Cortinarius sect. Brunnei ( Basidiomycota, Agaricales) in North Europe

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Cortinarius sect. Brunnei (Basidiomycota, Agaricales)in North Europe

Tuula NISKANENa,*, Ilkka KYTOVUORIb, Kare LIIMATAINENa

aDepartment of Biological and Environmental Sciences, Plant Biology, P.O. Box 65, FI-00014 University of Helsinki, FinlandbBotanical Museum, P.O. Box 7, FI-00014 University of Helsinki, Finland

a r t i c l e i n f o

Article history:

Received 23 January 2008

Received in revised form

12 September 2008

Accepted 10 October 2008

Corresponding Editor: Roy Halling

Keywords:

Barcoding

ITS

Mr Bayes

Taxonomy

TNT

a b s t r a c t

The section Brunnei was extensively studied based on material from North Europe. To sta-

bilise the nomenclature we studied the relevant types of taxa included in this section. Phy-

logenetic relationships and species limits were investigated using rDNA ITS sequences and

the results were compared with the morphological data. We recognised 11 species: Corti-

narius brunneus, C. clarobrunneus comb. nov., C. coleoptera, C. ectypus, C. gentilis, C. glandicolor

(neotypified), C. pseudorubricosus, and four species described as new C. caesiobrunneus, C. al-

bogaudis, C. carabus, and C. cicindela. They are described here and their taxonomy, ecology,

distribution, and relationships are discussed. In addition, a key to species of the section

Brunnei is provided. A total of 77 new sequences of 11 species are published including

nine type sequences. Also the taxonomic assignments of sequences in the public databases

belonging to the section Brunnei are revised.

ª 2008 The British Mycological Society. Published by Elsevier Ltd. All rights reserved.

Introduction

Cortinarius is the most species-rich mycorrhizal fungus genus

in the world. Recent molecular analyses, based on sequence

data from ITS1 and ITS2 and/or the adjacent LSU region,

have shown that Cortinarius itself is monophyletic, but many

of the traditional infrageneric groups are artificial (Garnica

et al. 2003; Garnica et al. 2005; Høiland & Holst-Jensen 2000;

Liu et al. 1997; Peintner et al. 2004; Seidl 2000). Two of these

studies included the type species of the section Brunnei,

namely C. brunneus, and based on the phylogenies in these

moleculal studies, the section Brunnei belongs to the clade

Telamonia (Høiland & Holst-Jensen 2000; Peintner et al. 2004).

This clade comprises most of the species traditionally classi-

fied in Telamonia and is from here onwards refered as Telamo-

nia s. str. The clade is characterised at the molecular level by

one or two indels in the ITS1 region (Garnica et al. 2005;

Høiland & Holst-Jensen 2000; Peintner et al. 2004).

Using morphological studies, the section Brunnei (Melot

1989) has been treated in many different ways (Brandrud

et al. 1998; Consiglio et al. 2006; Kuhner & Romagnesi 1953;

Moenne-Loccoz et al. 1990). Kuhner & Romagnesi (1953) intro-

duced the name Brunnei and included ten species into the

group (Table 1). Of these, only C. brunneus and C. glandicolor

are included in the later classifications too. Melot (1989) vali-

dated the name of the section, but only the type species

C. brunneus was mentioned in his paper. In the three current

ongoing Cortinarius flora projects, two to six taxa are included

in the section Brunnei (Brandrud et al. 1998; Consiglio et al.

2006; Moenne-Loccoz et al. 1990). C. brunneus is common to

all classifications and C. ectypus is included in two of the

treatments, otherwise the classifications differ.

* Corresponding author.E-mail address: [email protected]

j ourna l homepage : www.e lsev ier . com/ loca te /mycres

m y c o l o g i c a l r e s e a r c h 1 1 3 ( 2 0 0 9 ) 1 8 2 – 2 0 6

0953-7562/$ – see front matter ª 2008 The British Mycological Society. Published by Elsevier Ltd. All rights reserved.doi:10.1016/j.mycres.2008.10.006

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https://www.researchgate.net/publication/280806920_Phylogenetic_relationships_within_Cortinarius_subgenus_Myxacium_sections_Defibulati_and_Myxacium?el=1_x_8&enrichId=rgreq-b229fbe0-1497-449b-bb56-a43073a66aa2&enrichSource=Y292ZXJQYWdlOzIzNDkxNzY3O0FTOjE5MTM1OTk2NzEyOTYwMUAxNDIyNjM1MTgwMDA0


No phylogenetic studies of the Cortinarius section Brunnei

have hitherto been undertaken. Based on our unpublished

phylogenetic tree including ITS sequences of ca 200 telamo-

nioid species, our delimitation of this section includes at least

15 species in Europe. Of these, 11 are also morphologically

well-delimited species and presented here. Our delimitation

includes all the four Brunnei species so far classified in this sec-

tion by Moenne-Loccoz et al. (1990). However, C. rheubarbari-

nus, C. sordidemaculatus (syn. C. furvolaesus), and C. anisatus,

previously included in this section by Brandrud et al. (1998)

and Consiglio et al. (2006), do not belong to Brunnei. Previous

molecular phylogenetic studies of the genus Cortinarius indi-

cated that C. gentilis is related to C. brunneus (Høiland &

Holst-Jensen 2000; Peintner et al. 2004). Our preliminary stud-

ies supported these results and C. gentilis was also included in

this study.

The delimitation of the fungal species is controversial and

many different species concepts have been proposed (e.g.

Kuyper 1988; Taylor et al. 2000). The dominant species concept

used in Cortinarius has been the morphological species con-

cept (MSC). It has been applied in varying ways, resulting in

different opinions about the number of species (e.g. in

Cortinarius compare Brandrud et al. 1989, 1992, 1994, 1998;

Bidaud et al. 1995, 1997).

Sequence data in combination with an evaluation of mor-

phological data have been used in only a few species level

studies in Cortinarius, for example in subgenus Phlegmacium:

Frøslev et al. (2006), and Moser & Peintner (2002a, 2002b), in

subgenus Telamonia: Ammirati et al. (2007), Kytovuori et al.

(2005), Lindstrom et al. (2008), Matheny & Ammirati (2006),

and Niskanen et al. (2006). Frøslev et al. (2007) undertook the

most thorough study; they analysed a total of 421 ITS se-

quences of 79 species of sect. Calochroi (subgenus Phlegma-

cium). These studies are based on the ITS regions, which

have successfully been used for species level studies in

many other genera of fungi, e.g. Hygrophorus (Larsson &

Jacobsson 2004), and Macrolepiota (Vellinga et al. 2003). In Corti-

narius, the intraspecific genetic variation of ITS regions is usu-

ally low and specimens from different continents might have

identical ITS sequences (e.g. Moser & Peintner 2002a).

Only one multi-gene study has so far been done in Cortinar-

ius by Frøslev et al. (2005). They showed that the species level

results from the RNA polymerase II genes, RPB1 and RPB2, are

in concordance with the results from the ITS, and in Frøslev

et al. (2007) it was concluded that the ITS seems to be an appro-

priate marker for species level identification in Cortinarius. In

only two cases, the morphologically accepted species C. atrovi-

rens/C. ionocholorus and C. xanthophyllus/C. claroflavus were

found to have identical ITS sequences, the former similarity

also noticed by Garnica et al. (2005). In other groups of fungi,

ITS sequences have sometimes been found to be too variable,

e.g. Leccinum (Den Bakker et al. 2004), and Xerocomus (Peintner

et al. 2003), or too conserved, e.g. Hebeloma (Aanen et al. 2000),

for species level studies.

Here we describe seven known species and present four

new species of Cortinarius, subgenus Telamonia s str., section

Brunnei based on morphology and molecular data. We also

propose a neotype for C. glandicolor and elevate C. clarobrun-

neus to a species level. The new nomenclature was first pre-

sented in Niskanen et al. (2007), but was not effectively

published, as that paper is an independent, non-serial work

being part of a thesis (Niskanen 2007), being called a ‘manu-

script’ (see ICBN, Art. 30.5). The rest of the species recognised

in the Nordic countries in our preliminary study require fur-

ther studies and will be presented later. Several (3 to 12) sam-

ples per species were sequenced to confirm correct species

delimitation, and a phylogenetic tree based on these ITS se-

quences is provided. If they were available, we studied type

material from all included species to clarify the taxonomy

and nomenclature of Cortinarius. The sequences deposited in

Table 1 – European Cortinarius species classified in the section Brunnei by European authors

Classification proposedin this study

Moenne-Loccozet al. (1990)

Brandrud et al.(1989, 1992, 1994, 1998)

Consiglio et al.(2006)

Kuhner & Romagnesi(1953)

Cortinarius brunneus C. brunneus (syn.

C. subtigrinus?)

C. brunneus var. brunneus C. brunneus C. brunneus

C. glandicolor C. brunneus var. glandicolor C. glandicolor

C. caesiobrunneus

C. clarobrunneus C. brunneus var. clarobrunneus

C. pseudorubricosus C. pseudorubricosus

C. albogaudis

C. ectypus C. ectypus C. ectypus

C. coleoptera

C. carabus

C. cicindela

C. gentilis

C. rheubarbarinus C. anisatus C. bovinus

C. furvolaesus C. punctatus

C. biformis

C. adalbertii

C. stemmatus

C. rigidus

C. fusisporus

C. striaepilus

Bold names indicate species included in the section Brunnei in this study.

Cortinarius sect. Brunnei in North Europe 183


the public databases belonging to the section Brunnei were also

revised. In this study we have applied the morpho-genetic

species concept used in Kytovuori et al. (2005) and Frøslev

et al. (2006) where species are delimited based on morpholog-

ical and molecular data. This work is a part of our larger study

on the boreal Telamonia species.

Materials and methods

Materials

A total of ca 230 specimens were studied. Most collections

were gathered by us from different parts of Fennoscandia

and some from Estonia and Central Europe (deposited in H

and TUR). Additionally some material of H, TUR, and UPS

were included. The types of Cortinarius brunneus, C. brunneus

var. clarobrunneus, C. coleoptera, C. ectypus, and C. pseudorubrico-

sus were studied. The acronyms follow those used by Holmg-

ren et al. (1990). Collectors are abbreviated by T.N., I.K., K.L.,

and P.K., in collections made by the authors and Pirjo

Kytovuori.

Several collections of the studied species (a total of 73 spec-

imens) from different geographical areas (Table 2, marked

with superscript ‘D’ in the lists of specimens examined) and

type collections were sequenced. In addition, one sequence

previously published by us was included in the study. All the

species were represented by three to 12 sequences.

Morphological studies

Macroscopic characteristics were observed from fresh fruit

bodies and several collections were used for the descriptions.

Some representative collections were also photographed in

fresh condition. Colour codes were not used, instead photo-

graphs of the new species C. albogaudis, C. caesiobrunneus,

C. carabus, and C. cicindela, as well as that of the lesser known

C. pseudorubricosus and the neotype of C. glandicolor, are

provided.

Microscopic characteristics were observed from dried

material mounted in Melzer’s reagent (MLZ) and compared

with observation made from dried material mounted in 5 %

potassium hydroxide (KOH). Unless otherwise stated, the

colours reported for various structures are those observed in

MLZ. Measurements were made from Melzer’s preparations

with a�100 oil immersion lens. Twenty spores, from collections

marked with a superscript ‘S’ in the list of examined specimens,

were measured from one fruit body, from the top of stipe or veil.

Length and breadth were measured from the same spore, and

the length:breadth ratios (Q-value) were calculated for individ-

ual spores. The extreme measurements were excluded from

the final values. The hyphae of the lamellar trama, basidia,

and pileipellis were examined as in Kytovuori et al. (2005). In

addition, the pileipellis structure was studied from radial

freehand sections from midway to the pileus centre.

Molecular analyses

Total DNA was extracted from a few milligrams of dried mate-

rial (a piece of lamella) using the NucleoSpin Plant kit

(Macherey-Nagel). The primers ITS 1F and ITS 4 (Gardes &

Bruns 1993; White et al. 1990) were used to amplify the ITS

regions of the rDNA. For problematic material the primer com-

binations ITS 1F/ITS 2 and ITS 3/ITS 4 were also used. PCR am-

plifications were performed in a 25 ml reaction mix with about

70 ng extracted DNA, 1 U Phusion High-Fidelity DNA polymer-

ase and 1�HF buffer (Finnzymes), 200 mM of each dNTP and

0.4 mM of each primer. The PCR reactions were run on a MBS

0.2 G Thermal Cycler (Thermo Hybaid) with the following set-

tings: denaturation for 30 s at 98 �C, followed by 35 cycles of

denaturation for 10 s at 98 �C, annealing for 30 s at 50 �C, and

extension for 30 s at 72 �C. The PCR products were purified

using an ExoSAP-IT purification kit (Amersham Biosciences).

