-violet, hygrophanous, with or without blue tinges in young lamellae and the upper part of the stipe, with various amounts of veil remnants on the stipe and on the pileus margin where it often forms a continuous covering or discontinuous patches. Smell indistinct or weak. Spores broadly or narrowly ellipsoid, (6–)6.5–11(–14.4) × (3–)4–7(–8.5) μm (on average: 8.6 × 4.9 μm), verrucose. Gregarious to densely cespitose, rarely solitary, typically fruiting under hygrophilous deciduous trees (Salix, Populus, Betula) but also under Quercus and Cistus in the Mediterranean area, rarely under coniferous trees.

Cortinarius saturninus (Fr.) Fr., Epicr. Syst. Mycol.: 306. 1838 — MycoBank MB#177635

• Basionym. ≡ Agaricus saturninus Fr., Syst. Mycol. 1: 219. 1821.

• = Cortinarius fulvorimosus Carteret & Reumaux, Atlas des Cortinaires XVII, 1: 1178. 2008.

• = Cortinarius cohabitans var. urbicoides Bidaud & Fillion, Bull. Soc. Mycol. France 119, 1–2: 70. 2004.

• = Cortinarius urbicus var. sporanotandus Bidaud & Fillion, Atlas des Cortinaires XII: 695. 2002.

• = Cortinarius denseconnatus Rob. Henry, Bull. Soc. Mycol. France 99, 1: 65. 1983.

• = Cortinarius gramineus Rob. Henry, Bull. Soc. Mycol. France 99, 1: 64. 1983.

• = Cortinarius rastetteri Rob. Henry, Bull. Soc. Mycol. France 97, 3: 177. 1981.

• = Cortinarius dissidens Reumaux, Bull. Soc. Mycol. France 96, 3: 356. 1980.

• = Cortinarius marginatosplendens Reumaux, Bull. Soc. Mycol. France 96, 3: 356. 1980.

• = Cortinarius salicis Rob. Henry, Bull. Soc. Mycol. France 93, 3: 364. 1977.

• = Cortinarius umbrinoconnatus Rob. Henry, Bull. Soc. Mycol. France 73, 1: 53. 1957.

• [= Cortinarius dionisiae Bidaud ad int., Atlas des Cortinaires XXIII: f. 1451. 2015, nom. inval. (no diagnosis, no type designated)].

• – Cortinarius subtorvus Lamoure, Schweiz. Z. Pilzk. 47, 9: 169. 1969, sensu auct.

• – Cortinarius bresadolae Schulzer, Hedwigia 24, 4: 138. 1885, sensu Lamoure (1978).

• – Cortinarius cohabitans P. Karst., Bidrag Kannedom Finlands Natur Folk 32: 388. 1879, sensu auct.

• – Cortinarius urbicus (Fr.) Fr., Epicr. Syst. Mycol.: 293. 1838, sensu Bidaud et al. (2002) p.p.

Type. Sweden, Västergötland, Eggby, Drottningkullen, deciduous forest on calcareous ground (Corylus, Tilia, Quercus), 17 Sept. 1986, T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos CFP514, S, neotype designated here, MBT#373160. ITS sequence deposited in GenBank under KX964584.

Illustrations — Bidaud et al. 2015: pl. 983–989; Brandrud et al. 1994: pl. C09, but also 1990: pl. A04 (as C. subtorvus).

Taxonomic descriptions — Bidaud et al. 2015: f. 1448–1457; Niskanen et al. 2012: 847–848; Brandrud et al. 1994: pl. C09, but also 1990: pl. A04 (as C. subtorvus).

