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Research Article
New species in Cortinarius section Cortinarius (Agaricales) from the Americas and Australasia
expand article infoEmma Harrower, Neale L. Bougher§, Caitlin Winterbottom|, Terry W. Henkel|, Egon Horak, P. Brandon Matheny
‡ University of Tennessee, Knoxville, United States of America
§ Western Australia Herbarium, Kensington, Australia
| Humboldt State University, Arcata, United States of America
¶ Unaffiliated, Innsbruck, Austria
Open Access

Abstract

Five new species from Cortinarius sect. Cortinarius are formally described, four from the Americas (Cortinarius palatinus Harrower, sp. nov., Cortinarius atrotomentosus Harrower, sp. nov., Cortinarius altissimus Harrower & T.W. Henkel, sp. nov., Cortinarius neotropicus Harrower, sp. nov.) and one from Australasia (Cortinarius carneipallidus Harrower & E. Horak, sp. nov.) based on molecular, morphological, and ecological circumscription. Additional collections of the Australasian species Cortinarius hallowellensis Wood and Cortinarius kioloensis Wood reveal wider host associations and geographic ranges than previously recorded. Morphological descriptions, photomicrographs and a dichotomous key to all species in Cortinarius sect. Cortinarius are provided. This work raises the number of species in sect. Cortinarius to twelve.

Key words

Australia, Costa Rica, Guyana, New Zealand, North America, South America, systematics, taxonomy

Introduction

Recent phylogenetic analyses of Harrower et al. (2015) have revealed greater species-level diversity in Cortinarius (Pers.) Gray sect. Cortinarius (also known as the C. violaceus group) than previously thought (Bougher and Syme 1998, Halling and Mueller 2005, Gasparini and Soop 2008). Phylogenetically defined species within the section exhibit continental scale disjunct distributions and form ectomycorrhizal associations with different plant partners (Harrower et al. 2015). Taking their morphological differences into consideration, a taxonomic revision of the section is warranted.

Cortinarius subgenus Cortinarius sect. Cortinarius is an easily identifiable section within the very large genus Cortinarius. All basidiomata have a deep violet color, the pigment of which has been identified as (R)-3′,4′-dihydroxy-β-phenylalanine [(R)-β‐dopa] in Cortinarius violaceus (L.) Gray (von Nussbaum et al. 1998). The pigment is vacuolar, encrusting (Brandrud 1983) and oxidizes quickly to brown (von Nussbaum et al. 1998). The following features for the section were noted by Brandrud (1983), the circumscription of which as based on European taxa only at the time. The pileus is tomentose to finely scaly (comprising a trichoderm), neither hygrophanous nor viscid. KOH on any surface of a basidiome produces a red reaction. Pleurocystidia and cheilocystidia are present, none of which differ in size, shape and abundance and caulocystidia are present. The basidiospores of all species are verrucose and bear a plage.

Linnaeus (1753) described the first species in the section as Agaricus violaceus L. Persoon (1801) recognized A. violaceus (“ad margines sylvarum”) and A. hercynicus Pers. (“in pinetis”). However, the latter was synonymized with A. violaceus by Fries (1821). Moser (1967; 1969) later recognized Cortinarius hercynicus (Pers.) M.M. Moser as an autonomous species. Brandrud (1983) treated the two species as varieties and later as a subspecies (Brandrud 1990), but Harrower et al. (2015) did not find a genetic or ecological basis for distinguishing these taxa.

Clements and Shear (1931) designated C. violaceus (L.) Gray as the type species of the genus Cortinarius. The subgenus Cortinarius was circumscribed by Orton (1958) to include C. violaceus, C. orellanus, C. cotoneus and other allied species. Moser (1967, 1969) transferred members of the C. orellanus-cotoneus group into subgenus Leprocybe sensu Moser, leaving subgenus Cortinarius represented by only C. violaceus and C. hercynicus. Niskanen et al. (2008) included sections Cortinarius, Dermocybe, Veneti, Limonii, Orellani and Humicolae within subgenus Cortinarius.

As many as twelve species have been recognized in sect. Cortinarius, but the inclusion of some these species is suspect. Members are known on every continent except for Africa and Antarctica (Moser 1968, Brandrud 1983). Cortinarius violaceus occurs throughout North America, Asia and Europe in association with members of the Fagales, Salicaceae and Pinaceae (Brandrud 1983; Nezdoiminogo 1996; Harrower et al. 2015). Horak (1980) described C. gayi Horak from Nothofagus in Chile for C. violaceus Fr. sensu Montagne (Montagne 1989). Singer et al. (1983) described Cortinarius kerrii Singer & I.J.A. Aguiar from the Amazon in campinarana vegetation. Moser (1987) documented C. violaceus Fr. as occurring in mixed deciduous forests in Malaysia and Papua New Guinea. In the same paper, he described C. atroviolaceus M.M. Moser, C. subcalyptrosporus M.M. Moser and C. paraviolaceus M.M. Moser from Mt. Kinabalu, Sabah, Malaysia. Malaysian plant associates were likely in the plant families Fagaceae and Myrtaceae (Beaman et al. 2000, Beaman and Anderson 2004), and Moser also documented C. atroviolaceus and C. subcalyptrosporus as occurring with Nothofagus in New Zealand (Moser 1987). He was uncertain whether C. paraviolaceus belonged in sect. Cortinarius due to the lack of cheilocystidia, atypical pleurocystidia, a pileus that is not strongly squarrose-fibrillose-squamulose and basidiomata that are less intensely violet (Moser 1987). Moser also described C. atrolazulinus M.M. Moser with Nothofagus in New Zealand (Moser 1987). Gasparini (2001) added C. austroviolaceus Gasparini from Australia. This species was excluded from the C. violaceus group by Harrower et al. (2015) due to a lack of pleurocystidia and phylogenetic placement outside the clade containing C. violaceus. Wood (2009) described C. jenolanensis, C. kioloensis and C. hallowellensis from Eucalyptus forests in Australia, but the former lacks pleurocystidia suggesting to us an alliance outside the C. violaceus group. Assuming C. hercynicus is treated as a synonym of C. violaceus then twelve species can be recognized in sect. Cortinarius.

Harrower et al. (2015) produced a multi-gene phylogenetic tree that delimited eight different species present in sect. Cortinarius in the Americas and Australasia. Species were separated by at least a 2% difference in the ITS region, which conforms to the benchmark in Cortinarius established by Stephani et al. (2014). Of the eight species level lineages detected, two new were found in Costa Rica, one in North America, one in New Zealand, and one in Guyana. Here we provide descriptions for all species recognized in this previous study (except for C. violaceus). Additional collections of C. kioloensis and C. hallowellensis (Wood 2009) have been examined extending their known host association and geographic ranges. The purpose of this study is to revise the taxonomy of species in the C. violaceus group found in Australasia and the Americas and refine sect. Cortinarius based on morphological data previously confirmed by molecular phylogenetic results. A key to twelve species in the section is provided.