Sequencing was performed on both strands using a BigDye

Terminator v1.1 Sequencing kit (Applied Biosystems). Reac-

tions were performed in 10 ml with 1 ml of PCR product,

1.3 mM of primer (ITS 1F or ITS 4), 1 ml 5� sequencing buffer,

and 1 ml of Terminator Ready Reaction Mix. Reactions were

run for 1 min at 96 �C, followed by 30 cycles of 30 s at 96 �C,

15 s at 50 �C, and 4 min at 60 �C. Unincorporated dye termina-

tors and primers were removed by Sephadex G-50 DNA Grade

Fine (Amersham Biosciences) purification system and the re-

actions were analysed by a MegaBace (Amersham Biosci-

ences) automatic sequencer. Sequences were assembled and

edited using Sequencher 4.1 (Gene Codes, Ann Arbor, MI).

Intragenomic polymorphism was observed as mixed peaks

in chromatographic data. Base polymorphisms are marked

with ambiguous IUB codes and length polymorphisms with

N (further information is provided upon request). Intrage-

nomic variation with Cortinarius has previously been reported

by Frøslev et al. (2007). The sequences were compared with the

material in the public databases (GenBank: http://

www.ncbi.nlm.nih.gov/ and UNITE: http://unite.ut.ee/) using

BLAST searches.

In order to understand the closeness of the studied se-

quences, a sequence of every species was compared with all

the other sequences of the Brunnei species using BLAST

search. The sequences of the studied species and the se-

quences of the closest ones were then aligned using the Clus-

talW 1.8 program (Thompson et al. 1994) on the European

Bioinformatics Institute server (http://www.ebi.ac.uk/

clustalw/index.html). The following differences in the species

were counted from the alignments: (1) the observed number of

variable sites showing how many sites contained infraspecific

and/or -genomic polymorphisms, and (2) the maximum pair-

wise distance informing the biggest pairwise distance

observed between the two most distant sequences within

the species (differences due to intragenomic polymorphisms

are not counted). In Cortinarius this value is usually very low.

The differences between the closest species were also counted

as minimum evolutionary events including indels, transitions

and transversions, and e.g. multiple base indels were treated

as one change. If the amount of differences was 15 or less

the evolutionary events were presented in more detail in pa-

renthesis. Only differences shared by all the specimens of

the same species were counted. The results are presented in

point ‘ITS regions’ under every species.

For the preliminary analyses [POY (Gladstein & Wheeler

2001; Wheeler 1996), result not shown] 200 of our own unpub-

lished ITS sequences covering all the major clades of

184 T. Niskanen et al.

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Table 2 – Specimens used in the DNA study

Species Voucher Herb. Locality Sequenceno.

GenBankaccessionnumber

Cortinarius pseudorubricosus (holotype) GK16165 G France, Foret de Belval 696-05 EU266709

C. pseudorubricosus TN03-944 H Sweden, Jmt, Berg 379-04 EU266705

C. pseudorubricosus IK00-014 H Norway, S&F, Luster 499-04 EU266707

C. pseudorubricosus IK01-022 H Finland, EnL, Enontekio 319-03 EU266706

C. pseudorubricosus IK99-708 H Finland, PH, Virrat 360-04 EU266708

C. albogaudis (holotype) IK98-1233 H Finland, PeP, Runteli 014-02 EU266635

C. albogaudis TN02-1089 H Finland, Ks, Kuusamo 278-03 EU266633

C. albogaudis IK97-1545 H Finland, Kn, Puolanka 1163-07 EU266634

C. ectypus (holotype) GK13318 G Switzerland, foret de Sauaidas 705-05 EU266689

C. ectypus CFP710 S Sweden, Jmt, Undersaker 554-04 EU266688

C. ectypus IK97-080 H Finland, U, Espoo 304-03 EU266685

C. ectypus IK00-011 H Finland, EH, Virrat 315-03 EU266687

C. ectypus IK99-085 H Finland, Kn, Kuhmo 314-03 EU266686

C. gentilis (Fr.) Fr. CFP178 S Norway, Oppl, Dokka 544-04 EU266692

C. gentilis IK99-396 H Finland, PeP, Rovaniemi 085-03 EU266691

C. gentilis TN06-250 H Finland, St, Sakyla 1278-07 EU266690

C. brunneus (Pers. : Fr.)Fr. var. brunneus (neotype) CFP587 S Sweden, Ang, Sabra 557-04 DQ117927.2

C. brunneus var. brunneus IK94-1429 H France, Languedoc-Roussillon, Lozere 306-03 EU266640

C. brunneus var. brunneus TN03-1708 H Slovakia, Tatry Mts, Podbaske 286-03 EU266636

C. brunneus var. brunneus IK96-542 H Finland, PH, Virrat 305-03 EU266641

C. brunneus var. brunneus IK00-012 H Finland, St, Aetsa 317-03 EU266642

C. brunneus var. brunneus TN04-929 H Finland, U, Kirkkonummi 914-05 EU266637

C. brunneus var. brunneus TN04-932 H Finland, U, Kirkkonummi 919-05 EU266638

C. brunneus var. brunneus TN05-018 H Finland, ES, Joutsa 948-06 EU266639

C. glandicolor (Fr. : Fr.) (neotype) TN06-247 H Finland, St, Sakyla 1187-07 EU266700

C. glandicolor CFP615a S Sweden, Jmt, Hallesjo 545-04 EU266704

C. glandicolor TN04-931 H Finland, U, Kirkkonummi 918-05 EU266699

C. glandicolor TN04-914 H Finland, V, Vihti 1130-06 EU266698

C. glandicolor TN04-809 H Finland, U, Sipoo 1119-06 EU266697

C. glandicolor TN04-803 H Finland, U, Sipoo 1131-06 EU266696

C. glandicolor TN04-586 H Finland, PeP, Pisavaara 916-05 EU266695

C. glandicolor TN02-991 H Finland, Ks, Kuusamo 308-03 EU266694

C. glandicolor TN02-989 H Finland, Ks, Kuusamo 318-03 EU266693

C. glandicolor IK94-557 H Finland, EK, Anjalankoski 303-03 EU266701

C. glandicolor IK97-1750 H Finland, PK, Ilomantsi 310-03 EU266702

C. glandicolor JV2953 TUR Sweden, Sm, Hyltebruk 1279-07 EU266703

C. caesiobrunneus (holotype) TN05-127 H Finland, Kn, Suomussalmi 976-06 EU266653

C. caesiobrunneus TN05-094 H Finland, Kn, Puolanka 1271-07 EU266645

C. caesiobrunneus TN05-041 H Finland, EH, Juupajoki 1269-07 EU266644

C. caesiobrunneus TN04-621 H Finland, PeP, Tornio 1270-07 EU266643

C. caesiobrunneus CFP615b S Sweden, Jmt, Hallesjo 546-04 EU266652

C. caesiobrunneus IK02-019 H Finland, PH, Saarijarvi 512-04 EU266651

C. caesiobrunneus IK00-013 H Sweden, Dlr, Idre 302-03 EU266650

C. caesiobrunneus IK99-706a H Finland, Kn, Kuhmo 300-03 EU266648

C. caesiobrunneus IK99-706b H Finland, Kn, Kuhmo 301-03 EU266649

C. caesiobrunneus IK97-1970 H Estonia, Vorumaa, Vatseliina 295-03 EU266647

C. caesiobrunneus IK97-1793 H Finland, PK, Ilomantsi 307-03 EU266646

C. clarobrunneus (holotype) CFP458 S Sweden, Ang, Graninge 353-04 EU266679

C. clarobrunneus CFP1442 S Sweden, Hrj, Hede 354-04 EU266678

C. clarobrunneus IK95-1584 H Finland, EH, Orivesi 321-03 EU266676

C. clarobrunneus IK97-434 H Sweden, Ang, Solleftea 324-03 EU266677

C. clarobrunneus TN02-591 H Finland, Kn, Suomussalmi 320-03 EU266670

C. clarobrunneus TN02-986 H Finland, Ks, Kuusamo 325-03 EU266671

C. clarobrunneus TN05-136 H Finland, Ks, Kuusamo 1084-06 EU266675

C. clarobrunneus TN05-128 H Finland, Kn, Suomussalmi 1086-06 EU266674

C. clarobrunneus TN05-113 H Finland, Kn, Puolanka 1085-06 EU266673

C. clarobrunneus TN03-1717 H Slovakia, Liptovska kotlina basin, Va�zek 309-03 EU266672

C. coleoptera (holotype) F12650 S Sweden, Hrj, Hede 239-03 EU266684

C. coleoptera TN05-137 H Finland, Ks, Kuusamo 1137-06 EU266680

C. coleoptera TN06-137 H Finland, Kn, Suomussalmi 1274-07 EU266681

C. coleoptera IK01-024 H Finland, KiL, Kolari 013-02 EU266682

C. coleoptera IK02-020 H Finland, Kn, Kuhmo 1282-07 EU266683

(continued on next page)



subgenus Telamonia s. str. were chosen. The analyses were run

with C. norrlandicus (subgenus Phlegmacium) as an outgroup. In

all the analyses the 15 species of the section Brunnei formed

a monophyletic group and of these, 11 also morphologically

well-delimited species were included in further analyses.

Based on the preliminary analyses, C. anisatus and C. neofurvo-

laesus were chosen as outgroup species. An alignment of 82

sequences for phylogenetic analyses was produced with the

Muscle program (Edgar 2004) under default settings and

followed by manual adjustments in BioEdit (www.mbio.nc-

su.edu/BioEdit/bioedit.html). The original alignment was 580

nucleotides long (including gaps). After the exclusion of areas

with ambiguous alignment (positions 108–117, 188–201,

387–400 and 482–486), 537 positions were used for analysis.

Some Cortinarius species have fairly similar ITS regions

(Frøslev et al. 2007). When ambiguous areas were excluded

from the alignment and/or gaps were coded as missing data,

phylogenetic analyses were not always able to recognise

them as separate taxa. Thus a smaller dataset including

C. brunneus and C. glandicolor only (altogether 22 sequences,

and C. caesiobrunneus as outgroup species) were developed to

explore the sensitivity of the results to the inclusion of ambig-

uously aligned regions. Alignment was produced as above and

included 541 characters. Alignments are available in TreeBASE

(http://www.treebase.org/treebase/index.html).

MP analyses were performed with TNT v.1.0 (Goloboff et al.

2000). The analyses were run using one dual processor of

2.2 GHz. A heuristic search strategy with tree bisection–recon-

nection (TBR) branch swapping, 10 K random replicates, and

maxtrees of 1 K was employed. Two different analyses, treat-

ing gaps as missing data or as a fifth character were run for

a whole dataset. Gaps were treated as a fifth character for par-

titioned dataset. BS values were counted with 10 K replicates.

Bayesian inference (BI) was done with the program

Mr.Bayes v.3.1.1 (Huelsenbeck & Ronquist 2003). The program

FindModel (http://hcv.lanl.gov/content/hcv-db/findmodel/

findmodel.html) was used to choose an optimal model of

DNA substitution in both cases. The analyses were run using

one dual processor of 1.2 GHz. Gaps were treated as missing

data. The whole dataset was analysed with the GTR model in-

cluding a gamma shape parameter and estimating the propor-

tion of invariable sites. Two independent runs with four chains

in each were carried out with 1 M generations and sampling at

every 100th generation. All trees sampled before stationarity

were discarded using a 25 % safety margin [burn-in of 2500

trees (250 K generations)]. The partitioned dataset was

analysed with the HKY85 model without the gamma shape

parameter. The analysis was run as above with 500 K genera-

tions and burn-in of 1250 trees (25 %). The sampled trees from

both runs were combined in a 50 % majority rule consensus

phylogram with PPs. All analyses were run with the computer

clusters of the CSC, IT Centre for Science, Espoo, Finland.

Results

DNA studies

All of the 11 species are distinct as defined by their ITS data

and their morphological differences. Most of them have

more than 18 evolutionary event differences from the closest

species, but a few have less: C. brunneus/C. glandicolor five

evolutionary events and C. carabus (clade 1)/C. cicindela eight

evolutionary events. However, all of the species had much

smaller intraspecific pairwise distances compared with the

interspecific distances.

Table 2 – (continued)

Species Voucher Herb. Locality Sequenceno.

