Type studies and fourteen new North American species of Cortinarius section Anomali reveal high continental species diversity

Section Anomali is a species-rich group in North America belonging to Cortinarius, the most diverse genus in the Agaricales. This study is based on extensive morphological investigations and molecular methods using 191 nrDNA ITS sequence data and recovered 43 phylogenetic species from which 14 are described here as new to science. We sequenced ten type materials which belonged to eight species. The synonymy of C. caesiellus with C. albidipes and C. copakensis with C. albocyaneus is proposed here. The North American occurrence of four species (C. albocyaneus, C. anomalus, C. caninus, and C. tabularis), so far known only from Europe, was confirmed. Thirteen species were not formally described here due to lack of relevant information. An identification key to the known Anomali species in North America is provided.


Introduction
Cortinarius is an important ectomycorrhizal genus in the Agaricales and is known to be the largest agaric genus worldwide (e.g., Garnica et al. 2016;Liimatainen et al. 2017Liimatainen et al. , 2020Varga et al. 2019). Species recognition based on morphology is difficult in Cortinarius lineages due to overlapping species characteristics and variation within species (e.g., Frøslev et al. 2007;Niskanen et al. 2009Niskanen et al. , 2013Niskanen et al. , 2016Liimatainen et al. 2014Liimatainen et al. , 2015. Section Anomali represents a monophyletic, globally distributed group of Cortinarius (e.g., Soop et al. 2019) characterized by a telamonioid/sericeocyboid appearance, often with yellowish to brownish universal veil remnants on the stipe, typically bluish young lamellae, and subgloboid to broadly ellipsoid or rarely ellipsoid, verrucose spores. In section Anomali, more than 50 binomials were introduced in the last century, mainly from Europe with fewer from North America and elsewhere, but only ca. 20% of these names have been in general use . Classical European species were examined and typified in Dima et al. (2016) prior to studying species from other parts of the world, such as North America, where knowledge of sect. Anomali has been limited. Some earlier workers (e.g., Peck 1873Peck , 1874Peck , 1878Peck , 1888Peck , 1912Murrill 1946;Kauffman 1905Kauffman , 1932Kauffman and Smith 1933;Smith 1939Smith , 1944, and a few recent ones (Ammirati 2014;Ammirati et al. 2017;Liimatainen and Niskanen 2021) described a dozen species from the North and Central America. However, to date, the classical European names such as C. anomalus, C. albocyaneus, C. tabularis, and others have been applied to most of the taxa in North Section editor: Zhu-Liang Yang 1 3 America (Harrower et al. 2011). This is a general problem based on recent phylogenetic revisions in Cortinarius (e.g., Liimatainen et al. 2014Liimatainen et al. , 2020Garnica et al. 2016;Soop et al. 2019) and similar to what has occurred in other groups of fungi (e.g., Kropp et al. 2010).
To extend the knowledge of the taxonomy and distribution of species in section Anomali in North America, we present here an extensive nuc rDNA ITS1-5.8S-ITS2 (ITS barcode) dataset as a phylogenetic and taxonomic framework, based on sequences from basidiomata, including type materials and environmental samples belonging to this group of Cortinarius. Our taxonomic revision is based on both phylogenetic data and detailed macro-and microscopical observations.

Molecular analysis
The universal fungal DNA barcode region (Schoch et al. 2012), the ribosomal internal transcribed spacer (ITS1-5.8S-ITS2) was amplified, sequenced and analyzed. A total of 81 ITS sequences belonging in sect. Anomali were generated in this study, of which 77 originated from specimens collected in North America (Table 1). For DNA extraction, PCR, and sequencing, we followed the protocols described in Liimatainen et al. (2014) and Dima et al. (2016). Primer pairs of ITS1F-ITS4 or ITS1F-ITS4B, and alternatively in case of old type materials ITS1F/ITS2 and ITS3/4 (Gardes and Bruns 1993;White et al. 1990), were used to amplify the whole or partial ITS region. Sequencing was performed at the University of Helsinki (Finland) or at LGC Genomics (Berlin, Germany). GenBank accession numbers of the novel sequences are listed in Table 1.

Phylogenetic analyses
Sequences were aligned in MAFFT v7 (http:// mafft. cbrc. jp/ align ment/ server) choosing the E-INS-I method (Katoh and Standley 2013) under default settings. SeaView 4 (Gouy et al. 2010) was used to inspect the alignment. Based on the work of Nagy et al. (2012), phylogenetically informative indels were coded as presence/absence data with FastGap 1.2 (Borchsenius 2009) following the simple indel coding algorithm (Simmons et al. 2001). The final alignment including nucleotide and binary data was analyzed in RAxML (Stamatakis 2014) and MrBayes 3.1.2 (Ronquist and Huelsenbeck 2003). Maximum Likelihood (ML) phylogenetic reconstruction was performed in raxm-lGUI (Silvestro and Michalak 2012) using rapid bootstrap analysis with 2000 replicates. Three nucleotide partitions (ITS1, 5.8S, ITS2) were set to the GTRGAMMA substitution model in addition to one binary partition (indel characters) that was set to default. In the Bayesian Inference (BI) phylogeny, the alignment was divided into four partitions (ITS1, 5.8S, ITS2, and indels) as well. The GTR + Γ substitution model was applied to the nucleotide characters, while the two-parameter Markov model was set for the indels. Two independent runs of four Markov Chain Monte Carlo (MCMC) were performed each for 5,000,000 generations, sampling every 1000th generation. The first 30% of the trees was discarded as burn-in. For the remaining trees, a 50% majority rule consensus phylogram with posterior probabilities as nodal supports was computed. Intra-and interspecific genetic differences were calculated by dividing the number of differences (substitutions and/or indels) found in the whole ITS region by the length of the region (ca. 610-618 bases long). The best scoring ML tree from Maximum Likelihood analysis was further edited in MEGA 7 (Kumar et al. 2016) and Adobe Illustrator CS4 and shown in Figs. 1 and 2.

Morphological study
The morphological descriptions of the species are based on notes taken from fresh collections and associated photographs, fungarium specimens, and previously published descriptions of the holotype specimens and other sequenced materials or collections that are not sequenced. Microscopic characteristics are taken from air-dried specimens. Dextrinoid basidiospore reactions were recorded from pieces of lamellae placed in Melzer's reagent for 5 min. Lamella trama hyphae were examined for encrusting pigment in Melzer's reagent and 3% KOH. The pileipellis and basidia were examined in 3% KOH. Basidiospore measurements were made in 3% KOH from Table 1 The nrDNA ITS sequences of Cortinarius species used in the phylogenetic analysis. Newly generated sequences are in boldface.    basidiospores on lamella surfaces and those deposited on the stipe apex, pileus surface, and/or veil. Basidiospore measurements, averages, and Q values (L/W) are based on 20-30 basidiospores per collection, and measurements in parentheses (e.g., (10)) are exceptional. The category "Additional specimens examined" are those collections that were examined morphologically and microscopically but do not have any molecular data associated with them. Herbarium designations follow Thiers (continuously updated).