Methods

Dried specimens of collections labeled as C. violaceus, C. subcalyptrosporus, C. atroviolaceus, C. atrolazulinus, and C. austroviolaeus were obtained from the TENN, ZT, HSC, NY, PDD and PERTH herbaria. Additional collections were provided from Ian Dodd (West Kempsey, Australia). Requests for type collections of C. subcalytrosporus, C. atroviolaceus, C. atrolazulinus and C. paraviolaceus were denied from the IB herbarium. Previous attempts to sequence C. subcalyptrosporus and C. paraviolaceus were purportedly unsuccessful (Ursula Peintner, personal communication).

Morphological analysis

Macroscopic descriptions were taken from fresh material as described by collectors where possible. Color documentation of basidiocarps follows the Methuen Handbook of Colour (Kornerup and Wanscher 1967). When no descriptions were made from fresh material, macroscopic descriptions were assembled from dried material and from photographs of fresh material. By measuring the pileus diameter and stipe length of MEL 2351101 (C. kioloensis) before and after drying, it was estimated that basidiomata were reduced by a factor of 1.75 upon drying. Macroscopic measurements were taken from dried specimens of C. atrotomentosus sp. nov., C. neotropicus sp. nov., C. kioloensis and C. carneipallidus sp. nov. Dry measurements were multiplied by a factor of 1.75.

Microscopic features were observed from sections of dried material rehydrated in 5% KOH on a Nikon Eclipse 80i microscope. Basidiospore measurements were taken from spores trapped in the cortina on the stipe. However, for specimens that did not have any remaining cortina, basidiospore measurements were taken from lamellar tissue, and mature basidiospores (judged by the presence and development of ornamentation) were measured at random. Measurements were taken with a Nikon DS-Fi1 camera and Nikon NIS Elements 3.1 software. Twenty-five basidiospores were measured per collection. Outliers are placed in parentheses. Five basidia, cheilocystidia, and pleurocystidia were measured per specimen. The following abbreviations are used: Q = quotient of length divided by width; Q mean = average value of Q values. Scanning electron microscope images were taken on a Zeiss Auriga scanning electron microscope. Lamellae were placed on double-sided carbon adhesive tape and covered with two layers of a gold sputter coating.

Phylogenetic study

Methods for DNA extraction, PCR amplification, and sequencing are given in Harrower et al. (2015). Fig. 2 in Harrower et al. (2015) is summarized here in part as Fig. 1, showing the phylogenetic relatedness of species within sect. Cortinarius.

Figure 1. 

Maximum likelihood (ML) phylogenetic tree of Cortinarius subgen. Cortinarius sect. Cortinarius using combined nuclear ITS, 28S and RPB2 sequences. ML bootstrap support above 50% is shown above nodes.

Figure 2. 

Habit of the following Cortinarius species: a C. kioloensis (TENN 069666, photo I. Dodd) b C. palatinus (NY 796168, TYPE photo R.E. Halling) c C. atrotomentosus (TENN 069922, photo A. Bessette); d C. hallowellensis (MEL 2300544) e C. neotropicus (NY 75934, photo R.E. Halling) f C. carneipallidus (PDD 71219, photo E. Horak) g C. altissimus (TENN 069829, TYPE photo T. Henkel); h C. gayi (ZT 75/82, ISOTYPE photo E. Horak) i C. austroviolaceus (MEL 2231689, photo K. Syme). Scale bar = 1 cm.

Taxonomy

Cortinarius Fr. subgen. Cortinarius sect. Cortinarius

Synonyms: Agaricus (ser. Cortinaria) trib. Inoloma Fr.: Fr. 1821: 217. – Agaricus subgen. Inoloma (Fr.: Fr.) Loudon 1836: 1000. – Cortinarius Fr. 1836 trib. Inoloma (Fr.: Fr.) Fr. 1836: 279. – Inoloma (Fr.: Fr.) Wünsche 1877: 87. – Type species: Cortinarius violaceus (L.:Fr.) Gray.

Type species

Cortinarius violaceus (L.:Fr.) Gray

Basidiomata tricholomatoid, deep violet; pileus squamose to tomentose, rarely greasy, then simply innately fibrillose; KOH+ red; basidiospores subglobose, ellipsoid to amygdaliform, weakly to strongly verrucose, with suprahilar plage (may only be visible in SEM); pleurocystidia and cheilocystidia present; caulocystidia present or usually absent; pileipellis a trichoderm of hyphae (6–22 µm wide) or rarely an ixocutis with gelatinized hyphae (2–11 µm wide); pigment vacuolar and soluble in water, violet often turning brownish in age.

Key to species of Cortinarius sect. Cortinarius

1 Pileus smooth, greasy; pileipellis an ixocutis C. hallowellensis
Pileus scaly, dry; pileipellis a trichoderm 2
2 Stipe more than two times longer than the width of the pileus C. altissimus
Stipe less than twice as long as the width of the pileus 3
3 Basidiospores with a visible perisporium C. subcalyptrosporus
Basidiospores without a loosening of the perisporium 4
4 Cheilocystidia 25–30 × 9.5–11.5 µm C. kerrii
Cheilocystidia larger than above 5
5 Basidiospores 9–12.5 × 5.5–6.5 µm; Q=1.86; basidia 32–38 × 7.5–8 µm C. atrolazulinus
Basidiospores often longer, less elongate; Q<1.86; basidia width > 8 µm 6
6 Growing on calcareous soil with Quercus virginiana C. atrotomentosus
Growing on a variety of soils; not with Quercus virginiana 7
7 Basal mycelium white C. neotropicus
Basal mycelium pale violet 8
8 Caulocystidia present C. violaceus
Caulocystidia absent 9
9 Occurring with Quercus in Costa Rica C. palatinus
Occurring with Myrtaceae or Nothofagus in Australasia or Fagaceae in Indonesia 10
10 With Nothofagus C. carneipallidus
Not with Nothofagus 11
11 Occuring with Myrtaceae in Australia and New Zealand C. kioloensis
Occuring with Fagaceae in Indonesia C. atroviolaceus

Cortinarius kioloensis Wood, 2009

Figs 1, 2a, 3a, 4a

Type

AUSTRALIA. New South Wales: Batemans Bay, Kioloa State Forest, Eucalypt woodland, 22 Jun. 1980, S. Lowry (holotype: UNSW 83/781).