Notes — All contemporary and past authors agree on the fact that C. saturninus is a collective species, that Fries himself contributed to confuse through multiple diagnoses across his successive monographs, which, in addition, do not fit the plates he later directed. The French mycologist Robert Henry devoted decades of his life trying to sort out this complex, adding to the literature many new names and interpretations (for review, see Bidaud et al. 2015). The simplest way to clarify this issue would undoubtedly be to consider C. saturninus as a nomen dubium and readily discard it. However, the wide use of the name that pertained throughout modern literature and the general consensus about the species illustrated on the plate C09 of the CFP, prompted us to fix C. saturninus in its current, Nordic concept, through the neotypification of the name with the CFP514 collection. Our phylogenetic analysis reveals a tremendously polymorphic species, with no less than 9 holotypes previously thought to be unrelated to sect. Saturnini, falling as later synonyms of C. saturninus. Cortinarius subtorvus and C. cohabitans, usually considered as akin to C. saturninus, are most likely two additional synonyms, although their respective type material could not be sequenced to ascertain conspecificity. This work also establishes that C. oxytoneus, 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 (D_intra 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 D_intra max in C. saturninus is actually of 4 nt changes, a value that stems from three French collections (PML 75 in one hand and AB 04-10-344 and XC 2002-167 in the other) which may deserve taxonomic autonomy – at the infraspecific rank – when more thoroughly sampled (Table 3, Fig. 3). Although its suspected association with Salix is here demonstrated by the presence in the clade of several ectomycorrhizal sequences isolated from willow roots (within SH094324.07FU, Table 2, Fig. 3), C. saturninus may also be associated with other deciduous, but also coniferous trees. Morphologically, the species displays unprecedented levels of variability that represent a serious issue for field diagnosis. Practically, one should consider C. saturninus as a possible hit – and check the numerous aspects of this species in the last release of the ADC for instance (Bidaud et al. 2015) – whenever collecting a cespitose or gregarious medium-size Telamonia: i) under Salix spp. or other hygrophilous deciduous trees (and Dryas octopetala in the alpine zone), with or without blue hues at the stipe apex and with veil remnants ranging from none to white patches or covering at the cap margin, to copious and web-like covering the whole young fruit body; or ii) under coniferous trees and in this case with a ring and with short (L < 10 μm), ellipsoid spores. Highest risks of confusion are with other members of the revised sect. Saturnini (see notes under C. confirmatus, C. cyprinus and C. imbutus), and, for blue-lacking and densely veiled basidiomata collected under Salix spp. (referred to as C. saturninus ‘aspect’ salicis, ‘aspect’ urbicoides and ‘aspect’ sporanotandus in the ADC), with C. urbicus. The latter species displays more whitish hues on the fresh pileus and is typically less hygrophanous than C. saturninus, with no ‘Kuehneromyces-like’ dehydration.

Cortinarius confirmatus Rob. Henry, Bull. Soc. Mycol. France 99, 1: 67. 1983 — MycoBank MB#818598 (var. confirmatus); MycoBank MB#818597 (var. plesiocistus)

• = Cortinarius assiduus var. plesiocistus A. Ortega et al., Mycotaxon 101: 140. 2007.

• = Cortinarius assiduus Mahiques, A. Ortega & Bidaud, Bull. Féd. Mycol. Dauphiné-Savoie 162: 42. 2001.

• = Cortinarius bulbosovolvatus Rob. Henry & Contu, Doc. Mycol. XVI, 61: 32. 1985.

• [= Cortinarius kuehneri Bidaud ad int., Atlas des Cortinaires XXIII: f. 1440. 2015, nom. inval. (no diagnosis, no type designated)].

• [= Cortinarius spurcatocephalus Carteret ad int., Atlas des Cortinaires XXIII: f. 1439. 2015, nom. inval. (no diagnosis, no type designated)].

• [= Cortinarius paracohabitans Bidaud ad int., Atlas des Cortinaires XXIII: f. 1437. 2015, nom. inval. (no diagnosis, no type designated)].

• [= Cortinarius subcylindratus Carteret ad int., Bull. Soc. Mycol. France 128(3–4): 280. 2014, nom. inval. (no diagnosis, no type designated)].

• [= Cortinarius cistoadelphus Bidaud ad int., Bull. Féd. Assoc. Mycol. Méditerranéennes 6: 41 (1994), nom. inval. (no diagnosis, no type designated)].

• – Cortinarius cypriacus Fr., Epicr. Syst. Mycol.: 307. 1838, sensu Consiglio (1999) non Moënne-Loccoz & Reumaux (1989).