GenBankaccessionnumber

C. carabus (holotype) TN05-140 H Finland, Ks, Kuusamo 993-06 EU266661

C. carabus IK95-355 H Finland, InL, Inari 277-03 EU266654

C. carabus IK01-023 H Sweden, TL, Kiiruna 1283-07 EU266657

C. carabus IK01-025 H Finland, KiL, Kolari 015-02 EU266659

C. carabus IK04-032 H Finland, U, Hanko 1280-07 EU266658

C. carabus IK97-680 H Sweden, Jmt, Bracke 1284-07 EU266655

C. carabus IK99-707 H Finland, Kn, Kuhmo 276-03 EU266660

C. carabus IK01-026 H Finland, PS, Nilsia 1281-07 EU266656

C. cicindela (holotype) TN06-036 H Norway, Troms, Bardu 1273-07 EU266669

C. cicindela TN03-173 H Finland, Kn, Puolanka 249-03 EU266662

C. cicindela TN05-070 H Finland, St, Ikaalinen 1272-07 EU266663

C. cicindela IK95-037 H Finland, EH, Virrat 274-03 EU266664

C. cicindela IK95-303 H Finland, SoL, Sodankyla 1276-07 EU266665

C. cicindela IK97-1780 H Finland, PK, Ilomantsi 275-03 EU266666

C. cicindela IK98-637 H Finland 404-04 EU266667

C. cicindela IK99-592 H Finland, PeP, Rovaniemi 1277-07 EU266668

C. neofurvolaesus (holotype) CFP1438 S Sweden, Hrj, Hede 592-04 DQ139999

C. neofurvolaesus IK94-570 H Finland, EK, Anjalankoski 619-04 DQ140003

C. anisatus (holotype) CFP1200 S Sweden, Ang, Sabro 611-04 DQ117931

C. anisatus TN04-550 H Finland, PeP, Runteli 710-05 DQ120754

The bolded GenBank numbers were generated in this study. CFP, Cortinarius Flora Photographica; IK, I. Kytovuori; TN, T. Niskanen; JV, J. Vauras.

For acronyms of biological provinces see, e.g. Hansen & Knudsen 1992: Nordic Macromycetes 2: 24, 25.



https://www.researchgate.net/publication/10618138_MRBAYES_3_Bayesian_Phylogenetic_inference_under_mixed_models?el=1_x_8&enrichId=rgreq-b229fbe0-1497-449b-bb56-a43073a66aa2&enrichSource=Y292ZXJQYWdlOzIzNDkxNzY3O0FTOjE5MTM1OTk2NzEyOTYwMUAxNDIyNjM1MTgwMDA0

https://www.researchgate.net/publication/8667716_MUSCLE_Multiple_Sequence_Alignment_with_High_Accuracy_and_High_Throughput?el=1_x_8&enrichId=rgreq-b229fbe0-1497-449b-bb56-a43073a66aa2&enrichSource=Y292ZXJQYWdlOzIzNDkxNzY3O0FTOjE5MTM1OTk2NzEyOTYwMUAxNDIyNjM1MTgwMDA0

https://www.researchgate.net/publication/248583635_TNT_Tree_analysis_using_New_Technology?el=1_x_8&enrichId=rgreq-b229fbe0-1497-449b-bb56-a43073a66aa2&enrichSource=Y292ZXJQYWdlOzIzNDkxNzY3O0FTOjE5MTM1OTk2NzEyOTYwMUAxNDIyNjM1MTgwMDA0



Most species were genetically polymorphic, and in all cases

infraspecific polymorphisms were only or also observed intra-

genomically. Over 30 % of the samples had intragenomic poly-

morphisms. The majority of these were single base

polymorphisms, but in about 20 % of the samples one to four

base length polymorphisms were observed. In some species,

two or even three sites of intragenomic length polymorphisms

were simultaneously present in the same sequence. Polymor-

phisms were not observed in two species, C. albogaudis and

C. ectypus. For more information see point ‘ITS regions’ in

each species.

The 50 % majority rule phylogram resulting from the BI

analysis of the whole data is shown in Fig 1 with PPs indicated

above the branches. Trees from MP analyses are not shown.

The topology of MP trees corresponds with the BI tree in parts,

marked by the BS supports below the branches (gaps as miss-

ing data/gaps as fifth character). The branches without BS

support were not stable and topology was different in differ-

ent analyses.

Based on all the analyses, the ingroup representing the

sect. Brunnei is supported with 100 PP and 98/100 BS. Also C.

gentilis, which was previously placed in the sect. Limoni (e.g.

Brandrud et al. 1992; Moser 1983) based on morphology, genet-

ically belongs to this group, as indicated already, e.g. in the

study by Høiland & Holst-Jensen (2000). Within the sect.

Brunnei only a few infrageneric species groups were well sup-

ported, C. albogaudis/C. pseudorubricosus clade (98 PP, 79/98 BS),

C. brunneus/C. glandicolor (100 PP, 75/90 BS), and C. coleoptera/C.

cicindela/C. carabus clade (100 PP, 81/99 BS).

In the Bayesian analyses, treating gaps as missing data, C.

albogaudis was only supported by a PP value of 72, and two spe-

cies (C. carabus and C. glandicolor) did not form monophyletic

groups. All the other species were supported by PP values

>90. In the MP analyses only five species had>90 % BS support

and ten species >50 % BS. When the gaps were treated as

a fifth characteristic the figures for MP analyses were remark-

ably better, ten species were supported with >89 % BS values,

and C. carabus, not recognised in the Bayesian analysis,

achieved 99 % BS support, and clade I and II were separated

as well (>95 % BS). The only species not recognised in any of

these analyses was C. glandicolor; however, in separate BI (Fig 2)

and MP analyses of a partitioned dataset with no excluded

sequence areas, this species also formed a monophyletic

group, although with low support values (62 PP/65 BS).

In the comparison of GenBank and UNITE sequences with

ours, 22 were found to belong to the sect. Brunnei (Table 3) rep-

resenting six of the 11 species studied here. Of them 41 % (nine

sequences) were deposited in public databases under the

wrong name and 27 % (six sequences) without any name, leav-

ing only 32 % correctly identified (only C. brunneus and C. gen-

tilis sequences). No new Brunnei species, not sampled by us,

were found among these sequences.

Taxonomy

Cortinarius sect. Brunnei Kuhner & Romagn. ex Melot, Doc.

Mycol. XX(77): 97 (1989) em. Kytov., Niskanen & Liimat.

Type species: Cortinarius brunneus (Pers.) Fr., Epicr. Syst. mycol.:

298 (1838).

Fruit bodies are usually dark brown, less often brown, yel-

low brown to reddish brown. About half of the species have

a more or less bluish tint in flesh or mycelium. The pileus in

most species is more or less innately fibrillose, strongly hygro-

phanous or with hygrophanous streaks. The universal veil is

whitish to brownish. Exsiccata are usually� blackish, but if

the fruit body has already dried in the field, the exsiccata

can remain more brownish. Spores are subglobose to ellipsoid

to somewhat dacryoid, in one species somewhat ovoid. La-

mellar trama hyphae are often somewhat olivaceous, smooth

to strongly encrusted, best seen in the exsiccata. All the

known species grow with coniferous trees, which makes the

section Brunnei very unique among Cortinarius. We do not

know of any other group with as many species that are

restricted to conifers.

Most species in the sect. Uracei, and C. disjungendus and

related species also have blackish exsiccata. Many Uracei spe-

cies, however, have metallic or greenish tints in fresh fruit

bodies, vinaceous or metallic tints in exsiccata, and the spores

are fairly strongly verrucose, amygdaloid, or are very big (11–

12 mm long) and obovoid-ellipsoid. C. disjungendus and allied

species are easily distinguished by their long spores >9.5 mm.

Also, many species in the sect. Bovini s. lat., C. malachius, and

C. suberi have very dark exsiccata, but these are never totally

blackish, and especially the stipe is often paler. Small Brunnei

species might resemble the dark species in the sect. Hydrocybe

or Incrustati, but the latter usually have more distinct veils,

paler stipes or lamellae, different spores, or a well developed,

strongly pigmented hypoderm.

Cortinarius brunneus (Pers.) Fr., Epicr. Syst. mycol.: 298 (1838)

(Figs 1, 2, 4A, 5)

Basionym: Agaricus brunneus Pers., Syn. meth. fung.: 274 (1801):

sanctioned in Fr., Syst. mycol. 1: 211 (1821).

Type: Sweden: Angermanland: Sabra, Furuhultsan, 2 km north ofInnerbran, in wet spruce forest (Picea, Betula), 23 Aug 1987, Lind-strom et al. CFP587 [S d neotype, designated in Cortin. Fl. Photogr.II (Swedish version): 9, 1992]. GenBank no. DQ117927.2.

Illustrations: Brandrud et al. (1992: pl. B07), Consiglio et al.

(2004: pl. B31), Dahncke (1993: 818), Marchand (1983: fig 778),

Nylen (2000: 451), Ryman & Holmasen (1992: 522), Salo et al.

(2006: 192).

Pileus: 3.5–8 cm, hemispherical, later low convex with

a rounded or sometimes conical umbo, sometimes narrowly

pellucid-striate, somewhat innately fibrillose, dark red brown

to umber, sometimes with bluish tints, strongly hygropha-

nous. Lamellae: medium-spaced, strongly to weakly emargin-

ate, fairly thick, fairly broad to broad, dark brown, edge first

whitish, then concolorous. Stipe: 6–13� 0.8–1.5 cm, cylindrical

to clavate, greyish white fibrillose, later brown. Universal veil:

white to brownish white, usually forming a distinct girdle on

stipe. Mycelium: white or bluish. Context: dark red brown,

sometimes with a bluish tint, with age blackish brown, often

initially darkening at the base of the stem. Smell: indistinct or

slightly raphanoid, sometimes rubberlike. Exsiccata: blackish.

Spores: 7.5–9.5� 5.5–7 mm, Q¼ 1.24–1.44, X¼ 8.2–9.4� 6.3–

6.8 mm, XQ¼ 1.28–1.40 (eight collections; Fig 4), ovoidly to



0.1 substitutions/site

CFP1200

TN04-550 C. anisatus10099/100

CFP1438

IK94-570 C. neofurvolaesus

10098/100

68

9650/-

9469/99

10075/90

100

52

91/99

999493/9865/64

9963/89

80

10081/99

10097/99

99

100

87/98

85/95

100

100

100/100

100/100

100

72

9899/99

63/92

79/98

IK98-1233

IK97-1545TN02-1089

C. albogaudis

GK16165

TN03-944IK99-708IK00-014IK01-022

C. pseudorubricosus

TN05-140

IK04-032

IK95-355IK01-023IK01-025 C. carabus

IK97-680IK99-707IK01-026

IK98-637IK99-592IK97-1780

TN03-173TN05-070TN06-036

IK95-037IK95-303 C. cicindela

TN05-137TN06-137

F12650

IK01-024IK02-020

C. coleoptera

C. ectypusCFP71013318

IK97-080IK99-085IK00-011

C. clarobrunneusTN05-113TN03-1717

TN02-591TN02-986

TN05-128TN05-136

CFP1442CFP458

IK97-434IK95-1584

C. glandicolorTN06-247

TN04-931

CFP615a

TN04-914TN04-809TN04-803TN04-586TN02-991TN02-989

IK94-557IK97-1750Jv2953

C. brunneus

CFP587

IK94-1429

TN03-1708

IK96-542IK00-012

TN04-929TN04-932TN05-018

C. gentilis

C. caesiobrunneus

CFP178IK99-396

TN06-250

TN05-127

TN05-094TN05-041TN04-621

CFP615bIK02-019

IK00-013

IK97-1793

IK97-1970IK99-706a

IK99-706b

II

I

GK

I

II

I

II

Fig 1 – The Bayesian 50 % majority-rule consensus tree inferred from ITS regions. PPs >50 are indicated above branches and

BS supports are marked below the branches (gaps as missing data/gaps as a fifth characteristic). The bolded voucher

numbers represent type collections.



obovoidly broadly ellipsoid to subglobose, fairly finely to mod-

erately, sharply verrucose, more strongly at the apex, faintly

to moderately dextrinoid, brown and slightly dark-coloured

in KOH. Lamellar trama hyphae: olivaceous to sepia in MLZ

and KOH, weakly to fairly strongly encrusted (with zebra-

stripes and dark spots). Lamellar edge: mostly sterile, with

clavate marginal cells, 13–24� 6.5–9 mm, and relatively few

basidia. These cells are strongly umber in MLZ, mostly with

a dark umber brown, extracellular, amorphous substance; in

KOH they are fairly hyaline to olivaceous brown and without

extracellular substance. Basidia: 4–spored, 32–44� 7.5–11 mm,

in MLZ olivaceous brown to sepia: in KOH hyaline to oliva-

ceous brown, with granulose content. Pileipellis: Epicutis fairly

thin, upper hyphae 3–6 mm wide, marked with small to large

encrusted spots on the surface, and containing sepia, granu-

lose or foamy pigment in MLZ and KOH; lower hyphae wider

(>15 mm), pale sepia to light coloured, smooth to encrusted.

The transition to hypoderm is obscure. Hypoderm fairly

poorly differentiated, hyphae of the hypoderm 40–80� 25–

40 mm, in MLZ and KOH light-coloured to pale sepia brownish.

Hyphae of the context 8–23 mm wide, zebra-striped enrusted

and/or with dark sepia spots or pigment clumps. Stipitipellis

hyphae: at the stipe top dark, encrusted. Clamp connections:

present.

ITS regions (including 5.8S region): 516–520 bases long (based

on a total of eight sequences, including the type, Table 2). All

the variation is intragenomic length polymorphisms observed

at four different sites in two sequences, so the maximum

pairwise distance between two sequences is 0. One of the

variable sequences is the neotype collection, in which the

polymorphisms were overlooked (Kytovuori et al. 2005). In

the public databases we found four sequences of which

two are identical with our material (Table 3). The other two

have unique base changes and differ from one another,

although they are noted as part of the same collection

(AJ236076, UDB000158). The difference compared with C. glan-

dicolor is at least five evolutionary events (one indel, three

transitions and one transversion) and to C. clarobrunneus at

least 30 events.