Description

Pileus 45–120 mm wide, convex to plano-convex with a low, broad umbo with age, surface dry, tomentulose-squamulose, dark violet (17F8), lighter concolorous (17E3) near margin, red in KOH; margin entire, split and broadly undulating with age. Lamellae close, sinuate, thick, broad to ventricose, deep blue (19D–E8). Stipe 105–165 mm long, 7–28 mm thick at apex, 9–31 mm thick at base, bulbous, light violet (18A3–5) turning deep blue (18F8) upon touch because of the copious universal veil. Universal veil pale violet (17A3). Basal mycelium pale violet (17A3). Context at apex dark violet to black (18F8).

Basidiospores 10.5–14 µm × 6.5–9 µm, means = 11.5–12.5 µm × 7.5–8 µm, Q = 1.41–1.63, Q means = 1.45–1.60 (125 spores, 5 specimens), ellipsoid to amygdaloid, strongly verrucose, plage present under SEM. Basidia 4-spored, clavate, (27) 35–49 µm × 8–12 µm. Cheilocystidia ventricose to lageniform, sometimes strangulated, brown in KOH, (50–) 60–100 (–127) µm × (10–) 15–30 (–35) µm. Pleurocystidia narrowly conical to lageniform, sometimes strangulated, brown or grey in KOH, (50–) 60–90 (–105) µm × (15–) 20–25 (–30) µm. Caulocystidia absent but brown vacuolar pigment in some hyphae of the caulopellis. Pileipellis a trichoderm composed of fasicles of multiseptate hyphae, 8–20 µm wide, smooth, terminal cells clavate to lanceolate, with brown contents in KOH. Clamp connections present.

Figure 3. 

Light micrographs of basidiospores from the following Cortinarius species: a C. kioloensis (PDD 99307) b C. palatinus (NY 00795933) c C. altissimus (TENN 069830) d C. hallowellensis (MEL 2300544) e C. neotropicus (NY 34729 TYPE) f C. carneipallidus (PDD 95444 TYPE) g C. atrotomentosus (TENN 065535 TYPE) h C. violaceus (TENN 062899) i C. austroviolaceus (MEL 227499). Scale bar = 10 µm.

Figure 4. 

SEM micrographs of basidiospores from the following Cortinarius species: a C. kioloensis (MEL 2351101) b C. palatinus (NY 796168 TYPE) c C. altissimus (TENN 069830) d C. hallowellensis (MEL 2300544) e C. neotropicus (NY 34729 TYPE) f C. carneipallidus (PDD 71219) g C. atrotomentosus (TENN 065535 TYPE) h C. violaceus (TENN 063104) i C. austroviolaceus (MEL 2121961 PARATYPE). Scale bar = 2 µm.

Ecology and distribution

With Eucalyptus and/or Allocasuarina in southeast Australia. With Leptospermum in New Zealand. Fruiting April to July.

Other specimens examined

Australia. Tasmania: Florentine River Valley, Pagoda Hut, on soil under Eucalyptus and Allocasuarina, 23 Apr. 2003, D.A. Ratkowsky (MEL 2351101). New South Wales: Swans Crossing State Forest (Eucalyptus), 21 May 2003, I. Dodd 70845 (TENN 069666). New Zealand. Auckland: Waitakere Ranges, Little Huia, Donald Mclean Track, 10 July 2010, C. Shirley AK375 (PDD 99307). Auckland: Waitakere, Piha Rd, Upper Nihotupu Dam track, (Leptospermum), 24 July 2010, C. Shirley AK373 (PDD 99309). Auckland: Waitakere, Mountain Rd, Opanuku Pipeline Track, (Leptospermum), 17 July 2010, C. Shirley AK370 (PDD 99308).

Discussion

The collections examined here match the protologue of C. kioloensis Wood in all particulars. One collection (TENN 069666) was seen to have dark purple, nearly black flesh at the apex. Flesh color at the base was not observed. We have not studied the type of this species (the collection was not readily available at UNSW) but are confident for now applying the name C. kioloensis to our material. As such, the species range has been extended from N. S. W. into Tasmania and New Zealand and a new host association with Leptospermum has been revealed. Cortinarius atroviolaceus and C. kioloensis cannot be distinguished morphologically, but the two are geographically separated and likely ecologically differentiated. Cortinarius atroviolaceus is found at 1700 m on Mt. Kinabalu, Malaysia where Leptospermum (Myrtaceae), Trigonobalanus (Fagales), Quercus (Fagales), Lithocarpus (Fagales), and Castanopsis (Fagales) may occur (Beaman et al. 2000; Beaman and Anderson 2004). Cortinarius kioloensis is found with Eucalyptus (Myrtaceae) and/or Allocasuarina (Casuarinaceae) in southeastern Australia and with Leptospermum (Myrtaceae) in New Zealand. Herbarium collections of C. kioloensis from New Zealand have been misidentified as C. subcalyptrosporus. Cortinarius kioloensis differs from C. subcalyptrosporus by the non-calyptrate basidiospores. It differs from C. atrolazulinus by association with Myrtaceae, not Nothofagaceae. Cortinarius kioloensis is also similar to C. carneipallidus, a Nothofagus association, but differs from this latter species by association with Myrtaceae. Phylogenetic results (Fig. 1) also support these separations. Unique molecular synapomorphies are present at pos. 11, 35, 203 (ITS1), 475, 554, 577, 578, 623 (ITS2) of our alignment. Cortinarius kioloensis was treated as “C. sp. AU1” in Harrower et al. (2015).

Cortinarius palatinus Harrower, sp. nov.

MycoBank No: MycoBank MB 811657
Figs 1, 2b, 3b, 4b

Diagnosis

Similar to Cortinarius neotropicus sp. nov. but differs in having shorter basidiospores and absence of caulocystidia. Unique molecular synapomorphies at pos. 39 (ITS1), 524, 618, 649, 651, 672 (ITS2) of our alignment.

Type

COSTA RICA. San Jose: Perez Zeledon, Villa Mills, CATIE Experimental Forest, 9°33'03"N; 83°40'56"W, 2880 m alt., (Quercus costaricensis), 21 June 2003, R.E.Halling 8411 (holotype: NY 796168).

Etymology

Named for the color palatinate, a shade of violet.

Description

Pileus 30–50 mm wide, convex to plano-convex, surface dry, densely squamulose, appressed toward margin, erect on disc, violet (15C5–8, 15D5–8, 15E5–8, 16C5–8, 16D5–8, 16E5–8), red with KOH. Lamellae adnexed, close, violet, concolorous with pileus, soon assuming brown colors, up to 1 cm broad, edges even to uneven. Stipe 80–120 mm long, 10–14 mm broad, equal or subclavate, strict or curved, dry, upper half violet, fibrillose–striate; lower half pale violet, fibrillose, base violet (15C5–8, 15D5–8, 15E5–8, 16C5–8, 16D5–8, 16E5–8). Universal veil pale violet. Context pale violet, unchanging. Smell mild. Taste mild.