Phylogeny
A total of 209 sequences, 191 of which belong to sect. Anomali, were included in the analyses ( Table 1). The ITS alignment comprises 774 characters. After gap coding, a binary set of 183 characters was added to the nucleotide alignment, so the data matrix prior to phylogenetic analysis was composed of 957 characters (alignment is deposited as Supplementary information). Cortinarius bolaris (Pers.) Fr. (sect. Bolares) served as outgroup. Phylogenetic trees from ML and BI analyses showed congruent topologies, at least the grouping of the species agreed well in both phylogenetic trees. However, some topological differences in the deeper branches were observed.
Based on our phylogenetic analyses, most species are monophyletic and strongly supported. There are, however, two cases of species limits not fully resolved. In the first case, the monophyletic European C. lepidopus and the very closely related C. modestus are paraphyletic. In the second case, C. tabularis, C. anomalopacificus, and C. sp6 form a clade with C. nettieae within the /tabularis clade, but C. nettieae itself does not form a monophyletic lineage. In both cases, we recognize these lineages as separate species based on differences in the ITS region, morphology, and/or distribution.
The majority (94%) of the studied North American species have intraspecific sequence variation between 0 and 0.5% which means 0-3 substitution and/or indel differences in the ITS region. Cortinarius anomalovelatus, C. rarus, and C. sp1 have 0.6% and 0.8% intraspecific variations (4 and 5 substitution and/or indel differences), respectively.
Most of the species (92%) have 1-5% interspecific sequence dissimilarity which means 6-30 substitution and/ or indel differences in the ITS region. The three species that have interspecific difference from their closest relatives less than 1% (i.e., C. modestus: 0.3%, C. nettieae 0.6%, and Cortinarius sp6: 0.6%) have zero intraspecific variability. The average ITS variation within the studied species was 0.15% in contrast to the significantly larger genetic dissimilarity (1.93%) between the species. The ten sequenced North American type materials (see Taxon sampling part above) clustered in eight of the 43 recognized lineages, viz. C. albidipes, C. albidoavellaneus, C. brevissimus, C. caesiifolius, C. clintonianus, C. deceptivus, C. modestus, and C. perviolaceus, while C. caesiellus and C. copakensis later become synonyms of C. albidipes and C. albocyaneus, respectively. Cortinarius albomalus, C. anocorium, C. anomalovelatus, and C. barlowensis, recently described from North America, with available published ITS sequences (Ammirati 2014;Liimatainen and Niskanen 2021), were confirmed as separate species. The four classical European species C. albocyaneus, C. anomalus, C. caninus, and C. tabularis have been shown to occur in North America. Fourteen of the 27 remaining species are described as new to science based on molecular, morphological, and ecological data. Thirteen species were not described formally in this work due to lack of sampling, morphological, and/or ecological information. These were treated as Cortinarius sp1-sp13 in the phylogeny (Fig. 2 Description: Basidiomata small to medium or less commonly large in size, often somewhat fragile; young lamellae often bluish to violaceous, less commonly whitish to pallid; universal veil white or buff to yellowish or ochraceous, poorly to strongly developed; stipe often discoloring yellowish, base often enlarged; basidiospores globoid to subgloboid or broadly ellipsoid to ellipsoid, mostly moderately to more coarsely verrucose, often similar in size across species; lamella trama hyphae smooth or slightly encrusted; pileipellis typically with a distinct hypocutis below a ± well defined epicutis; clamp connections present; associated with broadleaf trees, conifers, shrubs, and subshrubs. Peck, Bull. N.Y. St. Mus. 157: 57 (1912)   decurved, surface viscid when fresh, glabrous or fringed with silvery white fibrils along the margin, shiny when dry, color evenly pale silvery gray to pale buff or ± gray vinaceous brown, the disc slightly more brownish, cinnamon buff to brownish cinnamon, then ± brownish over margin in age. Lamellae adnate with decurrent tooth to slightly adnexed, ± close, pale violet lilac to pale blue to pale drabgray, finally ± pale brown to cinnamon. Stipe (40) 60-100 mm long, 8-15 mm thick above, equal to somewhat clavate or clavate-bulbous, base 12-15 mm thick, dry, white to whitish silky fibrillose over a lilac violet or dull gray lilac ground color when fresh, basal mycelium white becoming ochraceous to watery gray, veil forming distinct pale orange buff, pale yellow buff or pale ochraceous zones and patches on lower stipe or these less commonly very pale (whitish) to nearly absent, sometimes disappearing in age. Context of pileus firm, mottled with violet to gray violet becoming white, in stipe solid, above pale bluish drab to pale violet lilac, below some violet at first then whitish or discolored. Odor of lamellae and/or context not distinctive, taste of context mild, not distinctive or fungoid. .5 (11) × 6.7-7.8 µm, av. 9 × 7.5 µm, Q = 1.1-1.3, av. Q = 1.2, broadly ellipsoid to subgloboid, moderately to coarsely verrucose, slightly to moderately dextrinoid. Basidia 4-spored, clavate, colorless to yellowish, some with refractive particles. Lamella trama hyphae not encrusted. Pileipellis duplex: epicutis well developed, hyphae ± cylindrical, moderately to strongly interwoven, 2.5-7.5-µm wide, colorless or yellowish, a few slightly encrusted; hypocutis well developed, hyphae ± cylindrical to ± enlarged, (5.5) 7.5-23 µm wide, colorless to yellowish, smooth or slightly encrusted. Clamp connections present.