Type material: The neotype consists of abundant material

including four very young fruit bodies, two and two half young

fruit bodies, two half mature fruit bodies, nine pilei, and one

stipe, all in good condition.

Ecology and distribution: Very common in mesic to damp

Picea abies forests. Known from Northern Europe and moun-

tainous areas of Central and Southern Europe.

Differential diagnosis: Cortinarius brunneus is recognised by

the fleshy fruit body, a distinct, brownish white girdle on the

stipe formed by the universal veil, and habitat with Picea. C.

glandicolor, C. clarobrunneus, and C. caesiobrunneus are similar

to C. brunneus; however, C. glandicolor has more slender fruit

bodies and usually a sparse universal veil, which rarely forms

a distinct girdle on the stipe. C. clarobrunneus has the same

habitus as C. brunneus, but has a paler pileus, smaller spores,

no distinct veil girdle, and grows with Pinus sylvestris. C. caesio-

brunneus is much more slender, has no velum girdle, and nar-

rower spores. C. subtigrinus Reumaux (1982) might be a later

synonym for C. brunneus. Our collection from Slovakia was de-

termined to be C. subtigrinus by Bidaud during the XXI J.E.C.

congress. The picture in Moenne-Loccoz et al. (1990) fits well

with C. brunneus, but we have not yet had the opportunity to

study the type specimen.

Specimens examined: Finland: Uusimaa: Kirkkonummi,

Meikotrasket, 23 Sep 2004, T.N. et al., TN04-929DS (H); loc. cit.,

T.N. et. al. TN04-932DS (H). Satakunta: Aetsa, Hoipo, 7 Aug 2000,

I.K., IK 00-012DS (H). Etela-Savo: Joutsa, Koivuranta, 30 Aug

2005, T.N. et al., TN05-018DS (H). Pohjois-Hame: Virrat, Hauhuu,

22 Aug 1996, I.K., IK96-542DS (H). d France: Languedoc-Roussillon:

Lozere, le Pont-de-Montvert, 18 Oct 1994, P.K. & I.K., IK 94-1429DS

(H). d Slovakia: Vysoke and Zapadne: Tatry Mts, township of

Podbanske, Liptovsky kosiar, 2 Oct 2003, T.N., TN03-1708DS (H,

as C. subtigrinus det. A. Bidaud). d Sweden: Smaland: Femsjo,

SE of Mellan Fargen, 12 Sep 1994, M. Moser 94/240 (IB).

Angermanland: Sabra, Furuhultsan, 23 Aug 1987, H. Lindstrom

et al. CFP587DS (neotype, S).

Cortinarius glandicolor (Fr.) Fr., Epicr. Syst. mycol.: 298 (1838)

(Figs 1, 2, 3E, 4B, 5)

Basionym: Agaricus gentilis ß glandicolor Fr., Syst. mycol. 1: 213

(1821).

Type: Finland: Satakunta: Sakyla commune, Virttaankangas, Por-saanharju, Pinus sylvestris heath forest on sandy soil, 18 Oct 2006,

Fig 2 – The Bayesian 50 % majority-rule consensus tree in-

ferred from ITS regions. PPs >50 are indicated above

branches and BS supports are marked below the branches.

The bolded voucher numbers represent type collections.




Liimatainen & Niskanen F06-247(H d neotype, NY d isoneotype.Neotypus hic designates). GenBank no. EU266700.

Illustrations: Brandrud et al. (1992: pl. B35, mixed photo col-

lection: the herbarium collection CFP615 includes more fruit

bodies than in the photo (C. glandicolor: five whole fruit bodies,

nine half fruit bodies, four pilei, C. caesiobrunneus: two whole

fruit bodies, four half fruit bodies, two pilei), and it is not pos-

sible to say with certainty which fruit bodies represent which

species. Presumably at least the pilei on the upper left, the 3rd,

4th, and 5th fruit body from the left, and the upper most pileus

of the three pilei on the right could be C. caesiobrunneus), Fries

(unpubl. plate 0317, S).

Pileus: 2.5–7 cm, conical to hemispherical, later low convex

to almost plane, with a small but distinct umbo, up to 1/2 pel-

lucid-striated, dark yellow brown to dark red brown, strongly

hygrophanous. Lamellae: medium spaced, strongly to weakly

emarginate, fairly thick, moderately broad to broad, pale yel-

low brown, soon dark brown, edge concolourous or whitish.

Stipe: 6–14� 0.4–1.1 cm, cylindrical to slightly clavate, greyish

white fibrillose, later brownish. Universal veil: white to brown-

ish white, often sparse, rarely forming a distinct girdle on the

stipe. Mycelium: white, thick, attaching to mosses at the base

of the stipe. Context: dark red brown, becoming blackish brown

with age, often first darkening at the base of the stem. Smell:

indistinct or slightly raphanoid. Exsiccata: blackish.

Spores: 8–10� 5.5–7 mm, Q¼ 1.3–1.54, �X¼ 8.5–9.5� 6.2–

6.7 mm, �XQ¼ 1.34–1.48 (13 collections, Fig 4), broadly ellipsoid

to ovoid-ellipsoid, fairly finely (to fairly strongly), densely,

sharply verrucose, weakly to fairly strongly dextrinoid, brown

and sometimes slightly dark-coloured in KOH. Lamellar trama

hyphae: in MLZ and KOH pale olivaceous brownish, smooth

to finely scabrous, often with olivaceous spots. Lamellar edge:

almost sterile to fully fertile, marginal cells mostly clavate

11–25� 5.5–10 mm, umber in MLZ, mostly with amorphous,

umber, extracellular substance; in KOH without extracellular

substance and fairly hyaline to olivaceous brown. Basidia: 4–

spored, 31–45� 8–11 mm, in MLZ olivaceous brown, in KOH hy-

aline to olivaceous brown, with granulose content. Pileipellis:

Table 3 – Sequences of Cortinarius sect. Brunnei species in the public databases

Species Name in database Locality Ecology Herbarium Collectionnumber

Basedifferenceto the type

Accessionnumber

Cortinarius

brunneus

C. brunneus Norway, Oslo, Oslo O 1 UDB000158,

AJ236076

C. brunneus Sweden, Umea Mixed forest UPS AT2001068 0 UDB001138

C. brunneus Germany Picea abies 0 AF430261

C. glandicolor C. brunneus Germany P. abies 0 AF430292

C. brunneus Scotland, Moray, Culbin

Forest

Pinus sylvestris and P.

nigra, few Betula

pubescens

AMG DG05-65 0 UDB001542


Forest

P. sylvestris and P. nigra,

few Betula pubescens

AMG DG25 0 UDB001541


Forest



AMG DG104 2 UDB001540

C. brunneus Norway, Troms,

Skibotndalen

P. sylvestris IB IB19950084 0 UDB001019,

AF325590

C. brunneus Germany IH-P17 0 AF430287

C. pseudorubricosus sp., ectomycorrhiza Canada, BC 0 DQ481698

sp., ectomycorrhiza Canada, BC 0 DQ481670

sp., ectomycorrhiza Canada, BC 1 DQ481676

C. coleoptera C. umbrinolens Scotland, Moray, Culbin

Forest



AMG 2006/217 0 UDB002396

C. umbrinolens Scotland, Moray, Culbin

Forest



AMG DG05-64 0 UDB001539

C. umbrinolens Scotland, Moray, Culbin

Forest



AMG DG103 0 UDB001538

sp., ectomycorrhiza Sweden 4 AY839207

(partial)

sp., ectomycorrhiza Sweden 1 AY839225

(partial)

C. carabus sp., ectomycorrhiza Canada, BC 2 DQ481758

C. gentilis C. gentilis Russia, Western Sibiria,

Ob river

Betula sp., Picea sp. IB 19960825 1 UDB001038,

AF325589

C. gentilis Norway, Oslo, Oslo Picea forest O 5 UDB000163,

AJ238034

C. gentilis Scotland, Invernesshire,

Glen Strathfarrar

Pinus sylvestris forest AMG 050818-14 5 UDB001563

C. gentilis USA, Washington WTU JFA10057 4 U56026

AMG, Aberdeen Mycorrhiza Group.



epicutis fairly thin, upper hyphae 3–6 mm wide, smooth, in

MLZ and KOH with olivaceous brownish to sepia, granulose

to foamy pigment, lower hyphae wider (>15 mm), smooth to

very finely scabrous, sepia to pale coloured. The transition to

the hypoderm is obscure. Hypoderm present, but fairly poorly

differentiated, hyphae of the hypoderm 40–70� 20–35 mm, in

MLZ and KOH light coloured. Hyphae of the context 7–25 mm

wide, finely to moderately zebra-striped encrusted. The pilei-

pellis generally paler than in C. brunneus. Stipitipellis hyphae: at

the stipe top pale to moderately olivaceous, weakly to dis-

tinctly encrusted. Clamp connections: present.

ITS regions (including 5.8S region): 517 bases long (based on

a total of 12 sequences, including the type, Table 2). Intrage-

nomic base polymorphisms were observed in five different

sites and two of these also included infraspecific variations.

The maximum pairwise distance is one base change. In all

the characteristic sites that differ, intermediate individuals

also occur. The infragenetic variation should be studied

more closely, before reaching a more reliable conclusion.

From sequence databases we found six C. glandicolor se-

quences, all deposited there as C. brunneus (Table 3). One of

them differs from the others by two bases but we have not

seen the chromatogram. The difference compared with C.

brunneus is at least five evolutionary events (one indel, three

transitions and one transversion) and to C. clarobrunneus at

least 28 events.

Type material: The neotype includes two whole fruit bodies,

one young fruit body, and six pilei and stipes, and the isoneo-

type includes three whole fruit bodies and two half mature

fruit bodies, all in good condition.

Ecology and distribution: Cortinarius glandicolor has a wide

ecological amplitude growing in dry to damp coniferous for-

ests with Pinus, and maybe also with Picea. We have seen col-

lections or sequences of this species from Northern Europe,

Great Britain, and Central Europe (Table 3). At least in

Northern Europe it is very common.

Differential diagnosis: The protologue of Agaricus ß glandi-

color (Fries 1821) is very brief. The fungus is described as um-

ber, with less distant lamellae, the same colour as Cortinarius

brunneus, stature like the former (C. gentilis), in coniferous for-

ests. A more extensive description is provided later in Epicrisis

Fig 3 – Photos of the new and less known Cortinarius. (A) C. albogaudis, Kytovuori 98-1233 (type, H). (B) C. caesiobrunneus,

Liimatainen & Niskanen F05-127 (type, H). (C) C. carabus Liimatainen & Niskanen F05-140 (type, H). (D) C. cicindela Liimatainen

& Niskanen F06-036 (type, H). (E) C. glandicolor, Liimatainen & Niskanen F06-247 (type, H). (F) C. pseudorubricosus, Kytovuori

98-1227 (H). (B–E) Photographs by K.L.; (A, F) photographs by I.K.



(Fries 1838) in which the taxon is presented at the species level

near C. brunneus. C. glandicolor was described by Fries as conco-

lourous umber brown, pileus umbonate, greyish brown when

dry, lamellae distant, stipe equal and slender, veil white, and

forming a temporary girdle on the stipe. He also mentions

that the species can sometimes have a pellucid-striate pileus

margin, violet stipe, and pale lamellae when young. The later

descriptions of Fries (1863, 1874) do not provide any new infor-

mation. At the moment we know of two Nordic species that

could fit into the protologue, one very common in dry to

damp coniferous forests with Pinus and presumably also Picea,

with big (8.5–10� 6–7 mm) spores, and without bluish tints,

and another less common in mesic Picea forests, often with

bluish tints, and smaller (7.5–9� 5–6 mm) spores. Fries’ unpub-

lished plate of C. glandicolor (0317, S), presents a slender, um-

ber brown fungus without bluish colours. In the latest

Nordic literature (Brandrud et al. 1992; Soop 2004) the name

C. glandicolor has been used for a slender species, which occurs

mostly in oligotrophic, sandy habitats with Pinus. The photo

collection in Brandrud et al. (1992) includes both of the species;

however, the overall description better fits our big spored one.

As neither the protologue nor the unpublished plate of C. glan-

dicolor include bluish colours, and the name in the recent Nor-

dic literature is mostly used for the Pinus associated species,

we propose that a neotype of C. glandicolor is the more com-

mon species in dry to damp coniferous forests with big spores

(for the other species, see C. caesiobrunneus). A collection from

Finland was chosen as neotype because it was the most repre-

sentative one, but our studies also include material from

Sweden, near Fries’s collection sites. C. glandicolor is recog-

nised by the slender fruit body (stipe often <8 mm broad),

sparse universal veil, and fairly big spores (8.5–10� 6–7 mm).