Basidiospores 11–15 µm × 7–9.5 µm, means = 12–13 µm × 8–8.5 µm, Q = 1.39–1.59, Q means = 1.40–1.53 (150 spores, 6 specimens). Amygdaloid, strongly verrucose, plage present under SEM. Basidia 2- and 4-spored, clavate, (20–) 30–40 × 10–15 µm. Cheilocystidia present but not abundant in younger specimens, lageniform, brown or grey in KOH, (35–) 60–95 (–100) µm × (11–) 13–25 (–30) µm. Pleurocystidia abundant, lageniform, brown or grey in KOH, (40–) 50–80 (–105) µm × (10–) 15–25 (–40) µm. Pileipellis a trichoderm composed of hyphae up to 20 µm wide, multi-septate, with brown contents in KOH, terminal hyphae blunt-ended, sometimes aciculate. Clamp connections present.

Ecology and distribution

Known from Perez Zeledon and San Gerardo de Dota in Costa Rica. Associated with Quercus copeyensis, Q. seemannii, and Q. costaricensis on acidic soils. Occurring mid-June to mid-July. Elevation 2220–2280 m.

Other specimens examined

Costa Rica, San Jose, Perez Zeledon, Villa Mills, CATIE Experimental Forest, 9°33'03"N; 83°40'55"W, 2880 m, (Quercus costaricensis), 22 June 1995, R.E.Halling 7450 (NY 79537). San Jose, San Gerardo de Dota, Albergue de Montana, Savegre, ~5 km SW of Cerro de la Muerte, 9°33'02"N; 83°48'27"W, 2500 m, (Quercus copeyensis, Q. seemannii), 20 June 1994, R.E.Halling 7307 (NY 34724). San Jose, Perez Zeledon, Villa Mills, CATIE Experimental Forest, 9°41'56"N; 83°56'31"W, 2850 m, (Quercus costaricensis), 12 June 2000, R.E.Halling 8184 (NY 795933). San Jose, San Gerardo de Dota, Albergue de Montana, Savegre, ~5 km SW of Cerro de la Muerte, 9°33'02"N; 83°48'27"W, 2220 m, (Quercus copeyensis, Q. seemannii), 10 July 2001, R.E.Halling 8004 (NY 460906). San Jose, Perez Zeledon, Villa Mills, CATIE Experimental Forest, (Quercus costaricensis), 23 June 2001, E. & A. Horak (ZT 10422).

Discussion

Cortinarius palatinus sp. nov. is treated as ‘C. sp. CR2’ in Harrower et al. (2015). It occurs in the same oak forests and at the same elevation as Cortinarius neotropicus sp. nov. It can be distinguished from C. neotropicus sp. nov. by its larger and more heavily verrucose basidiospores and pale violet context. Caulocystidia were found in C. violaceus and C. neotropicus sp. nov., but not in C. palatinus sp. nov. Cortinarius atrotomentosus sp. nov. differs from C. palatinus sp. nov. by its wider and olive colored stipe. These two species appear to be most closely related (Fig. 1). Cortinarius atrotomentosus sp. nov. occurs on limestone soil and C. palatinus sp. nov. on acidic soil. This is the same species that was referred to as C. violaceus by Halling and Mueller (2005) from the Talamanca Mountains of Costa Rica (NY795933 illustrated).

Cortinarius altissimus Harrower & T.W. Henkel, sp. nov.

MycoBank No: MycoBank MB 811658
Fig. 1, 2g, 3c, 4c

Diagnosis

Similar to Cortinarius kerrii Singer & I.J.M.Araujo but differs by its larger basidia and less frequent, much larger pleurocystidia. Unique molecular synapomorphies at pos. 46, 108, 156, 212, 216, 259, 260, 261, 262, 264 (ITS1), 440, 503, 506, 532, 545, 554, 583, 617 (ITS2) are present in our alignment.

Type

GUYANA. Region 8 Potaro-Siparuni: Pakaraima Mountains, Upper Potaro River Basin, vicinity of Potaro base camp at 5°18'04"N; 59°54'40"W, 710 m alt., on lateritic soils; 2 km southeast of base camp near Dicymbe plot 1, on humic mat of forest floor under Dicymbe corymbosa, May 25 2001, T.W.Henkel 8211 (holotype: BRG 41220; isotype TENN 069829, HSU G1168)

Etymology

Refers to the exceptionally tall basidiomata of the species, due to the long stipe.

Description

Pileus 24–56 (–87) mm wide, 17–29 mm tall, convex to plano-convex with a low, broad umbo with age, surface dry, erect tomentulose-squamulose throughout, especially over disc, under hand lens squamules subpyramidal and subacuminate, 0.5 mm tall, dark violet (16F7–16F8, 17F7–17F8), lighter concolorous (16E8) near margin, red in KOH; margin entire, broadly undulating with age. Lamellae subclose, shallowly adnexed with short decurrent tooth, subthick, 2 mm broad at margin, 8–10 mm centrally, 7 mm at stipe, concolorous (16F5–17F5), browning with basidiospore development; edges concolorous, hispid under hand lens; lamellulae usually 3 (2 short at 1–3 mm, 1 long at 6–21 mm), occasionally 5, rarely 7. Stipe 132–220(–263) mm long, (2–) 5–11 mm thick at apex, (3–) 7–17 mm thick at center, (7–) 12–24 mm thick at base, subequal, tapering gradually from base to apex, concolorous (16F5–16F6) or slightly lighter concolorous (16D7–16E7, 17D7–17E7) over lower 4/5, apex slightly more greyish violet (15D6–15D7), finely longitudinally striate throughout, with appressed longitudinal fibrils visible under hand lens, cartilaginous, snapping easily, red with KOH. Partial veil cortinate, rather scant, concolorous, minimally retained as scattered, rust-brown fibrils on upper stipe and occasionally pileus margin. Basal mycelium a pale purple (16B3–16B4) matted tomentum. Pileus context subsolid, off white to light purple (17A4–17B4), unchanging, 0.5–1 mm thick at margin, 1–1.5 mm centrally, 6 mm above stipe. Stipe context cartilaginous and concolorous in outer rind, core hollow and off-white, reddening instantly with KOH. Smell mild, slightly fruity to musty. Taste minimal, indistinct.