Cortinarius albidipes
Ecology and distribution: Cortinarius albidipes is associated with a broad range of broadleaf trees including Carpinus, Carya, Fagus, Quercus, and Ostrya, sometimes in combination with Fraxinus and Vaccinium, or with Pinus and Populus, occurring in temperate, boreal, and subalpine forests indicating a broad ecological range. It has an extensive distribution, occurring in Northeastern USA, Québec, Canada, and the Rocky Mountains. According to our unpublished sequence data, C. albidipes also occurs in mixed conifer and broadleaf forests of Northern Europe.
Comments: This is a rather large, stout species with ± persistent violet to lilac colors in the lamellae and stipe apex and rather large basidiospores (av. 9 × 7.5 µm) that are commonly broadly ellipsoid. Smith (1944) separated C. caesiellus from C. albidipes by the presence of the orange buff veil zones on the stipe of the former. Peck in the type description of C. albidipes emphasized the white stipe and absence of a colored veil. The additional collections of C. albidipes we examined all have some orange buff veil zones on the stipe surface. In our phylogeny, it belongs to the /albidipes clade together with two European species, C. lividomalvaceus and C. pelerinii, and 5 additional species from North America (C. tetonensis, C. huddartensis, C. perrotensis, C. kranabetteri, and Cortinarius sp5). Its most closely related species is European C. lividomalvaceus from which it differs by 13 substitutions and indel positions, with a similarity of 97.9%.
Ecology and distribution: From type description: "gregarious in wet grassy area in cedar swamp." Photo shows leaves of broadleaf trees. Known only from the Great Lakes region (type material). Cedar swamps in Michigan are mainly composed of Thuja occidentalis; however, the type material has leaves of broadleaf trees at the base of the stipe, and likely it was a mixed forest.
Comments: Cortinarius albidoavellaneus has a rather slender habit and long stipe, lacks distinct violaceous coloration and the veil leaves only slight, pale-colored remnants on the stipe surface. The basidiospores are darkly pigmented, coarsely verrucose, relatively large (av. 9.5 × 8 µm) and subgloboid. Phylogenetically, this species belongs to / albocyaneus clade together with C. albocyaneus, the European C. epsomiensis, C. jonimitchelliae, the North American C. caeruleoanomalus and Cortinarius sp7. The latter is the most closely related species differing by 10 substitutions and indel positions, with a similarity of 98.4%.  Dima et al. 2016). GenBank ITS: KX302206.
Ecology and distribution: Associated with Betula and Fagus or other broadleaf deciduous trees, such as Tilia and Populus. Widespread in Europe and known from northeastern North America.
Comments: Cortinarius albocyaneus is characterized by its very pale bluish gray to whitish, weakly hygrophanous pileus and the stipe is often with sparse veil remnants. Bluish tinges are present on the pileus, lamellae, and stipe apex in young specimens. The description of the type collection of C. copakensis is rather rudimentary, so parts of the above descriptive data here are adapted from Dima et al. (2016). Basidiospores of the type of C. copakensis are similar to the measurements of those for C. albocyaneus . The ITS sequence of the type of C. copakensis is partial (containing ITS1 and ITS2, but lacking 5.8S), and it differs by only one indel position from the epitype of C. albocyaneus (KX302206). Therefore, both species are considered here as conspecific from both morphological and phylogenetic perspectives. The sister species of C. albocyaneus is C. caeruleoanomalus from which it differs by 7 substitutions and indel positions, with a similarity of 98.9%. The taxonomy of C. copakensis requires further research.
Ecology and distribution: Gregarious under Picea in boreal and subalpine habitats. Known from interior Alaska and the Rocky Mountains.
Comments: Distinguishing features are the moderately large spores that tend to be broadly ellipsoid and coarsely verrucose, rather slender stature, distinct, yellowish buff veil on the stipe and moderate violaceous coloration. Phylogenetically, it belongs to the /caninus clade with C. caninus, C. anomalomontanus, and three unnamed species, two from North America (Cortinarius sp8 and Cortinarius sp9), and one from Japan. The most closely related species is Cortinarius sp8 from which it differs by 8 substitutions and indel positions, with a similarity of 98.7%.