The sister species C. brunneus that grows in mesic Picea abies

forests is usually stouter and the universal veil forms a distinct

girdle on the stipe. The slender spruce forest species C. caesio-

brunneus, can easiest be distinguished by the narrower spores,

<6 mm wide, and it often also has bluish tints in the fruit body.

The Pinus associated C. coleoptera differs from the C. glandicolor

in having a downwards tapering stipe with only sparse

mycelium and small, obovoid-subglobose spores.

Specimens examined: Finland. Varsinais-Suomi: Vihti,

Nuuksio National Park, 21 Sep 2004, K.L. & T.N., TN04-914DS

(H). Uusimaa: Kirkkonummi, Meikotrasket, 23 Sep 2004, T.N.

et al. TN04-931DS (H); Sipoo: Paippinen, 14 Sep 2004, K.L. &

T.N., TN04-803DS (H); loc. cit., TN04-809DS (H). Satakunta: Sakyla,

Virttaankangas, 18 Oct 2006, K.L. & T.N., TN06-247DS (neotype,

H). Etela-Karjala: Anjalankoski, Kaipiainen, 15 Sep 1994, I.K.,

IK94-557 D (H). Pohjois-Karjala: Ilomantsi, Niemijarvi, 24 Sep

1997, I.K., IK97-1750DS (H). Pera-Pohjanmaa: Rovaniemi, Pisa-

vaara, 31 Aug 2004, T.N. et al. TN04-586DS (H). Koillismaa: Kuu-

samo, Kantojoki, 21 Sep 2002, T.N. TN02-991DS (H); loc. cit.

Fig 5 – Diagram showing the spore size of Cortinarius

brunneus, C. caesiobrunneus, C. clarobrunneus, C. gentilis, and

C. glandicolor. The lines are drawn on the basis of scatter

diagrams, and contain 95 % of the spore measurements of

each species. x-Axis: spore length; y-axis: spore breadth.

Fig 4 – Spores of (A) Cortinarius brunneus, (B) C. glandicolor, (C) C. caesiobrunneus, (D) C. clarobrunneus, (E) C. pseudorubricosus,

(F) C. albogaudis, (G) C. ectypus, (H) C. coleoptera, (I) C. carabus, (J) C. cicindela, and (K) C. gentilis. Drawings T.N. and I.K.



T.N. et al. TN02-989DS (H); loc. cit. T.N. et al. TN02-991 DS (H). d

Sweden: Smaland: Hyltebruk, Marieslatt, 18 Sep 1987, J. Vauras

2953DS (TUR). Jamtland: Hallesjo, 1 km S Ansjo, 8 Sep 1987,

H. Lindstrom et al. CFP615DS (S).

Cortinarius caesiobrunneus Kytov., Niskanen & Liimat. sp.

nov. (Figs 1, 3B, 4C, 5, 7)

MycoBank no.: MB 511418

Etym: caesiobrunneus (Latin), bluish brown. The name refers

to the colour of the fruit body, which is often brownish with

bluish tints.

Pileus 3–6 cm, obtuse conicus, tum explanatus, umbonatus,griseoumbrinus vel rubroumbrinus, interdum subcaesius, hygro-phanus. Lamellae subdistantae, brunneae vel umbrinae. Stipes6–10� 0.4–0.8 cm, cylindraceus vel clavatus, primo griseoalbidefibrillosus, dein obscure brunneus vel atrobrunneus, velo albido,sparsissimo. Caro atrobrunnea , interdum apice stipitis subcaeru-lea. Odor nullus vel obsolete raphanoides. Exsiccata nigrescentia.Sporae 7.5–9� 5–6 mm, ellipsoideae, subvalde verrucosae , apicevalde verrucosae. In silvis coniferis mediocriter humidis, cumPicea abiete.

Typus: Finland: Kainuu: Suomussalmi commune, Huuruvaara,Petajavaara, old, mossy, mesic grass-herb Picea abies forest withsome Betula, Pinus sylvestris, and Populus tremula, 16 Sep 2005,Liimatainen & Niskanen F05-127 (H d holotypus, NY d isotypus).GenBank no. EU266653.

Illustrations: Brandrud et al. [1992, pl. B35, mixed photo, see

C. glandicolor (2), illustrations].

Pileus: 3–6 cm, conical to hemispherical, later low convex

with a small umbo, greyish brown to reddish brown, some-

times with bluish tints, up to 1/2 pellucid-striate, strongly

hygrophanous. Lamellae: fairly distant, strongly to weakly

emarginate, fairly thick, moderately broad to broad, brown

to dark brown, sometimes bluish when young, edge concolo-

rous or whitish. Stipe: 6–10� 0.4–0.8 cm, cylindrical to clavate,

greyish white fibrillose, soon pale brown. Universal veil: white,

often very sparse, not forming a girdle on the stipe. Mycelium:

whitish. Context: dark reddish brown, darker towards the base

of stipe, sometimes bluish at the top of the stipe. Smell: indis-

tinct or slightly raphanoid, sometimes rubberlike. Exsiccata:

brownish black.

Spores: 7.5–9� 5–6 mm, Q¼ 1.38–1.70, �X¼ 7.9–8.7� 5.4–

5.8 mm, �XQ¼ 1.43–1.59 (16 collections, Fig 4), ellipsoid to

weakly ovoidly ellipsoid, finely to moderately verrucose,

strongly at the apex, moderately dextrinoid, dark, in KOH

brown and slightly dark-coloured. Lamellar trama hyphae:

smooth to very finely scabrous, seldom with some spot-like

encrustations; in MLZ pale olivaceous yellowish, in KOH pale

olivaceous brownish. Lamellar edge: fairly sterile to fertile,

marginal cells mostly clavate, 14–23� 7.5–10 mm, more to

less umber in MLZ, often with extracellular, amorphous, um-

ber substance; in KOH without extracellular substance and

fairly hyaline to olivaceous brown. Basidia: 4-spored, 32–

41� 8–9 mm, in MLZ olivaceous, in KOH hyaline to olivaceous

brown, with granulose content. Pileipellis: Epicutis fairly thin,

hyphae 3–20 mm wide hyphae, with in MLZ sepia and in KOH

olivaceous brownish, granulose or finely encrusted pigment

(mostly small spots), narrow hyphae present or not. The

transition to a hypoderm is obscure. Hypoderm present but

very poorly differentiated, hyphae of the hypoderm 30–

50� 15–25 mm, in MLZ and KOH light coloured. Hyphae of

the context 7–20 mm wide, finely to moderately zebra-striped

encrusted or spotted. Stipitipellis hyphae: at the stipe top oliva-

ceous, mostly distinctly encrusted. Clamp connections: present.

ITS regions (including 5.8S region): 527–531 bases long

(based on a total of 11 sequences, including the type, Table 2).

There are seven infraspesific variation sites in the sequences,

including four base and three length polymorphisms intrage-

nomic sites, of these two are also infraspecific polymorphism

sites. The maximum pairwise distance between the

sequences is two. Based on these two base sites most of the

specimens could be divided into two subgroups (as in Fig 1,

clade I and II) although intermediates also occur, thus there

could be a possibility of hidden taxa. This would require

more material and a more variable DNA region. No sequences

of this species exist in the public databases. The difference

compared with C. glandicolor is at least 18 evolutionary events

and to C. brunneus at least 21 events.

Type material: The holotype includes three whole and two

half mature fruit bodies, and the isotype includes three whole

and one half mature fruit bodies, all in good condition.

Ecology and distribution: In boreal to hemiboreal, mesic Picea

abies forests, fairly common. Known from Northern Europe

and Estonia.

Differential diagnosis: Typical for Cortinarius caesiobrunneus is

the often slender fruit body, sometimes with bluish tints,

sparse universal veil, habitat with Picea, and ellipsoid spores,

7.5–9� 5–6 mm, which are strongly verrucose at the apex.

The closely related and also slender C. glandicolor, has broader

spores,>6 mm, and lacks bluish colours. C. brunneus can some-

times have a distinct bluish tint, but it is stouter, the universal

veil forms a distinct girdle on the stipe, and the spores are

broader. In mixed coniferous forests, the pale C. clarobrunneus

can sometimes resemble C. caesiobrunneus, but the former is

usually stouter, lacks bluish tints, and has subglobose to

broadly ellipsoid spores, 7–8.5� 5.5–6.5 mm. Based on the ge-

netic studies C. caesiobrunneus may include two taxa. So far,

no morphological or ecological characteristics have been

found to support these clades and they are treated here as

one species. C. caesiobrunneus is fairly common in the Nordic

countries, but no name was found in the literature. Most likely

it has previously been included either in C. glandicolor or

C. brunneus (see also discussion under C. glandicolor).

Specimens examined: Estonia: Vorumaa: Vatseliina, Meeksi,

30 Sep 1997, I.K., IK 97-1970 DS (H). d Finland: Varsinais-Suomi:

Vihti, Salmenkartano, 30 Aug 2005, I.K. (H); Sipilanmaki, 7 Oct

2001, I.K. & H. Tuovila (H).Uusimaa: Espoo, Luukki, 17 Aug 2004,

T.N., TN04-062 (TUR); loc. cit., 23 Aug 2005, K.L. & T.N. TN05-004

(H); Sipoo, Paippinen, 16 Sep 2004, K.L. & T.N., TN04-827 (H);

Nuuksio National Park, 28 Aug 2004, I.K. (H); Tuusula, Klemet-

skog, 16 Sep 1940, N. Malmstrom (H). Etela-Karjala: Sippola,

Saaramaa, 19 Sep 1970, L. Fagerstrom (H). Etela-Hame: Juupajoki,

Hyytiala forestry station, 4 Sep 2005, T.N. et al. TN05-041 DS (H);



Kuru, Seitseminen National Park, 30 Sep 1982, K. Alho (TUR);

Seitsemisjoki. 28 July 2000, I.K. (H); Ruovesi, Moskulanmaki, 7

Sep 2005, I.K. (H); Susimaki, 9 Sep 2005, K.L. & T.N., TN05-077

(H). Pohjois-Hame: Hankasalmi, Ohenmaki, 13 Sep 2004, I.K.S

(H); Konnevesi, Pynnola, 7 Sep 2004, I.K. (H); Laukaa, Hiton-

hauta, 10 Sep 2004, I.K. (H); Aijala, 10 Sep 2004, I.K. (H); Saari-

jarvi, Pyha-Hakki, 10 Sep 2002, I.K., IK02-019DS (H); Virrat,

Hauhuu, 2 Aug 2004, I.K. (H). Pohjois-Karjala: Ilomantsi, Hattu-

vaara, 22 Sep 1997, I.K., IK97-1681a (H); Mekrijarvi, 25 Sep

1997, I.K., IK97-1793 DS (H); loc. cit., I.K., IK97-1794 (H); Kitee,

Hammaskallio, 20 Sep 2006, K.L. & T.N., TN06-145S (H). Kainuu:

Kuhmo, Elimyssalo, 29 Aug 1999, I.K., IK99-706a DS (H); loc. cit.,

99-706b DS (H); Hirvivaara, 14 Aug 2002, I.K. (H); Kalliojoki, 13

Aug 2002, I.K. (H); Puolanka, Paljakka, 15 Sep 2005, K.L. &

T.N., TN05-094 DS (H); Pihlajavaara, 15 Sep 2005, I.K. (H); loc.cit.,

K.L. & T.N., TN05-105S (TUR); Suomussalmi, Huuruvaara, 16

Sep 2005, K.L. & T.N., TN05-127 DS (holotype, H), loc. cit., K.L. &

T.N., TN05-135S (H). Koillismaa: Taivalkoski, Hautakyla, 27

Aug 1999, I.K. (H). Pera-Pohjanmaa: Rovaniemi, Pisavaara, 31

Aug 2004, K.L. & T.N., TN04-575 (H); Tornio, Kalkkimaa, 1 Sep

2004, K.L. & T.N., TN04-621 DS (H); loc. cit., K.L. & T.N., TN04-

639S (H). d Norway: Hedmark: Rendalen, Ovre Rendal, 6 Sep

2000, I.K. et al. S (H). d Sweden: Vastergotland: Undenas, Satra,

6 Sep 2003, T.N. et al., TN03-1259 (H). Dalarna: Idre, W of St.

Tranan, Karmorasen Nature Reserve, 5 Sep 2000, I.K., IK

00-013 DS (H). Jamtland: Hallesjo, 1 km S Ansjo, 8 Sep 1987,

H. Lindstrom et al., CFP615 DS (S, part of C. glandicolor collection).

Medelpad: Boflon O, 27 Aug 2003, T.N. et al., TN03-615 (H).

Cortinarius clarobrunneus (H. Lindstr. & Melot) Niskanen,

Kytov. & Liimat., comb. nov. (Figs 1, 4D, and 5)


Basionym: Cortinarius brunneus var. clarobrunneus H. Lindstr. &

Melot, Cortinarius Flora Photographica vol. 2 [Swedish ver-

sion]: 33 (1992).

Type: Sweden: Angermanland: Graninge, Viksmon, in sandy Pinusheath, 25 Aug 1986, Lindstrom et al. CFP458 (S d holotype). GenBankno. EU266679.