Type. France, unknown locality and collection date, under Quercus ilex, R. Henry 3195, PC, holotype, MycoBank MBT#69663. ITS sequence deposited in GenBank under KX964438.

New combination. Cortinarius confirmatus var. plesiocistus (A. Ortega, Vila & Bidaud) Carteret, Bidaud, Reumaux & Bellanger, comb. nov.

Basionym. Cortinarius assiduus var. plesiocistus A. Ortega, Vila & Bidaud in Ortega et al., Mycotaxon 101: 140. 2007. ITS sequence deposited in GenBank under AM713178.

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 (D_intra 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:

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

2. 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

3. 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:

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

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

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

Cortinarius cyprinus Bidaud, Carteret & Reumaux, Atlas des Cortinaires XXIII: 1981. 2015 — MycoBank MB#815172

• [= Cortinarius saturninus var. bresadolae M.M. Moser, Kleine Kryptogamenflora von Mitteleuropa II: 162. 1953, nom inval. (ined.)].

• – Cortinarius cypriacus Fr., Epicr. Syst. Mycol.: 307. 1838, sensu 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.

Cortinarius imbutus Fr., Epicr. Syst. Mycol.: 306. 1838 — MycoBank MB#233557; Fig. 5d

• = Cortinarius laccatus Reumaux, Bull. Soc. Mycol. France 98, 4: 348. 1982.

• = Cortinarius betulaecomes Rob. Henry, Bull. Soc. Mycol. France 93, 3: 347. 1977.

• [= Cortinarius saturnalis Reumaux ad int., Atlas des Cortinaires XXIII: f. 1446. 2015, nom. inval. (no diagnosis, no type designated)].

Type. Finland, Perä-Pohjanmaa, Tornio, Arpela, Runteli, rich grass-herb spruce forest with deciduous bushes and some pines, slightly paludified depressions, calcareous ground, 10 Sept. 1997, I. Kytövuori 97-1162, H, neotype designated here, MycoBank MBT#373161 (Fig. 5d). ITS sequence deposited in GenBank under KX964498.

Illustrations — This study: Fig. 5d; Bidaud et al. 2015: pl. 976–982.

Taxonomic descriptions — Bidaud et al. 2015: f. 1445–1447.

Notes — The two major contemporary interpretations of C. imbutus are in marked contrast, as the CFP authors consider the species in sect. Bicolores, while those of the ADC place it in sect. Saturnini. The Friesian diagnosis of C. imbutus is, as often with old names, not precise enough to support a single, unequivocal interpretation. However, Fries described his species between C. saturninus and C. cypriacus, indicating that the original concept would be naturally placed in sect. Saturnini. Our work reveals that the French version of C. imbutus is one of the morphogenetic Saturnini, widely distributed across the northern hemisphere, whereas the CFP one corresponds to a blue-pigmented collection of C. cinnamoviolaceus (and is then conspecific with the French C. dolabratus, see notes under this species). We thus here stabilize the name in the revised sect. Saturnini, by neotypifying it with the sequenced IK97-1162 collection from Finland. Phylogenetically, C. imbutus is rather polymorphic at the ITS locus (D_intra max = 3 substitutions + 1 indel) and simultaneously very close from its closest species C. confirmatus (D_inter 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:

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

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

3. copious veil remnants on the stipe.

Macrochemistry may be useful as well to distinguish C. imbutus from C. confirmatus (gaïacol and silver nitrate), although the reliability of these reactions is still questionable.

Cortinarius lucorum (Fr.) Berger, Cat. Herb. III: 89. 1846 — MycoBank MB#818604

• Basionym. ≡ Cortinarius impennis var. lucorum Fr., Epicr. Syst. Mycol.: 294. 1838.

• ≡ Hydrocybe lucorum (Fr.) M.M. Moser, Kleine Kryptogamenflora von Mitteleuropa II: 162. 1953.

• ≡ Cortinarius lucorum (Fr.) Mussat: 101. 1901.