Cortinarius anomalomontanus
Etymology: Named for the mountain habitat in which it occurs.
Description: Pileus 20-24 mm diam., conic to obtusely conic, margin incurved to decurved, moist, subhygrophanous, innately fibrillose along edge, minutely appressed fibrillose scaly on margin, minutely scaly on disc, margin grayish brown with a slight grayish layer of fibrils, in age pale grayish brown with more brownish minute scales, disc grayish brown with slight yellowish to light brown cast and small brown scales. Lamellae adnexed, close to ± distant, light grayish lilac, gradually brownish, pale lilac color ± persistent, edges slightly undulate, paler than faces. Stipe 50-60 mm long, 6-7 mm thick above, base 8-11 mm thick, clavate, apex whitish or pale with very slight lilac tints, at first with whitish veil fibrils above, below whitish mixed with brown and sometimes faint grayish lilac tones, with slight pale ochraceous veil zones and fibrils, basal mycelium whitish. Context of pileus solid, firm, whitish, gray to watery gray over lamellae, stipe cortex solid, hard, pith soft, in apex light violet with white streaks, below whitish with grayish to brownish tones in cortex, whitish in pith. Odor and taste of context strongly fungoid.
Ecology and distribution: Known from the Rocky Mountain forests with Picea engelmannii and Pinus contorta.
Comments: This species features lilac tints in the lamellae and stipe apex, a gray brown pileus, and slight ochraceous veil bands on the stipe surface. The basidiospores are globoid to broadly ellipsoid and rather coarsely verrucose. Phylogenetically, it belongs to the /caninus clade with C. caninus, C. anomalodelicatus, and three unnamed species, Cortinarius sp8 and Cortinarius sp9 from North America and one from Japan. The most closely related species is Cortinarius sp8 from which it differs by 7 substitutions and indel positions, with a similarity of 98.9%.
Other Etymology: From Greek: "anomalo" meaning unusual and "pacificus" refering to the Pacific region of distribution.
Description: Pileus 25-70 mm diam., hemispheric to convex then plano-convex, margin involute, frequently upturned in age, undulate, disc ± even, surface smooth to silky, not hygrophanous, slightly gelatinous when moist, innately fibrillose in age, color pale tan with lilac hues, pale silvery gray along edge, with watery gray drab blotches, central area ± tinted light gray vinaceous to dark gray vinaceous, becoming darker tan with ochraceous discolorations, velar remnants often on margin. Lamellae sinuate to adnexed, close to crowded, light bluish or grayish lavender at first then pale tan to pale yellow, turning rusty brown mature, edges slightly crenate. Stipe 40-120 mm long, 9-20 mm thick, cylindrical to subclavate, fibrillose to silky at the apex, with a distinct annular zone, pale tan with bluish apex, discoloring to yellow tan or whitish, veil white, rarely pale yellow, partially covering the lower stipe, occasionally leaving buff to light buff zones or patches, sometimes indistinct. Context solid, firm, grayish to whitish or bluish, hollow in stipe, whitish to pale blue-lilac or pale lavender above, silvery along surface, below grayish to pale yellow brown to sordid brownish. Odor of context not distinctive or ± raphanoid, taste of context fungoid.
Ecology and distribution. Frequently associated with Notholithocarpus, sometimes also with Quercus, often in combination with Pseudotsuga, Pinus, Picea, and/or Tsuga. Moderately common at autumn in the northern California coastal forests and Sierra Nevada foothills.
Comments: Cortinarius anomalopacificus is a fairly typical representative of section Anomali. However, it lacks the strong violaceous coloration of some species. It has only a slightly colored veil and it has relatively small basidiospores. Phylogenetically, it is in the /tabularis clade with C. tabularis, 1 3 C. brevissimus, C.  Description: Pileus 11-45 mm diam., rounded conic to hemispheric then obtuse to plane, umbonate to subumbonate, margin incurved to decurved then plane to uplifted, rather fragile, surface appressed fibrillose, fibrillose tomentose or fibrillose scaly from a silvery to whitish or ochraceous buff veil, margin and disc grayish lavender to grayish blue beneath veil, in age becoming more strongly ochraceous overall, not hygrophanous. Lamellae adnexed, ± subdistant, grayish violet blue to light blue violet gray, gradually more grayish to brownish, eventually pale brownish cinnamon buff then medium brown, edges whitish at first then more conclorous with faces. Stipe 35-54 mm long, 3-6 mm above, 4.5-9 mm at base, clavate, ± tapered below, upper portions silky, shiny, lilac violet to violaceous at first, gradually becoming whitish to brownish, veil leaving heavy white to buff or creamy buff floccose zones on the surface. Context solid, whitish to grayish in disc, watery gray over lamellae, beneath pileus cuticle tinted cinnamon buff, stipe stuffed then hollow, above bright blue violet with some whitish streaks and at times a grayish cast, in lower stipe whitish to grayish, with age the blue violet color fades, eventually whitish to grayish or brownish, in a few places with orange brown cinnamon discolorations. Odor of context and/or lamellae fungoid, taste of context fungoid, mild or slightly astringent.
Ecology and distribution: Cortinarius anomalovelatus occurs in low-to mid-elevation forests on the western mountain slopes and coastal areas from southwestern Canada into northern California. It tends to prefer more mesic forests and occurs from late summer into the fall season. In conifer forests with Tsuga, Abies, Picea, and/or Pinus.
Comments: Cortinarius anomalovelatus is a very distinctive Pacific Northwest species due to the heavy universal veil and grayish blue to violet colors of young specimens. In older specimens, the veil zones may be less conspicuous and the pileus more ochraceous. It often occurs in the same forest, as C. barlowensis, but usually appears earlier in the season. Phylogenetically, it forms a strongly supported clade with C. deceptivus and C. harvardensis, sister to /lepidopus clade based on our analyses (Figs. 1 and 2). Its most closely related species C. deceptivus differs by 13 substitutions and indel positions, with a similarity of 97.9%. Description: Pileus 18-68 mm diam., convex, margin incurved, becoming broadly umbonate, margin incurved to decurved, not viscid, somewhat hygrophanous, violaceous to grayish buff with slight violaceus tones when moist, when dry buff with a sheen, becoming more brownish with maturity especially on disc. Lamellae adnexed, ± crowded, pale violet, violaceus gray, then clay brown with age. Stipe 45-75 mm long, 4-15 mm thick above, equal to clavate, apex violaceus, otherwise pale violaceus to bluish gray or white to buff, soon becoming grayish to grayish brown, veil ochraceous to grayish brown, leaving a zone above, basal mycelium white. Context in pileus off-white to pale buff or pale violaceous, in stipe apex pale violaceus, below violaceus to bluish or pale yellow, with hygrophanous streaks. Odor of lamellae raphanoid or indistinct.  µm, 8 × 6.5 µm, Q = 1.1-1.4, av. Q = 1.25, subgloboid to broadly ellipsoid, moderately to coarsely verrucose, slightly or moderately dextrinoid. Basidia 4-spored, rarely 2 or 3 spored, 30-40 × 8-9 µm, clavate, colorless or with yellow contents, some with refractive granules. Lamella trama hyphae colorless, smooth to slightly encrusted, scattered yellow pigmented hyphae. Pileipellis duplex: epicutis well developed, hyphae interwoven to entangled, radially arranged on margin, cylindrical, 5.2-9 µm wide, colorless or with yellow contents, smooth or slightly to coarsely encrusted; hypocutis well developed, some areas strongly cellular, otherwise hyphae more interwoven and radially arranged, cylindrical to very enlarged, 7.5-33.5 µm wide, colorless to yellowish, walls refractive, yellow to colorless, smooth or encrusted. Clamp connections present.
Ecology and distribution: Associated with broadleaf and broadleaf-conifer forests in northern and southeastern North America. Known from Québec south to North Carolina and 1 3 Tennessee. It is a widespread and frequent species in Europe in deciduous broadleaf and conifer forests as well.
Comments: Cortinarius anomalus features a violaceous to bluish pileus surface, lamellae and stipe apex when young, an ochraceous-tinted veil, and moderately large (8 × 6.5 µm) subgloboid to broadly ellipsoid, moderately to coarsely verrucose basidiospores. North American collections are similar to those from Europe. Phylogenetically, it belongs to the isolated /anomalus clade along with an unnamed species from Japan and C. sericeolazulinus known from Costa Rica, from which it differs by 18 substitutions and indel positions, with a similarity of 97.1%.
Ecology and distribution: Cortinarius barlowensis is characteristic of mature and old growth submesic to mesic conifer forests composed of Tsuga, Abies, Picea, and/or Pseudotsuga. Known from the Pacific Northwest coastal to low elevation forests, Oregon, Washington, and British Columbia.
Comments: Cortinarius barlowensis has a rather long, narrow stipe in relation to the size of the pileus. The pileus surface is vinaceous gray to purple gray but soon develops brown coloration. The lamellae and stipe are blue to violaceous, and the universal veil leaves faint to thin yellowish buff bands on the stipe below a distinct single zone. The basidiospores are large (av. 10.8 × 6.6 µm) and ellipsoid. Based on available data, it has a quite isolated position in the phylogenetic tree (Fig. 2). The most closely related species is C. rarus from which it differs by 18 substitutions and indel positions, with a similarity of 97.1%.
Context of pileus whitish, in stipe hollow, pale violaceous. Odor and taste not recorded.
Ecology and distribution: Known only from New York, on ground in woods, and a GenBank sequence (JX030219) which shows an association with Castanea dentata; otherwise, we know very little about its host trees.
Comments: The description of the type collection is rarther short and cryptic and there is no illustration. No additional collections are known to date. Therefore, this species is poorly known. The whitish to gray brown pileus, violaceous lamellae and stipe, and its medium-sized basidiospores (av. 8 × 5.5 µm) are not particularly diagnostic in this clade. This is assuming that the short stature of the type collection is not a diagnostic feature. Phylogenetically, it is in a distinct position within the /tabularis clade. Cortinarius nettieae is the most closely related species from which it differs by 23 substitution and indel positions, with a similarity of 96.3%.
Etymology: The epithet refers to the bluish basidiomata of the species.
Ecology and distribution: In broadleaf and broadleafconifer forests, oak and birch or birch and hemlock. Known only from southeastern North America.
Comments: A striking blue to violaceous species with a white veil and rather small (av. 7 × 6 µm), mostly subgloboid basidiospores. Phylogenetically, it belongs to the /albocyaneus clade which contains C. albocyaneus, C. albidoavellaneus, C. sp7, C. epsomiensis, and C. jonimichelliae. It is a close sister species to C. albocyaneus from which it differs by 7 substitution and indel positions, with a similarity of 98.9%. Description: Pileus 50-90 mm diam., obtusely rounded to convex, expanding to nearly plane, sometimes broadly umbonate, edge inrolled, margin incurved, not hygrophanous, surface dry, innately fibrillose at first, nearly glabrous in age, margin usually fringed with yellowish fibrils, at first near clay color to cinnamon buff or more yellowish, becoming darker in age, tawny olive, medium brown, vinaceous brown, dark brown or rich reddish brown, outer margin usually paler, at times edge with faint violaceous cast. Lamellae adnate or slightly adnexed, close to crowded, at first pale lilac, pale violaceous, gray lilac, pale gray blue or grayish blue, becoming dull brownish, edges even. Stipe 62-130 mm long, 9-20 mm thick, clavate, ventricose with tapered base or ± bulbous, base 18-25 mm thick, silky fibrillose, pale violaceus or white with pale lilac tones, veil distinct, forming yellow ocher to rich ochraceous zones and patches, basal mycelium white, sometimes with white rhizomorphs in substrate. Context of pileus solid, slightly violaceous or whitish to pallid brownish drab, in stipe solid, violaceus to grayish lilac, white towards base. Odor of context and/or lamellae indistinct, fetid but soon vanishing or raphanoid. Taste of context mild.
Ecology and distribution: In conifer forest, Pseudotsuga, Tsuga, Abies, Pinus, and/or Larix, with broadleaf trees and shrubs, Quercus, Populus, Alnus, and Salix sometimes present. Rather common species, known from British Columbia to California and east to Idaho and Montana.
Comments: The distinctive yellow ochre veil, persistent lilac lamellae, and rather small basidiospores (av. 7.2 × 5.6 µm) are helpful in distinguishing C. caesiifolius from other Western species in section Anomali. Based on available knowledge, this is a singleton species with an isolated position in the phylogenetic tree (Fig. 2). The most closely related species are C. rarus and C. huddartensis from which it differs by 20 substitution and indel positions, with a similarity of 96.8%.
Ecology and distribution: Often with Picea, or combinations of Picea, Pinus, Pseudotsuga, Tsuga, and/or Abies, sometimes mixed with Populus in boreal habitats. This transcontinental species is rather common in boreal forests across North America, in western subalpine conifer forests and west coastal mesic forests. Widespread in Europe as well, under Picea.
Comments: Cortinarius caninus is a medium to larger species that sometimes appears in large numbers, especially with Picea. The pileus is usually dominated by brown colors, especially by maturity. However, white or gray to violaceous colors may be present in fresh, young specimens, especially along the pileus edge. The cream buff veil patches and zones on the stipe surface may be poorly formed in some specimens. Basidiospore size is rather consistent between collections with an average of 8.5 × 7 µm in North American material. Phylogenetically, it forms the /caninus clade together with C. anomalodelicatus, C. amomalomontanus, Cortinarius sp8, and Cortinarius sp. from Japan. The most closely related species is Cortinarius sp8 from which it differs by 8 substitution and indel positions, with a similarity of 98.7%.  Etymology: Named for Clackamas County, Oregon. Diagnosis: Medium to large basidiomata; pileus gray brown, vinaceous brown or purplish brown; lamellae grayish brown to light brown or purplish tinted; stipe silky white to watery brown; veil buff to brownish; basidiospores av. 9.7 × 6.5 µm, ellipsoid to broadly ellipsoid, rarely subgloboid, moderately to coarsely verrucose.
Description: Pileus 20-53 mm diam., convex to obtusely convex, margin decurved, edge narrowly incurved, surface moist, somewhat hygrophanous, pale silky fibrillose, disc minutely punctate and finely rimose, color evenly light purplish brown to gray brown or purplish brown, edge paler, disc more brownish or deeper brown to dark vinaceous brown, margin ± hygrophanous, often becoming more grayish. Lamellae adnexed, close to ± subdistant, pale grayish to very pale gray violaceous then light brown, sometimes with slight grayish tones, eventually deeper brown, edges somewhat uneven. Stipe 30-90 mm long, 4-10 mm thick above, base narrowly clavate, 5-11 mm thick, at times somewhat narrowed downwards, surface shiny, silky whitish, similar below but developing watery brown coloration, basal mycelium whitish, veil fibrils, zones and patches slightly buff to brownish, nearly lost in age. Context in pileus solid, firm to ± soft fragile, pale purplish brown to grayish purple, watery gray or brown, faded area whitish to pale brownish, in stipe becoming hollow, in apex light gray purple, below very pale brown to whitish, becoming watery brownish in age. Odor of lamellae and/or context fungoid, indistinct or slightly raphanoid, taste mild.
Ecology and distribution: Gregarious in deep humus under mixed conifers, Picea, Pinus, and Abies. In Pacific Northwest conifer forests.
Comments: This is a very elegant species, somewhat resembling C. barlowensis. The pileus is somewhat hygrophanous, vinaceous brown to purplish brown, with the margin becoming more grayish when dehydrating. The lamellae and stipe have limited violaceous coloration. The universal veil remnants on the stipe surface are tinted buff to brownish. The large basidiospores (av. 9.7 × 6.6 µm) are typically ellipsoid to broadly ellipsoid and rather coarsely verrucose. Phylogenetically, it is the sister species to the morphologically similar C. latiodistributus from which it differs by 12 substitution and indel positions, with a similarity of 98.1%. The two species form an isolated clade, tentatively named as the /latiodistributus clade.
Ecology and distribution: In conifer forests of Pseudotsuga and Tsuga, Pinus and Thuja or Pinus and Abies, sometimes with Populus and Betula. Known from the Pacific Northwest to the Great Lakes region and into the northeast, New York, Ontario, and Québec.
Comments: Cortinarius clintonianus appears to be a rather widespread species associated with mature conifer forests. The violaceous to gray violet colors of the young lamellae, pileus margin, and stipe are soon replaced by pallid to brownish colors or are nearly absent in some collections, giving the impression of a Telamonia s. str. Violaceous tones may persist in the stipe apex. The veil zones on the stipe surface are pale to buff. The basidiospores are coarsely verrucose, rather small (av. 7.5 × 6 µm) and subgloboid to broadly ellipsoid. There are two collections (syntypes) by C. H. Peck. One collection is from Croghan and the other one is from New Scotland, and the latter collection is selected as the lectotype for this species. Phylogenetically, it forms an isolated clade with a species known only from environmental sequence data from Japan (Fig. 2). The most closely related species are C. rarus and C. huddartensis from which it differs by 21 substitution and indel positions, with a similarity of 96.6%.
Other Description. Pileus 10-70 mm diam., suborbicular to hemispherical, becoming convex to convex-campanulate, basic color strongly grayish blue violet to fawn-colored tinged with lavender, the blue to lavender colors fading very quickly, becoming pure fawn or light tan, disc tan buff, initially covered by yellowish universal veil, which leaves distinct fibrillose patches on the margin or minute, brownish fibrillose squamules on the surface, eventually becoming subglabrous, somewhat hygrophanous, rugulose in age. Lamellae ± thick, crowded to moderately close, adnate, deep blue to lavender when young, becoming grayish then pale tan with age. Stipe 30-108 mm long, 4-10 mm thick, rather stout, clavate at first then elongate and slender to cylindrical and flexuous with a slightly enlarged base, entirely bluish, at first covered by the yellowish fibrillose veil, eventually with brownish scales and partial zones, these terminate at a zone with the partial veil which is pale at first and then covered by spores, basal mycelium violet. Context hygrophanous, blue to pallid with a strong lavender tinge when young, fading quickly, becoming cork color, thick on disc, thin towards margin, texture spongy, in stipe solid, soon becoming hollow, entirely blue with stronger blue colors at apex and cortex. Odor of lamellae and/or context not distinctive or faintly unpleasant, sweetish.
Ecology and distribution: C. H. Kauffman reported that it was common in hemlock woods, on ground and among remains of very rotten logs and brush heaps. It also occurs with Pinus, August to September. Known from northeastern North America.
Comments: Cortinarius deceptivus was apparently collected more than once. The type is from Coy Glen, New York. Kauffman stated: "The colors and shapes of the plant are very variable and deceptive." Based on the photos of collections, Kauffman was certainly correct about the coloration of basidiomata. Fresh, young specimens are strongly bluish throughout but eventually this color fades and becomes more grayish. The well-developed yellowish universal veil covers the pileus surface at first, leaving behind fibrillose patches as the pileus expands, and distinct yellowish to brownish zones on the stipe surface. Phylogenetically, it forms a strongly supported clade with C. anomalovelatus and C. harvardensis. The most closely related species is C. anomalovelatus from which it differs by 13 substitution and indel positions, with a similarity of 97.9%.
Etymology: The epithet refers to the Harvard Forest, the locality where the holotype was collected.
Ecology and distribution: In conifer and mixed coniferbroadleaf forests. Known from eastern North America, Quebec south to Massachusetts.
Comments: This medium-sized species has a violaceous pileus and lamellae when young but mature specimens become more grayish to brownish. The veil is sparse and whitish to yellowish, and the basidiospores are similar in size to those of C. deceptivus and C. anomalovelatus, which form the sister clade to this species within the /lepidopus clade. The most closely related species is C. anomalovelatus from which it differs by 21 substitution and indel positions, with a similarity of 96.8%.