Basidiospores dark orange–brown (7E7–7E8) in heavy deposit, 10–16 × 8–11 (–13) µm, means = 12–14 × 9.5–11 µm, Q = (1.0–) 1.10–1.44 (–1.56), Q means = 1.18–1.33 (70 spores, 5 specimens), ellipsoid to amygdaloid, verrucose, plage present under SEM. Basidia 4-spored, clavate, (40–) 50–60 (–65) × (8–) 9–15 (–16) µm. Cheilocystidia scattered to abundant, ventricose-rostrate to lageniform, opaque greyish or brown in KOH, (60–) 70–90 (–100) µm × (10–) 20–30 (–40) µm. Pleurocystidia infrequent, ventricose to ventricose–rostrate, grey or rarely brown in KOH, (60–) 70–110 (–125) × (10–) 25–40 (–45) µm. Pileipellis a trichoderm, organized into discrete, suberect fascicles; hyphae (10–) 15–30 (–35) µm wide, (225–) 270–500 (–550) µm high, light brown in KOH, multiseptate; terminal cells undifferentiated, rounded at apex, or occasionally subclavate. Clamp connections present.

Ecology and distribution

Solitary to scattered on humic mat of forest floor in forests dominated by Dicymbe corymbosa (Caesalpinioideae) on lateritic soils; also occurring in forests dominated by Dicymbe altsonii, Aldina insignis (Papilionoideae), and Pakaraimaea dipterocarpacea (Dipterocarpaceae) and Dicymbe jenmanii on white sand soils; known only from the Upper Potaro and Upper Mazaruni River Basins in the Pakaraima Mountains of Guyana, in the central Guiana Shield.

Other specimens examined

Guyana, Region 8 Potaro-Siparuni: Pakaraima Mountains, Upper Potaro River Basin, within a 15 km radius of Potaro base camp at 5°18'04"N; 59°54'40"W, 710 m alt., on lateritic soils; 1 km SE of Potaro base camp on Benny’s Ridge, (Dicymbe corymbosa), 7 July 2003, T.W.Henkel 8539 (BRG; HSU G1169; TENN 069831). ~15 km E of Potaro base in vicinity of Tadang base camp, (Dicymbe altsonii, Aldina insignis), 30 Dec 2009, T.W.Henkel 9180 (BRG; HSU G1170; TENN 069830). 200 m southwest of Tadang base camp (Dicymbe corymbosa, Dicymbe altsonii, Aldina insignis), 6 June 2013, T.W.Henkel 9752 (BRG; HSU G1171). Region 7 Mazaruni-Cuyuni: Pakaraima Mountains, Upper Mazaruni River Basin, within 1 km radius of base camp at 5°26'21"N, 60°04'43"W, ~800 m alt., on white sand soils; 1 km SW of base camp in monodominant stand of Pakaraimaea dipterocarpacea, 25 Dec 2010, T.W.Henkel 9543 (BRG; HSU G1172). Pakaraima Mountains, Upper Mazaruni River Basin, within 1 km radius of base camp at 5°26'21"N, 60°04'43"W, ~800 m alt., on white sand soils; 1 km SW of base camp in monodominant stand of Pakaraimaea dipterocarpacea, 5 June 2012, T.W.Henkel 9690 (BRG; HSU G1173).

Discussion

Cortinarius altissimus was treated as ‘Cortinarius sp. SA1’ in Harrower et al. (2015). The species has an exceptionally long stipe compared to the width of its pileus, and overall one of the longest stipe lengths recorded in the genus. The size of its basidiospores overlaps with that of C. kerrii, which is known from Amazonia. However, the size of the basidia and cystidia are twice that of C. kerrii. Additionally, pleurocystidia are infrequent in C. altissimus sp. nov. whereas they are abundant in C. kerrii. Cortinarius altissimus sp. nov. is a prominent member of the ECM fungal assemblage associated with Dicymbe monodominant forests in Guyana. In a long-term D. corymbosa plot study of Henkel et al. (2012), basidiomata of C. altissimus sp. nov. occurred in 5.2% of 630 quadrats sampled during the May–July rainy seasons over seven years. Phylogenetically (Fig. 1), C. altissimus sp. nov. is most closely related to other taxa of the C. violaceus group that also occur in the Americas.

Cortinarius hallowellensis Wood, 2009

Figs 1, 2d, 3d, 4d

Type

AUSTRALIA. Western Australia: Mount Hallowell Reserve, (Eucalyptus) 22 May 1993 K. Syme (holotype: PERTH005506794).

Description

Pileus 34–90 mm wide, not velvety or velvety smooth, not at all squamulose, greasy when wet, radially innately fibrillose when dry, dark violet (18F4), red in 5% KOH. Lamellae broadly adnexed to adnate, 4–9 mm deep, subdistant, dark violet (16F5), edge smooth, entire, lamellulae abundant in 2 tiers. Stipe 37–80 mm long, 10–30 mm thick, dry, longitudinally fibrillose, cylindrical to clavate with a swollen base, 16F4 (dark violet). Universal veil deep violaceus. Context firm in pileus, fibrous in mature stipe, pallid, tinged purplish (16A2) especially near outside; purple slate/deep purple in stipe and middle of pileus surrounded by speckled white in pileus; yellow-brown at base of stipe. Basal mycelium whitish. Odor not distinctive. Taste none.

Basidiospores (8.5–) 10–13.5 µm × (5–) 6–7.5 µm, means = 10–12 µm × 6–7 µm, Q = 1.55–1.95, Q means = 1.55–1.94, (150 spores, 6 specimens), ellipsoid to subamygdaliform, moderately verrucose, plage barely noticeable under SEM. Basidia 4-spored, clavate, (25–) 30–50 (–65) µm × 9–12 µm. Cheilocystidia abundant, narrowly fusiform, purple or brown in KOH, (50–) 55–95 (–130) µm × 10–20 µm. Pleurocystidia abundant, narrowly fusiform, purple or brown in KOH, (40–) 50–90 (–105) µm × (5–) 10–15 µm. Caulocystidia not observed. Pileipellis an ixocutis, of even thickness throughout. Hyphae 2.5–11.0 µm wide. Clamp connections present.

Ecology and distribution

With Eucalyptus, Corymbia, Melaleuca, Allocasuarina, Agonis, and/or Leptospermum in Western Australia and Tasmania. Fruiting May and June.

Other specimens examined

Australia. Tasmania: Scamander, Winifred Curtis Scamander Reserve, near Boundary Track (Allocasuarina verticillata and Eucalyptus), 8 June 2003, D.A. Ratkowsky (MEL 2350466). Hobart, Peter Murrell Nature Reserve. Site HEG. [55G, 524296, 5258469], (Allocasuarina monilifera, Eucalyptus amygdalina, Leptospermum scoparium), 10 July 2001, S. McMullan-Fischer (MEL 2300544). Western Australia: Two Peoples Bay Nature Reserve, Moates Lake access road (Allocasuarina monilifera, Eucalyptus amygdalina, Leptospermum scoparium), 7 May 1991, K. Syme (PERTH 3978729). Cemetery Road near Walpole-Nornalup National Park (Eucalyptus marginata, Corymbia calophylla, Agonis flexuosa, Agonis parvceps, Agonis hypericifolia, Allocasaurina fraseriana, Melaleuca sp.), 3 May 1992, K. Syme, N.L. Bougher & M. Hart (PERTH 7581696).