Illustrations: Brandrud et al. (1992: pl. B08).

Pileus: 3.5–8 cm, hemispherical, later low convex with

a rounded umbo, pale brown to brown, often narrowly pellu-

cid-striate, strongly hygrophanous. Lamellae: medium spaced,

strongly to weakly emarginate, fairly thick, fairly broad to

broad, yellowish brown, later brown, edge sometimes whitish.

Stipe: 6–13� 0.8–1.5 cm, cylindrical or clavate, greyish white

fibrillose, later pale brown. Universal veil: white to brownish

white, often sparse, not forming a girdle on the stipe. Myce-

lium: whitish. Context: red brown, later darker, often first dark-

ening at the base of the stem. Smell: indistinct or slightly

raphanoid, sometimes pleasant and aniseed-like. Exsiccata:

brownish black.

Spores: 7–8.5� 5–6.5 mm, Q¼ 1.23–1.44, �X¼ 7.5–8.1� 5.7–

6.1 mm, �XQ¼ 1.31–1.38 (nine collections, Fig 4), broadly ellip-

soid to subglobose, fairly finely to moderately verrucose,

somewhat more at the apex, weakly to moderately dextrinoid,

somewhat dark, in KOH brown and slightly dark-coloured.

Lamellar trama hyphae: in MLZ and KOH pale olivaceous,

smooth to very finely zebra-striped encrusted, seldom with

some spots. Lamellar edge: mostly fertile, marginal cells 13–

25� 5.5–9 mm, in MLZ umber, mostly with extra-cellular,

amorphous, umber substance, in KOH without extracellular

substance and fairly hyaline to olivaceous brown. Basidia: 4–

spored, 29–40� 7.5–10 mm, in MLZ olivaceous brownish, in

KOH hyaline to olivaceous brown, often with granulose con-

tent. Pileipellis: epicutis with upper hyphae of 3–8 mm wide, se-

pia in MLZ and KOH, granulose, very finely zebra-striped

encrustated and small-spotted, wider lower hyphae present,

but not distinct, pale. Hypoderm present, but fairly poorly dif-

ferentiated, hyphae of the hypoderm 30–60� 20–30 mm, light

coloured in MLZ and KOH. Hyphae of the context 10–25 mm

wide, smooth to very finely zebra-striped encrusted and/or

with some dark sepia spots or pigment clumps. Stipitipellis hy-

phae: at the stipe top pale to olivaceous, almost smooth to

finely encrusted. Clamp connections: present.


(based on a total of ten sequences, including the type, Table

2). There are four intragenomic polymorphism sites, includ-

ing three single base, and one length polymorphism sites,

and some correlation between these sites exists. The maxi-

mum pairwise distance is 0. More specimens and variable

DNA regions would be needed to study the genetic variation

more closely. No sequences of this species appear in the

public databases. The closest related species are Cortinarius

glandicolor and C. brunneus that differ by at least 28 evolution-

ary events.

Type material: The holotype is abundant and includes two

mature fruit bodies, one whole and four half young fruit bod-

ies, six pilei and three stipes, all in good condition.

Ecology and distribution: In dry to dryish Pinus forests, com-

mon. Known from Northern Europe and Slovakia.

Differential diagnosis: Cortinarius clarobrunneus resembles C.

brunneus, but it has a paler brown pileus, sparse universal

veil, smaller spores, and grows with Pinus. Typical for the spe-

cies seems to be that the spores in one fruit body can be of very

variable in size and form. Closely related, nameless species

seem to occur in coniferous forests and they might be more

difficult to distinguish from C. brunneus. The exsiccata of C.

clarobrunneus might sometimes be difficult to distinguish

from those of C. caesiobrunneus, because the spores of C. caesio-

brunneus can sometimes be fairly short and broad, but the lat-

ter mostly has more slender fruit bodies. The Pinus forest

species C. neofurvolaesus is reminiscent of C. clarobrunneus,

but has longer, ovoidly ellipsoid spores and dark, sordid

brown exsiccata. Cortinarius anisatus has the same pleasant

aniseed smell as some fruit bodies of C. clarobrunneus, but its

smell is usually stronger, the sparse, white universal veil

forms a sock-like sheet in young fruit bodies, the spores are

ovoid, and the species grows with Picea. Cortinarius clarobrun-

neus was first described as a variety of C. brunneus. Cortinarius

clarobrunneus, however, clearly differs from C. brunneus by

having a paler pileus, sparser universal veil, smaller spores

and a different ecology. As the genetic difference is also clear

(Fig 1), we combine C. clarobrunneus to the species level.



Specimens examined: Finland: Etela-Hame: Orivesi, Yros-

kulma, 26 Sep 1995, I.K., IK95-1584 DS (H). Kainuu: Puolanka,

Pihlajavaara, 15 Sep 2005, K.L. & T.N., TN05-113 DS (H). Suomus-

salmi, Hossa, 16 Sep 2002, T.N. et al., TN02-591 DS (H); Huuru-

vaara, 16 Sep 2005, K.L. & T.N., TN05-128 DS (H). Koilllismaa:

Kuusamo, Oulanka Biological Station, 17 Sep 2005, K.L. &

T.N., TN05-136 DS (H); Sompsanpera, 21 Sep 2002, T.N. et al.,

TN02-986 DS (H). d Slovakia: Liptovska kotlina basin:Va�zek,

Va�zecke luky, 3 Oct 2003, T.N. et al., TN03-1717 DS (H). d Swe-

den: Harjedalen: Hede, 1 Oct 1999, H. Lindstrom et al., CFP1442D

(S). Angermanland: Graninge, Viksmon, 25 Aug 1986, H. Lind-

strom et al., CFP458 DS (S, holotype); Solleftea, Junsele, 30 Aug

1997, I.K., IK97-434 DS (H).

Cortinarius pseudorubricosus Reumaux, Atlas des Cortinaires

1: 21 (1990) (Figs 1, 3F, 4E, 6, 8)

Type: France: Ardennes: Foret de Belval, sous epiceas, 2 Nov 1986,leg. et det. P. Reumaux no 304(G – holotype). GenBank no. EU266709.

Illustrations: Moenne-Loccoz et al. (1990: pl. 18).

Pileus: 4.5–9 cm, with a long down-curved margin, innately

fibrillose, brown, faintly hygrophanous. Lamellae: medium

spaced, strongly to weakly emarginate, fairly thick, at first

moderately broad, then broad, greyish brown, sometimes

with a bluish tint, soon dark chocolate brown, edge concolour-

ous or white. Stipe: 6–11� 0.8–1.3 cm, clavate, up to 2.5 cm

broad at base, fairly firm, greyish white fibrillose, later brown.

Universal veil: white, forming a sock-like sheath or thick girdles

on the stipe. Mycelium: white or sometimes bluish. Context:

dark brown, often with a distinctly bluish tint, strongest at

the top of the stipe, the whole context darkening with age, of-

ten first at the base of the stem. Smell: indistinct or slightly

raphanoid. Exsiccata: brownish to brownish black.

Spores: 6–8� 4–5.5 mm, Q¼ 1.36–1.63, �x¼ 6.8–7.3� 4.6–5 mm,�xQ¼ 1.43–1.53 (15 collections, Fig 4), slightly obovoidly ellip-

soid, with a gradually tapering base, not very thin-walled,

fairly finely verrucose, somewhat more at the apex, fairly

weakly dextrinoid, somewhat dark, in KOH brown and slightly

dark-coloured. Lamellar trama hyphae: very pale olivaceous to

pale olivaceous in MLZ and KOH, smooth to sometimes finely

zebra-striped encrusted. Lamellar edge: fairly sterile, with

mostly narrowly clavate marginal cells, 11–18� 4.5–6.5 mm,

and few basidia, these cells fairly light-coloured to umber in

MLZ, often with extra-cellular, amorphous, dark umber brown

substance, in KOH without extracellular substance and fairly

hyaline to olivaceous brown. Basidia: 4–spored, 25–35� 6–

8 mm, in MLZ almost concolourous to olivaceous brownish,

in KOH hyaline to olivaceous brown, often with granulose

content. Pileipellis: Epicutis with upper hyphae of 3–6 mm, in

MLZ and KOH pale sepia to sepia, granulose and small-spot-

ted, wider lower hyphae scanty. Hypoderm present, but very

poorly differentiated, hyphae of the hypoderm 25–60� 15–30

mm, thin-walled. Hyphae of the context 7–20 mm wide, smooth

to very finely zebra-striped encrusted and/or with some um-

ber brown spots. Stipitipellis hyphae: at the stipe top olivaceous,

smooth (to finely) encrusted. Clamp connections: present.


a total of five sequences, including the type, Table 2), all five

sequences are identical. Within these sequences the maxi-

mum pairwise distance is 0. In addition, we have sequenced

four other collections, but the quality of the seqeunces was

not good enough for publication. For three of them, the reason

was two intragenomic length polymorphisms occurring in the

same sequence. In the public databases we found three

sequences, one of which had one base pair difference

(Table 3). Thus there is some variation inside C. pseudorubrico-

sus that might be of taxonomic value, but this would need

further studies. The closest species C. albogaudis differs by at

least 18 evolutionary events.

Type material: The holotype includes a part of a mature fruit

body, which is in good condition.

Fig 6 – Diagram showing the spore size of Cortinarius albo-

gaudis, C. carabus, C. cicindela, C. coleoptera, C. ectypus, and C.

pseudorubricosus. The lines are drawn on the basis of scatter

diagrams, and contain 95 % of the spore measurements of

each species. x-Axis: spore length; y-axis: spore breadth.

Fig 7 – Distribution of Cortinarius caesiobrunneus in North

Europe according to the material examined.



Ecology and distribution: In mesic to dry coniferous forests,

mostly with Picea, also with Pinus, presumably fairly rare.

Known from Northern Europe and mountainous areas of

Central Europe. An ectomycorrhizal ITS sequence from Tsuga

heterophylla roots from Western Canada, Vancouver, was also

identified as this species (Table 3).

Differential diagnosis: Cortinarius pseudorubricosus is a fairly

big, fleshy Brunnei species with an innately fibrillose pileus,

and very small, obovoidly ellipsoid spores. Macroscopically it

can also resemble some Bovini species, but none of them

have such small spores, and their exsiccata are never totally

black. The exsiccata of C. suberi Soop (sect. Malachii) can some-

times also be very dark, but the white veil is more distinct, and

it has bigger spores (7.5–8–9� 5–5.5 mm). The original descrip-

tion corresponds well with our observations, except that the

bluish tint in the context is not mentioned, and the spores

are reported to be 6.5–7.5� 4–4.5 mm, narrower than in our

measurements (�X¼ 4.7 mm in the type). The ITS sequence of

the type is identical to our material.

Specimens examined: Finland: Varsinais-Suomi: Vihti,

Nummela, 9 Oct 2004, I.K.S (H). Uusimaa: Espoo, Nuuksio, 4

Sep 1998, I.K., IK98-1631S (H); Tuusula, Ruotsinkyla, 21 Sep

1986, L. SeppanenS (H, as C. malachius). Etela-Hame: Lammi,

Evo, 30 Aug 1979, R. Tuomikoski & M. Korhonen, 2834S (H);

Tammela, Syrja, 1877, P.A. Karsten, 295S (H, as C. bovinus), loc.

cit., Sep 1877, P.A. Karsten, 296 (H, as C. bovinus). Pohjois-Hame:

Virrat, Navettavuori, 19 Sep 1999, I.K., IK99-708DS (H). Pohjois-

Savo: Lapinlahti, Rasila, 21 Sep 1956, O. SchulmannS (H, as

Hydrocybe bovina). Pera-Pohjanmaa: Tervola, Raemaki 11 Sep

1997, I.K., IK97-1200 S (H); Tornio, Arpela, 20 Aug 1998, I.K.,

IK98-1227DS (H); Ylitornio, Kaitajarvi, 9 Sep 1997, I.K., IK97-

1042DS (H). Koillismaa: Kuusamo, Oulanka National Park, 23

Sep 2002, T.N. et al., TN02-1097DS (H); loc. cit., 19 Sep 2005,

T.N. et al., TN05-176DS (H). Enontekion Lappi: Enontekio,

Muotkajarvi, 9 Aug 2001, I.K., IK01-022DS (H). d France: Ard-

ennes: Foret de Belval, 2 Nov 1986, P. Reumaux, 304DS (holotype,

G). d Norway: Sogn og Fjordane: Luster, Solvorn, 9 Sep 2000, I.K.

& T.N., IK00-014DS (H). d Sweden: Uppland: Hallnas, Ornstenen,

19 Sep 2007, M. Toivonen & I.K. (H); Alvkarleby, 5 km E of Alvkar-

len, 5 Sep 1998, H. Lindstrom, 98.835 (UPS). Medelpad: Stode,

Soransjoberget, H. Lindstrom, 90.050 (UPS). Jamtland: Berg, E of

Ratan, 31 Aug 2003, T.N. et al., TN03-944DS (H).