Cortinarius huddartensis
Diagnosis: Basidiomata large to medium size; at first with pale lavender colors in the pileus and lamellae and a blue stipe apex; universal veil and cortina white; basidiospores av. 9.0 × 6.2 µm, broadly ellipsoid, coarsely verrucose.
Ecology and distribution: Thus far only known from California in association with Quercus. Its distribution may be limited to the warmer parts of the Pacific Coast.
Comments: Cortinarius huddartensis occurs under oaks, including Quercus agrifolia, Quercus douglasii, and Quercus wislizenii. The species exhibits pale lavender to pale blue coloration when fresh and has a white universal veil. The basidiospores are large (av. 9.0 × 6.2 µm), broadly ellipsoid and coarsely verrucose. Phylogenetically, it belongs to the /albidipes clade, with C. albidipes, C. kranabetteri, C. perrotensis, C. tetonensis, and C. sp5, as well as with two European species, C. lividomalvaceus and C. pelerinii. The most closely related species is C. kranabetteri from which it differs by 10 substitution and indel positions, with a similarity of 98.4%.
Ecology and distribution. Known from British Columbia and Alberta in late successional conifer or mixed forests with Populus.
Comments: Cortinarius kranabetteri has relatively small basidiospores (av. 7 × 6 µm) which it shares with the sister species C. perrotensis. Based on descriptions to date, this species does not have strong persistent blue coloration in the lamellae and stipe, and the veil zones on the stipe surface are pale ochraceous. Phylogenetically, it belongs to the /albidipes clade, with C. albidipes, C. huddartensis, C. perrotensis, and C. tetonensis, C. sp5, as well as with two European species, C. lividomalvaceus and C. pelerinii. The most closely related species is C. perrotensis from which it differs by 6 substitution and indel positions, with a similarity of 99%.