Discussion

Cortinarius hallowellensis was treated as ‘Cortinarius sp. AU2’ in Harrower et al. (2015). Based on sequencing the type, we have demonstrated that this species occurs in Tasmania as well as Western Australia. Cortinarius hallowellensis is unique in the C. violaceus group in that it has an ixocutis instead of a trichoderm and is the deepest diverging lineage in the C. violaceus clade (Fig. 1). Macroscopically, the pileus is not at all squamulose. The suprapellis is mucilaginous imparting a greasy appearance when wet. The only other species in the C. violaceus group found in Australia – C. kioloensis is readily distinguished from C. hallowellensis by its dry, tomentulose-squamulose pileus. Cortinarius jenolanensis, C. paraviolaceus and C. austroviolaceus lack a squamulose pileus, but unlike C. hallowellensis, they lack pleurocystidia and/or cheilocystidia. Cortinarius hallowellensis is also referred to as C. violaceus by Bougher and Syme (1998) (PERTH 5506794 illustrated). Unique molecular synapomorphies are found at pos. 63, 100, 101, 123, 132, 148, 163, 164, 180, 228 (ITS1), 462, 532, 540, 546, 577, 602, 611, 614, 643 (ITS2) in our alignment.

Cortinarius neotropicus Harrower, sp. nov.

MycoBank No: MycoBank MB 811660
Figs 1, 2e, 3e, 4e

Diagnosis

Similar to Cortinarius palatinus Harrower sp. nov. but differs by its longer basidiospores and presence of caulocystidia. Unique molecular synapomorphies are found at pos. 58, 161, 200, 205 (ITS1), 467, 566 (ITS2) in our alignment.

Type

COSTA RICA. San Jose: Perez Zeledon, Villa Mills, CATIE Experimental Forest, 9°33'03"N; 83°40'55"W, 2880 m, (Quercus costaricensis), 22 June 1994, R.E.Halling 7330 (holotype: NY 34729).

Etymology

Refers to where it is found: the neotropics.

Description

Pileus 28–88 mm wide, convex to plano-convex, surface dry, not hygrophanous, squamulose to appressed squamulose, (18D3-6, 18E3-6), dull violet to greyish violet, red with KOH. Lamellae subdistant, adnexed to adnate, average thickness, ventricose, dark violet (17F2). Stipe 61–149 mm long, 4–16 mm thick at apex, 7–48 mm thick at base, tapering gradually from base to apex, sometimes clavate, dry, 18(D–E)(3–6), dull violet to grayish violet. Universal veil pale violet. Basal mycelium white. Odor mild. Taste mild.

Basidiospores (11.5–) 13–17 µm × 7–10 µm, means = 13–15 × 8–9 µm, Q = 1.53–1.77, Q means = 1.59–1.78, (200 spores, 8 specimens), ellipsoid to amygdaloid, finely verrucose, plage present under SEM. Basidia 4-spored, clavate, (25–) 30–45 (–55) µm × 10–15 µm. Cheilocystidia abundant, narrowly fusiform to lageniform, brown or grey in KOH, (40–) 50–75 (–95) µm × (10–) 15–20 (–30) µm. Pleurocystidia abundant, narrowly fusiform to lageniform, sometimes subcapitate, brown or grey in KOH, (35–) 50–90 (–120) µm × 10–25 (–40) µm. Caulocystidia present. Pileipellis a trichoderm, composed of interwoven smooth hyphae, these 9–20 µm wide, 265–415 µm high, multiseptate, with brown contents in KOH, terminal hyphae mostly blunt-ended, some lanceolate. Clamp connections present.

Ecology and distribution

Known from Costa Rica and Colombia. With Quercus copeyensis, Q. seemannii, and Q. costaricensis in Costa Rica and with Q. humboldtii and Trigobalanus in Colombia. 2200–2880 m alt. Fruiting late May to the end of August in Costa Rica and November in Colombia.

Other specimens examined

Colombia. Huila: Finca Merenberg, (Quercus humboldtii), 21 May 1987, R.E.Halling 5284 (NY). Valle del Cauca: Municipio de Pance, Parque Nacional Los Farallones, above El Topacio, (Trigonobalanus), 17 Nov. 1988, A.E.Franco-Molano 151 (NY). Costa Rica. San Jose: Perez Zeledon, Villa Mills, CATIE Experimental Forest, 9°33'03"N 83°40'60 W (Quercus costaricensis), 30 June 1998, R.E.Halling 7787 (NY 75934). Perez Zeledon, Villa Mills, CATIE Experimental Forest, 9°33'03"N; 83°40'60"W, 2840 m, (Quercus costaricensis), 27 June 2000, R.E.Halling 7946 (NY 460484). San Gerardo de Dota, Albergue de Montana, Savegre, ~5 km SW of Cerro de la Muerte, 9°33'02"N; 83°48'27"W, 2200 m alt., (Quercus copeyensis and Q. seemannii), 9 June 2001, R.E.Halling 8154 (NY 795935). San Gerardo de Dota, Albergue de Montana, Savegre, ~5 km SW of Cerro de la Muerte, 9°33'02"N; 83°48'27"W, 2500 m alt., (Quercus copeyensis and Q. seemannii), 8 June 1997, R.E.Halling 7709 (NY 181476). La Guaria, 9°35'60"N; 83°58'60"W, 2300 m alt., (Quercus copeyensis and Q. seemannii), 21 Aug. 1995, A.E.Franco-Molano 1393 (NY).

Discussion

Cortinarius neotropicus sp. nov. was treated as ‘Cortinarius sp. CR1’ in Harrower et al. (2015). This species can be distinguished from C. palatinus sp. nov., with which it co-occurs in Costa Rica, by its smaller and finely verrucose basidiospores. In addition, C. neotropicus sp. nov. and C. violaceus both have caulocystidia, whereas other species in the C. violaceus group (such as C. palatinus sp. nov. and C. atrotomentosus sp. nov.) do not. Colors of the basal mycelium and context have not been recorded. There are no microscopic characters that can be used to distinguish it from C. violaceus, and the two species are supported as sister lineages (Fig. 1).

Cortinarius carneipallidus Harrower & E. Horak, sp. nov.

MycoBank No: MycoBank MB 811661
Figs 1, 2f, 3f, 4f

Diagnosis

Similar to Cortinarius kioloensis but differs by its association with Nothofagus. Unique molecular synapomorhies at pos. 33, 157, 170, 190, 233, 249 (ITS1), 472, 480, 489, 525, 607, 670 (ITS2).