Cortinarius albogaudis Kytov., Niskanen & Liimat.

sp. nov. (Figs 1, 3A, 4F, 6, 9)MycoBank no.: MB 511421

Etym: albogaudis (Latin), album¼white, gaudium¼ glad,

pleasure. The epithet refers to the similarity with C. pinigaudis.

Pileus 3–8 cm, convexus, dein planoconvexus, atroochraceo-brunneus vel atrogriseobrunneus, cum venis hygrophanis.Lamellae subdistantae, griseobrunneae, dein atrobrunneae. Stipes6–14� 0.8–1.5 cm, clavatus, primo griseoalbide fibrillosus, deinatrobrunneus, velo albido, cingulato. Caro brunnea vel umbrina,variegata. Odor nullus. Sporae 5.5–7.5� 5–6 mm, obovate subglobo-sae, subvalde echinate verrucosae. In silvis coniferis mediocriterhumidis, cum Picea abiete in solo calcareo.

Typus: Finland: Pera-Pohjanmaa: Tornio commune, Arpela,Korkiamaa, Runteli, rich grass-herb Picea forest with hardwoodbushes and some Pinus, slightly paludified depressions, calcare-ous ground, 20 Aug 1998, Kytovuori 98-1233 (H d holotypus,NY d isotypus). GenBank no. EU266635.

Pileus: 3–8 cm, hemispherical, later low convex with a low

and broad umbo, innately fibrillose, yellow brown to greyish

brown, with hygrophanous streaks. Lamellae: medium spaced,

strongly to weakly emarginate, fairly thick, at first moderately

broad, then broad, greyish brown, soon dark chocolate brown,

edge whitish, later concolorous. Stipe: 6–14� 0.8–1.5 cm, cla-

vate, up to 3 cm broad at base, greyish white fibrillose, later

brown. Universal veil: white, distinct, forming a girdle on the

Fig 9 – Distribution of Cortinarius albogaudis and C. ectypus in

North Europe according to the material examined.

Fig 8 – Distribution of Cortinarius pseudorubricosus in North

Europe according to the material examined.



stipe. Mycelium: white. Context: brown, with age and at the

base of the stipe dark brown to blackish brown, marbled

hygrophanous. Smell: indistinct. Exsiccata: blackish.

Spores: 5.5–7.5� 5–6 mm, Q¼ 1.15–1.30, �X¼ 6.6–6.7� 5.3–

5.5 mm, �XQ¼ 1.20–1.24 (4 collections, Fig 4), obovoid-subglo-

bose, not very thin-walled, echinate-verrucose, weakly

dextrinoid, dark, in KOH brown and slightly dark-coloured.

Lamellar trama hyphae: smooth to distinctly, finely, densely

scabrous, olivaceous yellowish to olivaceous brownish in

MLZ, pale olivaceous brownish in KOH. Lamellar edge: fairly

sterile with clavate marginal cells, 11–20� 5–9 mm, and fairly

few basidia, the cells fairly light to dark umber in MLZ, with

scanty to fairly abundant extra cellular, amorphous, dark um-

ber substance, in KOH without extracellular substance and

fairly hyaline to olivaceous brown. Basidia: 4–spored, 26–

32� 7–8 mm, in MLZ olivaceous, in KOH hyaline to pale oliva-

ceous brown, some olivaceous brown, often with granulose

content. Pileipellis: epicutis thin, upper hyphae 3–10 mm wide,

in MLZ and KOH sepia, granulose, smooth to very finely

encrusted, lower hyphae somewhat wider (>15 mm), and

paler. Hypoderm present, hyphae 20–60� 20–30 mm, fairly

poorly differentiated, but a bit more distinct than in most

other Brunnei species. Hyphae of the context 7–20 mm wide,

smooth to very finely zebra-striped encrusted and/or with

some umber brown spots. Stipitipellis hyphae: at the stipe top

pale olivaceous, mostly smooth. Clamp connections: present.


a total of three sequences, including the type, Table 2). All the

known specimens from different regions (collected before

2007) have been sequenced and they are identical without

any polymorphisms. We did not find this species from the

public databases. The difference compared with the closest

relative C. pseudorubricosus is at least 18 evolutionary events.

Type material: The holotype includes four whole and one

half fruit bodies, and the isotype includes two whole and

one half mature fruit bodies, all in good condition.

Ecology and distribution: The species is only known from five

localities in Finland growing with Picea abies, on calcareous

ground.

Differential diagnosis: Cortinarius albogaudis can be recog-

nised by the subglobose, echinate verrucose spores, and C.

pinigaudis-like appearance, but with a white universal veil.

This combination is so unique that C. albogaudis is mostly eas-

ily separated from the other Brunnei species.

Specimens examined: Finland: Kainuu: Puolanka, Vayryla, 15

Aug 1997, I.K., IK97-1545DS (H). Pera-Pohjanmaa: Tornio,

Kalkkimaa, 22 Aug 2007, I.K. (H); Korkiamaa, 20 Aug 1998,

I.K., IK98-1233D (holotype H, isotype NY); loc. cit., I.K.,

IK98-1234S (H); loc. cit., 22 Aug 2007, I.K. (H). Koillismaa:

Kuusamo, Oulanka National Park, Korvasvaara, 29 Aug 2007,

I.K. (H); Puukkorinne 23 Sep 2002, T.N. et al., TN02-1089aDS

(H); loc. cit., 20 Sep 2005, T.N. et al., TN05-186S (H); loc. cit., 1

Sep 2007, I.K. (H); loc. cit., 1 Sep 2007, T.N. & K.L., TN07-124 (H).

Cortinarius ectypus J. Favre, Ergebn. Wiss. Unters. Schweiz-

NatnParks 6, 42: 513 and 588 (1960) (Figs 1, 4 G, 6, 9)

Type: Switzerland: dans le vallon du haut-marais de la foret deSauaidas, 1550 m, au bord de ce marais, mais non sur celui-ci,sous epiceas, 22 Aug 1955, J. Favre no 625 (G d holotype). GenBankno. EU266689.

Illustrations: Brandrud et al. (1992: pl. B28, includes four C.

paragaudis fruit bodies: two fruit bodies from the right, the

lower right pileus of the three pilei, and one slender half fruit

body in the middle), Favre (1960: pl. III) (Fig 3).

Pileus: 3–7 cm, conical to hemispherical, later low convex

with a small umbo, coarsely innately fibrillose, greyish brown

to dark brown, not hygrophanous but with hygrophanous

streaks. Lamellae: medium spaced, strongly to weakly emar-

ginate, fairly thick, at first moderately broad, pale greyish

brown, later brown, edge first whitish, then concolorous. Stipe:

4–8� 0.6–1.1 cm, cylindrical, greyish white fibrillose, later

brownish. Universal veil: brownish, indistinct. Mycelium: white

or bluish. Context: brown, often with a bluish tint, especially at

the top of the stipe, marbled hygrophanous, darkening with

age, often first at the base of the stem. Smell: indistinct.

Exsiccata: blackish.

Spores: 5.5–7� 4.5–6 mm, Q¼ 1.17–1.36, �X¼ 6–6.7� 4.9–

5.3 mm, �XQ¼ 1.23–1.3 (seven collections, Fig 4), ovoid-subglo-

bose, thin-walled, finely, lowly verrucose, slightly more at

the apex, not to very weakly dextrinoid, brown and slightly

dark-coloured in KOH. Lamellar trama hyphae: in MLZ and

KOH pale olivaceous, smooth to finely, densely scabrous,

sometimes with olivaceous spots. Lamellar edge: fairly sterile

to fairly fertile, marginal cells clavate, 11–24� 5.5–8 mm, fairly

pale umber in MLZ, colour mostly intracellular or on cell walls,

extracellular amorphous umber substance scanty; in KOH

fairly hyaline to olivaceous brown and without extracellular

substance. Basidia: 4–spored, 29–44� 7–9 mm, almost concolo-

rous to olivaceous brownish in MLZ and KOH, often granulose.

Pileipellis: epicutis fairly thin, hyphae 10–20 mm wide, finely to

zebra-striped encrusted, sepia in MLZ and KOH, narrow hy-

phae very scanty. The transition to the hypoderm is obscure.

Hypoderm poorly differentiated and obscure, hyphae of the

hypoderm 35–60� 15–25 mm, in MLZ and KOH light coloured.

Hyphae of the context 6–20 mm wide, smooth to finely

encrusted and with some spots, slightly olivaceous in KOH.

Stipitipellis hyphae: at the stipe top pale olivaceous brownish,

smooth to very finely encrusted. Clamp connections: present.


a total of five sequences, including the type, Table 2), and all

the sequences are identical. We did not find this species in

the public databases. The closest known species, Cortinarius

gentilis, differs by more than 30 evolutionary events.

Type material: The type includes nine fairly young, half fruit

bodies in good condition.

Ecology and distribution: In mesic to damp Picea abies forests,

presumably rare. Known from Central and Northern Europe.

Differential diagnosis: Cortinarius ectypus is distinguished

from all other Brunnei species by the combination of dark sor-

did brown, innately fibrillose to finely squamulose pileus,

which is not hygrophanous but with hygrophanous streaks,

and small, finely verrucose, ovoid-subglobose spores.



Specimens examined: Finland: Ahvenanmaa: Jomala, Gottby,

21 Aug 2000, J. Vauras, 16669S (TUR, H dupl.). Uusimaa: Espoo,

Luukki, 3 Aug 1997, I.K., IK97-080 DS (H). Etela-Hame: Virrat,

Monoskyla, 26 Jul 2000, I.K., IK00-011 DS (H). Kainuu: Kuhmo,

Elimys, 24 Jul 1999, I.K., IK99-085 DS (H). Pera-Pohjanmaa:

Rovaniemi, Kaihua, 23 Aug 1999, I.K.S (H). d Sweden:

Halsingland: Los, N of Karlsberg, 16 Sep 1995, P.K & I. K., IK95-

1436 S (H). Jamtland: Undersaker, Edsasdalen, 14 Aug 1988,

H. Lindstrom et al., CFP710DS (S). d Switzerland: Dans le vallon

du haut-marais de la foret de Sauaidas, 22 Aug 1955, J. Favre,

625 DS (holotype G).

Cortinarius coleoptera H. Lindstr. & Soop, Journal des JEC,

Journees Europeenes du Cortinaire 2: 58 (1999) (Figs 1, 4H, 6, 10)

Type: Sweden: Harjedalen: Hede, Remmen, 24 Aug 1998, leg. K.Soop F12650 (S d holotype). GenBank no. EU266684.

Illustrations: Lindstrom & Soop (1999: 45), Soop (2004: pl. 23,

Fig. 89).

Pileus: 2–5.5 cm, conical to hemispherical, later low convex

with a small but distinct umbo, up to 1/3 pellucid-striated,

dark red brown, hygrophanous. Lamellae: fairly distant,

strongly to weakly emarginate, fairly thick, moderately broad

to broad, dark red brown, edge whitish or concolorous. Stipe:

4–9� 0.4–0.7 cm, cylindrical, greyish white fibrillose, later

reddish brown. Universal veil: white, very sparse. Mycelium:

white. Context: dark red brown, darkening with age, often

first at the base of the stem. Smell: indistinct. Exsiccata:

greyish black.

Spores: 6.5–8� 5–6 mm, Q¼ 1.19–1.4, �X¼ 6.7–7.4� 5.2–

5.5 mm, �XQ¼ 1.24–1.38 (17 collections, Fig 4), obovoidly subglo-

bose, moderately, densely, and sharply verrucose, weakly

dextrinoid, dark, in KOH brown and slightly dark coloured. La-

mellar trama hyphae: fairly strongly zebra-striped encrusted,

seldom with some spots. In MLZ olivaceous brownish, in

KOH pale olivaceous to olivaceous brownish. Lamellar edge:

fully fertile, marginal cells 13–22� 5–9 mm, somewhat umber

in MLZ, with extracellular, amorphous, umber substance or

not; in KOH fairly hyaline to olivaceous brown, without extra-

cellular substance. Basidia: 4–spored, 27–40� 7–9 mm, oliva-

ceous brown in MLZ, in KOH hyaline to olivaceous brown,

often with granulose content. Pileipellis: With epicutis of 4–

8 mm wide, granulose and very finely scabrous, in MLZ and

KOH pale sepia to sepia upper hyphae, and few wider

(>15 mm), very finely scabrous lower hyphae. Hypoderm pres-

ent, but fairly poorly differentiated, hyphae of the hypoderm

35–60� 15–25 mm, in MLZ and KOH light coloured. Hyphae of

the context 6–20 mm wide, strongly zebra-striped encrusted

and with few large spots, pale olivaceous brownish in

KOH. The pileipellis is distinctly paler than in C. carabus and

C. cicindela. Stipitipellis hyphae: at the stipe top pale

olivaceous, smooth to weakly encrusted. Clamp connections:

present.


(based on a total of 5 sequences, including the type, Table 2),

the type specimen has one intragenomic length polymor-

phism site, but otherwise all the other sequences are identical.