Cortinarius latiodistributus
Etymology: The name "latiodistributus" refers to the wide distribution of the species.
Description: Pileus 30-45 mm diam., convex, slightly umbonate, margin incurved, slightly inrolled at edge, dull white veil fibrils on edge of margin at first, surface moist, ± hygrophanous, margin primarily gray brown, at times violaceous, becoming more brownish with age, around disc reddish brown, disc gray brown violaceous, becoming brownish with age, faded areas watery gray to pallid brown. Lamellae adnexed, close, dull violet mixed with gray tones at first, then gray brown, eventually more brownish, edges pale, whitish, ± even. Stipe 70-73 mm long, above 5-6.5 mm thick, base clavate-bulbous, 10-14 mm thick, at first light blue to near base, becoming white in age (almost no hint of blue lavender color remains), at base more grayish, basal mycelium white, veil whitish to ochraceous or brownish ocharceous, leaving a distinct band mid-stipe, below and fibrils and patches below. Context of pileus solid, firm, watery concolorous, in buttons with violet tones in places, fading to whitish or pale brown, context of stipe hollow mature, light violet above with some whitish streaks, lower portion more grayish to white or watery brown, slight violet tints in places. Odor and taste of context mushroom-like.
Ecology and distribution: In mature and old growth mesic conifer forests, with Tsuga and Abies in the Pacific Northwest of North America. Also known from Japan, Finland, Norway, and Sweden with Picea, Pinus, sometimes mixed with Betula.
Comments: This species occurs in similar habitats as C. barlowensis in the Pacific Northwest. In the field, these two taxa can appear somewhat similar; however, C. latiodistributus has deeper violaceous colors, a more distinct veil, and significantly smaller basidiospores than C. barlowensis. Phylogenetically, it is distantly related to C. barlowensis, but forms an isolated clade with its sister species C. clackamasensis, which is similar in appearance but has significantly larger basidiospores and differs by 12 substitution and indel positions, with a similarity of 98.1%.
Other Description: Pileus convex or expanded, 20-70 mm broad, surface subfibrillose or with minute fibrillose patches on disc and inner margin, not hygrophanous, at first pale to light brown with gray violet tints on the outer margin, becoming light brown to pale brown with darker brown center and pale edge, with veil fibrils on outer margin. Lamellae adnexed, close, pale purplish gray at first then pallid to light brown or cinnamon mature, edges pale. Stipe narrowly clavate, rarely bulbous, 45-75 mm long, 4-8 mm thick above, 5-11 mm thick at base, subfibrillose, white overall or apex with pale violaceous tints at first, veil forming yellowish buff zones and patches on the stipe surface, basal mycelium white, sometimes with white rhizomorphs. Context in pileus solid, white, to pale brownish white, in stipe pale violaceous above, white to brownish white below, hollow or stuffed with white pith. Odor of lamellae somewhat raphanoid. Taste not recorded.
Ecology and distribution: Known from eastern North America from Québec to New York. In mixed forests of broadleaf and conifer, including Abies, Picea, and Pinus, sometimes with Betula, Populus, Alnus, and/or other broadleaf trees.
Comments: This species has a predominately brownish pileus with some gray violet tints on the margin at first. The lamellae and stipe apex are pale violaceous at first but the lamellae soon become brownish and the stipe is often white throughout when mature. The universal veil typically leaves yellowish to buff zone on the stipe surface. The basidiospores are rather small (av. 7.5 × 6.5), mostly subgloboid and ± coarsely verrucose. Phylogenetically, it is very closely related to the morphologically similar European C. lepidopus. They might represent the same species, since they only differ by 2 nucleotides in the ITS region (= 99.7% similarity); however, it is consistantly different. Intraspecific genetic variability is 0 in both species. We have not yet successfully sequenced the holotype of C. lepidopus Cooke from Kew Herbarium. Until the European name is fixed by a type sequence, we treat C. modestus as a separate North American taxon.