Type

NEW ZEALAND. Ruapehu: Tongariro National Park, Mt. Ruapehu, Blyth Track, (Nothofagus solandri) 21 April 2009, J.A. Cooper (holotype: PDD 95444).

Etymology

Means ‘pale flesh’, referring to the relative lack of purple pigment in the center context of this species compared to C. violaceus, C. atrotomentosus, C. atrolazulinus, C. atroviolaceus and C. subcalyptrosporus.

Description

Pileus 49–100 mm wide, surface dry, convex then plano–convex, tomentose to fine scaly, dark violet (16D5–8), paler at margin, red in KOH. Lamellae crowded, emarginate-adnexed, average thickness, broad, deep violet. Stipe 96–175 mm long, 5–11 mm thick at apex, 12–30 mm thick at base, subequal, sometimes slightly bulbous, tapering gradually from base to apex, pale violet (17C3–6), covered in copious veil material. Universal veil pale violet (same color as stipe), leaving bands on the stipe. Basal mycelium pale violet. Context white flecked with violet in pileus and lower half of stipe, upper half of stipe violet flecked with white, sometimes white with violet cortex, younger specimens violet flecked with white and only fully white in the base.

Basidiospores (9–) 10–12 (–13) µm × (7–) 8–9 (–9.5) µm, means = 10.5–12 µm × 8–8.5 µm, Q = 1.19–1.64, Q means = 1.30–1.45, (224 spores, 8 specimens). Ellipsoid to amygdaloid, strongly verrucose, plage present. Basidia 4 spored, clavate, (35–) 40–50 (–55) µm × (9–) 10–12 (–13) µm. Cheilocystidia abundant, narrowly fusiform to lageniform, brown in KOH, 60–90 (–100) µm × 15–20 (–25) µm. Pleurocystidia abundant, narrowly fusiform to lageniform, sometimes subcapitate, brown in KOH, (40–) 55–80 (–100) µm × (10–) 15–25 (–30) µm. Caulocystidia not seen. Pileipellis a trichoderm, composed of smooth hyphae 10–25 µm wide, 350–370 µm long, multiseptate, brown content in KOH, terminal cells bottle-shaped, clavate, lanceolate or ending bluntly. Clamp connections present.

Ecology and distribution

Under Nothofagus in New Zealand. Fruiting April to June.

Other specimens examined

New Zealand. Wellington: Kapiti Coast, Tararua Forest Park (Nothofagus), 16 May 2009, K. Hosaka (PDD 98057). Canterbury: Selwyn Klondyke Corner (Nothofagus cliffortioides), 5 June 2010, P. White (PDD 95823). Waikato: Taupo, Erua Forest National Park, Erua Forest rest area (Nothofagus), 8 Apr. 2005, L. Fischer (PDD 82693). Taupo, Mt. Ruapehu, Tongariro National Park, Whakapapanui Track, Buller 28 Apr. 2001, E. Horak (PDD 72636). West Coast: Buller, Springs Junction, Lake Christabel Track, 14 Apr. 2000, E. Horak (PDD 71219). Buller, Reefton, Victoria Forest Park, Tawhai Walk, (Nothofagus) 13 May 2006, E. Horak (PDD 88638). Buller, South Charleston, Tiropahi Track (Nothofagus) 8 May 2010, A. Roberts (PDD 88995).

Discussion

Cortinarius carneipallidus sp. nov. differs from C. subcalyptrosporus by the absence of calyptrate basidiospores. The association with Nothofagus distinguishes this species ecologically from C. atroviolaceus, which likely associates with members of the Myrtaceae and/or Fagaceae in Malaysia. The context is hygrophanous purple in young specimens. In older specimens, it is hygrophanous grey or white in the pileus and base and only purple at the stipe apex. The only other species described from Indonesia or Australasia to have whitish flesh is C. kioloensis. It differs from C. kioloensis, with which it forms a sister group relationship (Fig. 1), by its association with Nothofagus. C. carneipallidus has a wider pileus and longer stipe than C. atrolazulinus. The width of the basidiospores is wider than those in C. atrolazulinus as well.

Cortinarius carneipallidus sp. nov. has been variously identified as C. atroviolaceus, C. atrolazulinus, C. subcalyptrosporus, C. violaceus and Cortinarius sp. The species was treated as ‘C. violaceus sensu Moser 1986’ in Harrower et al. (2015) but differs from north temperate C. violaceus by the absence of caulocystidia and association with Nothofagus. However, the name ‘C. violaceus sensu Moser 1986’ was applied based on interpretation of the protologue only, and as no specimens of ‘C. violaceus sensu Moser 1986’ were examined, we do not intend to synonymize the former with C. carneipallidus.

Cortinarius atrotomentosus Harrower, sp. nov.

MycoBank No: MycoBank MB 811662
Figs 1, 2c, 3g, 4g

Diagnosis

Similar to Cortinarius violaceus (L.: Fr.) Gray but differs by having smaller spores and shorter basidia as well as an absence of caulocystidia. Unique molecular synapomorphies at pos. 71 (ITS1), and 606 (ITS2) are present in our alignment.

Type

USA, Florida: Wakulla Co., Crawfordville, Apalachicola National Forest (30°12'06"N; 84°26'33"W), on soil under Quercus virginiana, 4 Dec. 2010, TFB 13848, (holotype: TENN 065527).

Etymology

Meaning ’dark-tomentose’ in reference to the dark coloration of the fruiting body.

Description

Pileus 26–91 mm wide, surface dry, tomentose to fine scaly, dark violet (17F3) to dark brown in age (7F3), red in KOH. Lamellae adnexed, not sinuate, close to subdistant, thin, ventricose, dark violet (17F3). Stipe 75–131 mm long, 8–18 mm thick at apex, 17–26 mm thick at base, ventricose, silky-glabrous, olive brown (4E6) to brownish grey in age (5D2). Basal mycelium lilac (16B4) to greyish magenta (14D3). Context purple grey (13E2) to reddish brown (10D4). Smell mild. Taste none. Basidiospores (9–) 10.5–13 (–14.0) µm × (6–) 7–8 µm, means = 10.5–12 µm × 6–8 µm, Q = 1.43–1.67, Q means = 1.50–1.53, (70 spores, three specimens), ellipsoid to amygdaloid, strongly verrucose, plage present under SEM. Basidia 4-spored, clavate, 20–30 × 10–12 µm. Cheilocystidia abundant, narrowly fusiform to lageniform, brown in KOH, 50–70 (–80) µm × 15–25 µm. Pleurocystidia abundant, narrowly fusiform to lageniform, brown in KOH, 43–65 (–70) × 15–20 µm. Caulocystidia not seen. Pileipellis a trichoderm, hyphae 6–20 µm wide, 185–370 µm high, multiseptate, brown content in KOH, terminal hyphae mainly blunt-ended, some lanceolate or aciculate. Clamp connections present.