• ≡ Cortinarius impennis subsp. lucorum (Fr.) Sacc.: 951. 1887.

• = Cortinarius incarnatolilascens Rob. Henry, Bull. Soc. Mycol. France 97, 3: 170. 1981.

• = Cortinarius montis-dei Reumaux, Bull. Soc. Mycol. France 96: 357. 1980.

• = Cortinarius circumvelatus Reumaux, Bull. Soc. Mycol. France 96: 355. 1980.

• ? = Cortinarius umidicola Kauffman, Bull. Torrey Bot. Club 32, 6: 322. 1905.

Type. Norway, Vestfold, Moss, Jelöy, under Populus tremula, 13 Sept. 1986, T.E. Brandrud, H. Lindström, H. Marklund, S. Muskos CFP490, S, neotype designated here, MycoBank MBT#373173. ITS sequence deposited in GenBank under KX964585.

Illustrations — Bidaud et al. 2015: pl. 967–969; Brandrud et al. 1994: pl. C10.

Taxonomic descriptions — Bidaud et al. 2015: f. 1428–1431; Niskanen et al. 2012: 847; Brandrud et al. 1994: pl. C10; Matheny & Ammirati 2006.

Notes — In Nordic countries, this widespread species is tightly associated with Populus spp. and it is well known, in large part thanks to the plate C10 published in the CFP. North American mycologists, following Kauffman’s footsteps, sometimes name this species C. umidicola, even though the latter binomial has been originally applied to a mushroom fruiting in conifer forests, e.g., Tsuga (Kauffman 1932). French authors described it repeatedly, as C. circumvelatus, C. incarnatolilascens and C. montis-dei, on the basis of deviating 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 D_inter min far exceeding D_intra 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).

Cortinarius stuntzii S.A. Rehner & Ammirati, Mycologia 80, 6: 903. 1988 — MycoBank MB#135248

Type. USA, Washington, Grant County, Crab Creek, 5 Nov. 1981, S.A. Rehner 394, WTU, holotype, MycoBank MBT#78780. ITS sequence deposited in GenBank under KX964558.

Illustration — Rehner et al. 1988: f. 1.

Taxonomic description — Rehner et al. 1988: 904–906.

Notes — This stout species densely fruiting under Salix exigua and S. rigida, so far known only from a small location of North-western USA, was compared to C. umidicola and C. subtorvus in the original publication, compatible with a placement into sect. Saturnini. However, a positioning elsewhere in subg. Telamonia, or even in subg. Sericeocybe – due to its low hygrophaneity – has also been invoked. The present work unambiguously establishes C. stuntzii as a genuine Saturnini, phylogenetically most closely related to C. saturninus, from which it differs by 3 substitutions and 5 indels (Table 3). Not considering biogeography, so far restricted to the type locality, the species is easily distinguished from other Saturnini members by its unusually large spores, up to 14.4 μm long and 8.5 μm wide (on average: 11.5 × 6.7 μm, Table 3).

KEY TO SPECIES TREATED IN THE PRESENT STUDY

• 1. Alpine and arctic zone, under Salix spp. or Dryas octopetala . . . . . . . . . . . . . . C. saturninus

• 1. Mediterranean thermophilic area, under Quercus ilex or Cistus spp. . . . . . . . . . . . . . . C. confirmatus

• 1. Continental zone . . . . . . . . . . . . . . 2

• 2. Coniferous trees . . . . . . . . . . . . . . 3

• 2. Deciduous trees . . . . . . . . . . . . . . 20

• 3. Acidic soils, in or near peatlands, Picea or Abies . . . . . . . . . . . . . . 4

• 3. Dry to mesic acidic woodlands . . . . . . . . . . . . . . 6

• 3. Basic to neutral, often calcareous woodlands . . . . . . . . . . . . . . 13

• 4. Average spore length > 10 μm, blue tinges obvious, usually odourless . . . . . . . . . . . . . . C. evernius

• 4. Average spore length < 10 μm, usually smelling . . . . . . . . . . . . . . 5

• 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.

Acknowledgments

REFERENCES