Cortinarius nettieae
Etymology: Named for the late Nettie Laycock, a wellknown Pacific Northwest citizen scientist and mycologist.
Description: Pileus 40-70 mm diam., obtusely convex to convex, subumbonate with flattened disc, margin incurved, edge slightly inrolled, not hygrophanous, sometimes completely brownish in button stage, normally margin silvery lavender to grayish vinaceous or grayish violet streaked with brown, disc shaded slightly brownish to rusty brown, overall becoming more brownish when mature. Lamellae adnexed, close to crowded, violet when young becoming dull lavender vinaceous, then brownish. Stipe 50-120 mm diam., apex to 5-15 mm thick, somewhat clavate to narrowly clavate, surface white or violet or ± colored as pileus margin, cortina pale lavender at first, veil forming inconspicuous buff to yellowish bands on the stipe, basal mycelium white to lavender. Context in pileus solid, firm to watery mature, at first pale grayish brown, white mature, in stipe becoming narrowly hollow, with violet areas in cortex especially above, otherwise white with pale grayish brown colors below. Odor of lamellae and/or context rapahnoid or not distinctive, taste of context mild.
Ecology and distribution: Known from coastal Oregon and Washington in conifer forests.
of Picea, Pinus, Abies, and Tsuga, from Montana in subalpine forest, with Picea, Pinus, and Abies and from Québec under Betula with Populus, Salix, Abies, and Picea.
Comments: This is a relatively large species compared to other Anomali. The violet to lavender colors of the pileus are not particularly persistent, the pileus soon developing brown to grayish colors. The lamellae and stipe are violaceous at first, and the veil forms only slight buff bands on the stipe surface. The basidiospores average 8 × 6.3 µm, and are mostly broadly ellipsoid. It is in the /tabularis clade with C. tabularis, C. anomalopacificus, Cortinarius sp6, and C. brevissimus. Even though C. nettieae forms a paraphyletic lineage in our phylogenetic tree (Fig. 2), it is distinct from other Anomali species, except for two close sequences from Canada (Cortinarius sp6), which differ by 4 substitution and indel positions with a similarity of 99.4%. The relationship between C. nettieae and Cortinarius sp6 needs further study.
Ecology and distribution: Known from midwestern North America in broadleaf forests with Quercus and Tilia.

Cortinarius perviolaceus
Ecology and distribution: In broadleaf and broadleafconifer forests. Known from Florida, Georgia, Massachusetts, New Hampshire, New York, and Tennessee.
Comments: Cortinarius perviolaceus is a rather small, ± entirely purple to bluish species with small mostly subgloboid basidiospores. Our description fits very well with the type description of W. A. Murrill. We gained only ITS1 from the holotype. Genetically, this is the most isolated species in this study with almost 5% dissimilarity (30 changes) to the closest species, and it occupies a singleton position in Etymology: From Latin: "rarus" meaning rare, scarce. Diagnosis: Basidiomata rather large and robust; pileus, lamellae and stipe with strong grayish blue to bluish colors; universal veil and cortina white; basidiospores rather large, av. 8.5 × 6.5 µm, subgloboid to broadly ellipsoid or ellipsoid, coarsely to very coarsely verrucose.
Ecology and distribution: Infrequent, vernal species in mountain conifer forests of Abies, Pseudotsuga, Tsuga, and/or Pinus, sometimes mixed with Quercus. Known from California, Oregon, and Washington.
Comments: Cortinarius rarus is rare but widespread, known from numerous collections. It is a rather robust representative of sect. Anomali. Its most distinctive features are the somewhat larger spores, the persistent bluish coloration, the white veil, and its occurrence in the spring season. Phylogenetically, it is a singleton species based on our current knowledge. The most closely related species is the European C. pelerinii from which it differs by 16 substitution and indel positions, with a similarity of 97.6%.