Ecology and distribution

Under Quercus virginiana. Known only from Apalachicola National Forest, Crawfordville, Florida, USA. Fruiting early December.

Other specimens examined

USA. Florida: Wakulla Co., Crawfordville, Apalachicola National Forest (30°12'07"N; 84°26'33"W), (Quercus virginiana), 2 Dec. 2010, D.Lewis & B.Petty TFB 13840 (TENN 065527).

Discussion

Cortinarius atrotomentosus sp. nov. was treated as ‘Cortinarius sp. NA1’ in Harrower et al. (2015). This species can be differentiated from C. violaceus by its dark violet to dark brown pileus and the olive brown to brownish grey stipe. The context of C. violaceus is violet to greyish, not mauve as is C. atrotomentosus sp. nov. Caulocystidia were not found on this species whereas they are present in C. neotropicus sp. nov. and C. violaceus. The current Gulf Coast geographic distribution of C. atrotomentosus sp. nov., where it associates with Quercus, does not overlap with the more northerly distribution of C. violaceus. The species differs from C. palatinus sp. nov., with which it appears to be most closely related (Fig. 1), by its location (Gulf Coast region) and its dark exterior. The basal mycelium is the only part of the fruiting body that could be described as pale violet.

Species inquirendae

Cortinarius hercynicus (Pers.) M.M.Moser 1967

Cortinarius hercynicus differs from C. violaceus s.s. by having smaller and broader basidiospores and occurrence in coniferous forests. Harrower et al. (2015) did not find a molecular difference between C. hercynicus and C. violaceus, but taxon sampling of C. hercynicus was not adequate. Brandrud (1983) treated C. hercynicus as a variety of C. violaceus. However, in Harrower et al. (2015) samples of C. violaceus s.l. recorded from coniferous forests and separately in hardwood forests failed to form monophyletic groups suggesting that host association is of no taxonomic significance. Thus, current data suggest C. hercynicus is synonymous with C. violaceus.

Species excludendae

Cortinarius austroviolaceus Gasparini 2001

Fig. 2i, 3i, 4i

This species is phylogenetically placed outside of Cortinarius sect. Cortinarius (Harrower et al. 2015). In comparison to species within the section, the basidiospores lack a plage (Fig. 3i), the basidiospore ornamentation differs (less coarsely ornamented) at the SEM level, and pleurocystidia are absent.

Cortinarius gayi E. Horak 1980

Fig. 2h

The purple color and densely squamulose or fibrillose pileus is consistent with most other species in sect. Cortinarius. However, C. gayi lacks both pleurocystidia and cheilocystidia, and a plage is reportedly absent from the basidiospores. This species may have closer to affinities to C. austroviolaceus than sect. Cortinarius.

Cortinarius paraviolaceus M.M.Moser 1987

This species is described as having a pileus that is “rather intensely violaceus…the center becoming dull tawny ochraceus” (Moser 1987). The stipe is described as “violaceus tomentose at the base, otherwise brownish fibrillose with the cortina”. Pleurocystidia are rare and no cheilocystidia were observed (Moser 1987). The pileus is “not strongly squarrose-fibrillose-squamulose”. These characters, taken together, are not consistent with the C. violaceus group.

Cortinarius jenolanensis Wood 2009

This species has a smooth pileus and cheilocystidia, but lacks pleurocystidia. This species may have closer to affinities to C. austroviolaceus than sect. Cortinarius.

Cortinarius atrolazulinus M.M. Moser sensu Soop 2008

Known collections (not examined by the authors): New Zealand. Southland: Southland, Fiordland, Lake Gunn Track, (Nothofagus), 5 May 2001, K. Soop KS CO1223 (PDD103879; GenBank KF727372; KF727328). Taupo: Taupo, Kaimanawa Forest Park, Cascade Hut Track, (Nothofagus) 9 May 2009 K. Soop KS CO1917 (PDD97542).

C. atrolazulinus was described by Moser (1987) from Nothofagus forest in New Zealand. These two collections match Moser’s description well, which includes the presence of cheilocystidia and pleurocystidia. Phylogenetic analyses (not shown) place it in an unknown clade outside of Cortinarius sect. Cortinarius. Further morphological and genetic analyses are needed to correctly classify this species. Attempts to sequence the type collection of C. atrolazulinus (ZT 69-276) were unsuccessful.

Cortinarius atroviolaceus M.M. Moser sensu Shirley

New Zealand: Auckland, Kauri Point Reserve. (Leptospermum) C. Shirley AK 369 (PDD; GenBank KT444633; KT444634).

Pileus fibrillose, not tomentose. Epicutis not a trichoderm, hyphae 6-14 µm wide, blue extracellular pigment clumps. Purple intracellular pigment readily dissolving in KOH. Spores 7.5-8.5 x 4-5 µm. Cheilocystidia and pleurocystidia present. This species shares 90% ITS sequence identity with one member of Cortinarius sect. Cortinarius (C. carneipallidus) at best. It shows higher sequence identity to members outside of the section. Phylogenetic analyses (not shown) could not place it within the section with confidence. Cortinarius jenolanensis has wider spores and lacks pleurocystidia, compared to this species. Cortinarius atroviolaceus M.M. Moser has much longer and wider spores. As there is only one collection of this species, with inadequate documentation, more work needs to be done to document and classify this species.

Acknowledgements

We would like to thank the TENN, HSC, NY, PDD, FRIM, PERTH, ZT and MEL herbaria and staff for loans of collections. This research was funded by a Postgraduate-Doctoral Scholarship provided by the National Science and Engineering Research Council of Canada to E. Harrower, Chancellor’s Funds from the University of Tennessee, and U.S. National Science Foundation award DEB-1354802 to M.E. Smith and P.B. Matheny. Collection of Guyana specimens was supported by NSF DEB-0918591 and National Geographic Society’s Committee for Research and Exploration grants to T.W. Henkel. Collection of Costa Rican material was supported by NSF DEB-9972018 and NGS-7341-02 grants to Roy E. Halling. We thank Clive Shirley and Ian Dodd for also providing material. We acknowledge John Dunlap for training and assistance with use of SEM. We also thank Jerry Cooper, Karl Soop and Roy Halling for their correspondence. D. and P. Lewis and B. Petty were helpful in aiding in the recollection of C. atrotomentosus. A. and A. Bessette generously photographed C. atrotomentosus for this publication. Dr. Jeannine Cavender-Bares confirmed the identification of Quercus virginiana as the host for Cortinarius atrotomentosus.

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