Cortinarius tetonensis
Etymology: Name is taken from the Teton Mountains.
Ecology and distribution: Scattered to gregarious in moist soil and litter, subalpine conifer forest of Picea, Abies, and

Morphological characters
One of the most interesting characteristics of species in the section Anomali involves the violet to blue coloration of the lamellae, pileus, stipe and context of basidiomata.
There are a handful of species that have almost completely and persistently violet to blue basidiomata, including C. anomalovelatus, C. deceptivus, and C. harvardensis all in the same clade and C. caeruleoanomalus and C. perviolaceus each in separate phylogenetic groups. By comparison, most of the remaining species, with the possible exception of C. tetonensis, have at least some violet to blue coloration in the lamellae, stipe apex, and/ or pileus surface but in some instances, young, fresh specimens are essential to being certain of the coloration. This is especially true in relatively dry habitats where even young appearing basidiomata can lose violet to blue coloration rather quickly. Some species such as C. albocyaneus, C. anomalus, C. barlowensis, C. huddartensis, C. latiodistributus, and C. rarus tend to maintain violet to blue colors in the lamellae and/or stipe or even the pileus margin for some length of time. Typical of many species are gray to brown colors of the pileus surface, sometimes mixed with pale blue to violet colors, blue to violet colored lamellae that become brown as they mature, and violet to blue pigment in the stipe apex which can be lost with age. The various patterns and changes in coloration shared by many species are what make their identification so challenging. In general, these coloration patterns are found across many of the lineages. Several previous studies (e.g., Frøslev et al. 2007;Liimatainen et al. 2017;Clericuzio et al. 2017;Bellanger 2018;Bellanger et al. 2018) have already demonstrated that the presence/absence of blue/violet hues in basidiomata of Cortinarius groups is highly misleading to distinguish phylogenetically supported species.
While a number of species in Anomali have non-hygrophanous or slightly hygrophanous pilei, there are a few taxa that are more distinctly hygrophanous, these include C. albidoavellaneus, C. barlowensis, C. clackamasensis, and C. latiodistributus.
The size of the basidiomata appears to be somewhat helpful in separating species from one another. However, a full collection of different stages of development is necessary to determine whether or not this is a significant distinguishing characteristic because of the overlap in pileus diameter and stipe thickness of young to mature basidiomata among and within species. Cortinarius perviolaceus seems to be one species that is consistently smaller and more slender than others.
The universal veil in many species of Anomali occurs as buff to yellowish or ochraceous to light orange buff patches, zones and/or fibrils on the stipe surface, and sometimes the pileus surface as well. The degree of development and coloration of these veil remnants, while rather distinctive in many species, for example C. caesiifolius, are variable in others, for example, C. albidipes and C. caninus, and can lead to erroneous identifications. Only a few species, for example, 1 3 C. rarus and C. huddartensis, appear to have universal veil remains that are white and/or lack significant buff to yellowish coloration. As with other characteristics, universal veil color alone cannot be used for identification.

Microscopic characters
As for many groups of Cortinarius, section Anomali does not have many characteristics beyond those of the basidiospores that can be used for identification. The structure of the pileipellis is rather consistent across the section, being composed of a ± well-developed epicutis over a distinct hypocutis composed of cylindrical to enlarged hyphae that is sometimes cellular in appearance. The basidia and lamella tramal hyphae are ± consistently the same across the section, with the development of some yellow pigmentation, but no significant encrusting pigment on the tramal hyphae. The basidia are mainly 4-spored, rarely 2-spored.
Section Anomali, for the most part, produces basidiospores that are subgloboid to broadly ellipsoid in profile view. There are a few species which appear to consistently produce subgloboid basidiospores, but in most instances shape varies from subgloboid to broadly ellipsoid with differing proportions of the two shapes. Several species produce more ellipsoid basidiospores. Basidiospore size is helpful in identifying some species when used in correlation with other characteristics. For example, C. anomalopacificus has rather small basidiospores (av. 6.5 × 5.5 µm) and C. barlowensis has rather large basidiospores (av. 11 × 6.5 µm), as does C. clackamasensis (av. 9.7 × 6.5 µm). However, most species produce basidiospores that are 7-9 × 5.5-7 µm on average. The most accurate measurements are from mature basidiospores deposited on the stipe surface or veil fibrils. The basidiospore ornamentation is difficult to interpret without considerable observation using a 1000 × oil immersion lens and spores mounted in 3% KOH. Differences in the size and pattern of ornamentation may differ considerably within and among taxa so it is important to base comparisons on mature basidiospores.

Ecology and distribution
The patterns of distribution and ecology of most Anomali species in North America cannot be determined for certain because of the small sample size for almost all species treated here. Nonetheless, there are certain patterns of distribution that are noteworthy and to some degree are correlated with ecology and plant hosts. Cortinarius albocyaneus and C. anomalus occur in hardwood and/or mixed hardwood conifer forests in both eastern North America and Europe. By comparison, C. caninus and C. tabularis are known from Europe and across northern North America in conifer and/ or conifer hardwood forests. Cortinarius latiodistributus has an even more extensive distribution including the Pacific Northwest, Japan, and northern European conifer forests. Cortinarius tetonensis, for which we have relatively few samples, has a broad distribution and interesting ecology, being reported from Svalbard, western North America, and Alaska in subalpine conifer or arctic habitats with Dryas. More widely distributed species that occur across North America, often in conifer forests, include C. albidipes and C. clintonianus. Several species have regional patterns of distribution. Cortinarius albidoavellaneus is known only from the type locality in northern Michigan with conifers and one locality in North Europe (data not shown). Several species, including Cortinarius brevissimus, C. caeruleoanomalus, C. deceptivus, C. harvardensis, C. modestus, C. perrotensis, and C. perviolaceus, occur in hardwood, mixed hardwood conifer, and/or conifer forests in eastern North America. Western North America is the home of C. rarus, an unusual vernal species associated with montain conifer forests. Cortinarius anomalodelicatus, C. anomalomontanus, C. anomalovelatus, C. barlowensis, C. caesiifolius, C. clackamasensis, C. kranabetteri, and C. nettieae have a western distribution with conifers. California has two hardwood-associated species, Cortinarius huddartensis in oak forests and C. anomalopacificus in tan oak or oak conifer forests.