The maximum pairwise distance is 0. In the public databases

there are three identical sequences from Scotland (under the

name Cortinarius umbrinolens) and two partial ITS sequences

from Sweden from ectomycorrhizal root tips, which differ by

one and four base changes (Table 3). This latter variation

would be extensive for infraspecific variation but we have

not seen the chromatograms to be sure that the differences

are real. The closest known species, C. cicindela, differs by

more than 30 evolutionary events.

Type material: The holotype includes two half fruit bodies,

which are in good condition.

Ecology and distribution: In dry Pinus sylvestris heath forests

on sandy soil, less frequently in mesic Picea abies forests

with Pinus sylvestris. Known in Northern Europe and Great

Britain, common in Northern Europe.

Differential diagnosis: Cortinarius coleoptera can be easily

recognised by the small to medium sized, dark reddish

brown fruit bodies, fairly stiff, downwards tapering stipe

with sparse mycelium and small subglobose spores. The

other two small Brunnei species, C. carabus and cicindela,

have on average smaller, darker and less reddish brown fruit

bodies and longer spores, and the pileus is often thin fleshed

and slightly innately fibrillose. Cortinarius glandicolor can also

resemble C. coleoptera, but the stipe is clavate, often soft, and

with thick mycelium attaching mosses to the base of the

stipe, and the spores are much bigger. In the original de-

scription (Lindstrom & Soop 1999) the spores are mentioned

as ‘Sp ellipsoideis usque subglobosis, 7-8.7� 5-6 mm confertae ver-

rucosis’, but based on our observations the spores are 6.5–

8� 5-6 mm and subglobose. Cortinarius coleoptera is said to

be uncommon in dryish Pinus forest on calcareous ground,

also in the later description by Soop (2004), but we have

found the species to be common at least in oligotrophic,

sandy Pinus heath forests and occasional in mesic coniferous

forests with Pinus.

Fig 10 – Distribution of Cortinarius coleoptera in North Europe

according to the material examined.


https://www.researchgate.net/publication/265026439_Quelques_petits_Telamonia_calciphiles?el=1_x_8&enrichId=rgreq-b229fbe0-1497-449b-bb56-a43073a66aa2&enrichSource=Y292ZXJQYWdlOzIzNDkxNzY3O0FTOjE5MTM1OTk2NzEyOTYwMUAxNDIyNjM1MTgwMDA0


Specimens examined: Estonia: Hiiumaa: Tahkuna, 13 Sep

2001, T.N. & I.K. S (H). Puhaleba, Nomba, 13 Sep 2001, J. Vauras,

17807F S (TUR, as Cortinarius glandicolor). d Finland: Uusimaa:

Nurmijarvi, Kiljava, 13 Oct 2004, I.K. (H); Sipoo, SW of Hindsby,

21 Oct 2004, I.K. & Bonsdorff (H); Tammisaari, W of the railway

station of Raasepori, 15 Oct 2004, I.K.S (H). Etela-Karjala:

Virolahti, Lansikyla, 18 Sep 1993, L. Fagerstrom S (H). Satakunta:

Ikaalinen, Seitseminen Nature Reserve, 7 Sep 2005, I.K.S (H).

Etela-Hame: Orivesi, Konilampi, 17 Oct 2004, I.K. (H); Ruovesi,

Tuuhonen, 20 Sep 1999, I.K.S (H); Visuvesi N, 14 Oct 2001, I.K.

(H); loc. cit., 18 Oct 2004, I.K. (H). Etela-Savo: Valkeala,

Tuohikotti, 11 Oct 2005, I.K. (H). Pohjois-Hame: Aanekoski, Kon-

ginkangas, 9 Sep 2002, I.K. (H). Pohjois-Karjala: Ilomantsi,

Hattuvaara, 22 Sep 1996, I.K., IK97-1681b S (H); Lieksa, Kitsi, 7

Sep 1998, K. Salo (H). Keski-Pohjanmaa: Pyhanta, Itamaki, 14

Sep 2005, I.K. (H). Kainuu: Kuhmo, Viiksimo, 30 Aug 1999, I.K.

(H); Puolanka, Pirunkirkko, 16 Sep 2002, I.K., IK02-020 DS (H);

Suomussalmi, Hossa, 16 Sep 2002, T.N. et al., TN02-571 (H);

loc. cit., T.N. et al., TN02-573 (H); loc. cit., T.N. et al., TN02-574

(H); loc. cit., T.N. et al., TN02-582 (TUR); loc. cit., T.N. et al.,

TN02-624 S (H); loc. cit., 18 Sep 2006, K.L. & T.N., TN06-137 DS

(H); Huuruvaara, 16 Sep 2005, K.L. & T.N., TN05-134 (H); Siika-

jarvi, 14 Sep 1997, I.K. S (H); loc. cit., 8 Aug 2002, I.K. et al. (H);

Sarkka, 16 Sep 2002, T.N. et al., TN02-656 S (H). Oulun Pohjanmaa:

Haukipudas, Kello, 18 Aug 2001, I.K. (H). Koillismaa: Kuusamo,

Kantojoki, 2 Sep 2002, T.N. et al., TN02-513 (H); loc. cit., T.N.

et al., TN02-515 (H); loc. cit., T.N. et al., TN02-523 (H); loc. cit.,

T.N. et al., TN02-529 (H); Oulanka biological station, 31 Aug

2002, K.L. & T.N., TN02-291 (H); loc. cit., K.L. & T.N., TN02-392

(TUR); loc. cit., 18 Sep 2002 T.N. et al., TN02-697a (H); loc. cit.,

T.N. et al., TN02-712 (H); loc. cit., T.N. et al., TN02-757 (H); loc.

cit., T.N. et al., TN02-761 (H); loc. cit., 17 Sep 2005, K.L. & T.N.,

TN05-137 DS (H); loc. cit., K.L. & T.N., TN05-155 (H); Vuotunki,

21 Sep 2002, I.K. & T.N., TN02-898 (H); loc. cit., I.K. & T.N.,

TN02-921 (TUR). Salla, Kaunisharju recreation area, 2 Sep

2002, K.L. & T.N., TN02-501 (H); Naruska, 1 Sep 2002, K.L. &

T.N., TN02-456 (H). Pera-Pohjanmaa: Pello, Orajarvi, 17 Aug

2001, I.K. (H); Rovaniemi, Louejarvi, 28 Aug 2004, K.L. &

T.N., TN04-431 (H); Louevaara, 28 Aug 2004, K.L. & T.N.,

TN04-479 (H); Pisavaara, 26 Aug 2004, K.L. & T.N., TN04-275

(H); Ylitornio, Meltosjarvi, 21 Aug 1999, I.K., IK99-454 (H); Pal-

orommas nature reserve area, 2 Sep 2004, K.L. & T.N., TN04-

666 (H). Kittilan Lappi: Kolari, 15 Aug 2001, I.K., IK01-024 D (H).

Muonio, Laaksivaara, 11 Aug 2001, I.K. (H). d Norway: Hed-

mark: Eidskog, Magnor, 16 Sep 2000, I.K. (H). Troms: Storf-

jords, Skibotn, 24 Aug 2006, K.L. & T.N., TN06-066 (H). d

Sweden: Dalarna: Idre, Karmorasen Nature Reserve, 5 Sep

2000, I.K. et al. S (H); Alvdalen, Oxgrav, Kytovuori et al. (H). Hal-

singland: Skog, Tonnebro, 24 Aug 2003, T.N. et al., TN03-399

(H). Harjedalen: Hede, Remmen, 24 Aug 1998, K. Soop,

F12650 D (holotype S). Medelpad: Borgsjo, Alby, 28 Aug 2003,

T.N. et al., TN03-674 (H); Havero, Satervallen, 31 Aug 2003,

T.N. et al., TN03-881 (H). Jamtland: Berg, Roian, 3 Sep 2003,

T.N. et al., TN03-1121 (H); Klovsjo, Klovsjohojd, 25 Aug

2003, T.N. et al., TN03-437 (H); Ratan, Handsjo, 31 Aug 2003,

T.N. et al., TN03-912 (H), loc. cit., T.N. et al., TN03-975 (H),

loc. cit., T.N. et al., TN03-990 S (H); Rojan, 3 Sep 2003, T.N.

et al., TN03-1121 (H). Torne Lappmark: Jukkasjarvi, Paittas-

jarvi, 13 Aug 2001, I.K. S (H); W side of the village of Jukkas-

jarvi, 13 Aug 2001, I.K. (H). Norrbotten: Pajala, SW of

Aareavaara, 17 Aug 1998, I.K., IK98-1051 S (H); Kaunisjoen-

suu, 15 Aug 2001, I.K. S (H); Perajavaara, 16 Aug 1998, I.K.,

IK98-954 (H).

Cortinarius carabus Kytov., Niskanen & Liimat. sp. nov.

(Figs 1, 3C, 4I, 6, 11)


Etym: carabus (Latin), a beetle genus Carabus belonging to

Coleoptera. The name refers to the closely related C. coleoptera.

Pileus 0.5–2.5 cm, conicus, tum explanatus, umbonatus, atroum-brinus ad quasi nigricantem, hygrophanus. Lamellae subdistan-tae, brunneae vel umbrinae. Stipes 2–6� 0.2–0.4 cm,cylindraceus, primo griseoalbide fibrillosus, dein obscure brun-neus vel atrobrunneus, velo albido, sparsissimo. Caro atrobrunnea.Odor nullus. Exsiccata nigrescentia. Sporae 7.5–9� 4.5–5.5 mm,dacryoideae, subvalde verrucosae, apice valde verrucosae. In sil-vis coniferis aridis et semiaridis, cum Pino sylvestri.

Typus: Finland: Koillismaa: Kuusamo commune, Oulanka biolog-ical station; dry pine heath forest (Pinus sylvestris) on sandy soil, 17Sep 2005, Liimatainen & Niskanen F05-140 (H d holotypus, NY d iso-typus). GenBank no. EU266661.

Pileus: 0.5–2.5 cm, conical to broadly conical sometimes

hemispherical, later low convex to almost plane with a small

umbo, often very finely innately fibrillose, dark brown to al-

most blackish brown, hygrophanous. Lamellae: medium

spaced to fairly distant, strongly to weakly emarginate, me-

dium thick, moderately broad to fairly broad, dark brown,

edge concolorous or whitish. Stipe: 2–6� 0.2–0.4 cm, cylindri-

cal, greyish white fibrillose, later brown to dark brown. Univer-

sal veil: whitish, very sparse. Mycelium: white, sparse. Context:

dark brown, darkening with age, often first at the base of the

stem. Smell: indistinct. Exsiccata: greyish black.

Spores: 7.5–9� 4.5–5.5 mm, Q¼ 1.54–1.83, �X¼ 7.9–8.4� 4.7–

4.9 mm, �XQ¼ 1.62–1.79 (14 collections, Fig 4), narrowly

dacryoid to fusoid, with a low suprahilar depression,

Fig 11 – Distribution of Cortinarius carabus in North Europe




moderately verrucose, strongly at the apex, weakly dextri-

noid, often very dark, in KOH brown and dark-coloured. Lamel-

lar trama hyphae: pale olivaceous brownish to olivaceous

brownish in MLZ and KOH, somewhat zebra-striped and

very strongly spot-like encrusted (small to large spots). Lamel-

lar edge: fertile, marginal cells clavate 9–22� 5–8 mm, strongly

umber in MLZ, with extracellular, amorphous, umber sub-

stance; in KOH without extracellular substance and pale oliva-

ceous brown to umber. Basidia: 4–spored, 25–36� 6–9 mm,

olivaceous brownish to umber in MLZ, fairly hyaline to umber

in KOH. Pileipellis: Epicutis fairly thin, upper hyphae 3–6 mm wide,

granulose, sometimes zebra-striped encrusted, sepia in MLZ and

KOH, lower hyphae>10 mm wide, finely scabrous, pale sepia. The

transition to a hypoderm is obscure. Hypoderm present, but

fairly poorly differentiated, hyphae of the hypoderm 35–

60� 15–30 mm. Hyphae of the context 6–15 mm wide, sometimes

zebra-striped encrusted, umber brown spots abundant. In MLZ

an umber brown amorphous substance present in some fruit

bodies. Stipitipellis hyphae: at the stipe top pale olivaceous yellow-

ish to brownish, mostly distinctly encrusted. Clamp connections:

present.

ITS regions Cortinarius carabus clade I (including 5.8S re-

gion): 519–520 bases long (based on a total of five sequences,

including the type, Table 2). The type sequence has one

intragenomic base polymorphisms site (R) and the collection

IK01-023 has one intragenomic length polymorphism site.

Otherwise all the sequences are identical and the maximum

pairwise distance is 0. In the public databases there is one

ectomycorrhiza sequence from Canada (DQ481758) differing

by two base changes (Table 3). To study the meaning of this

variation we would need more material from North Amer-

ica. The difference compared with Cortinarius carabus clade

II is at least four evolutionary events (three transitions and

one four basepair transversion) and with C. cicindela, at least

eight events (four indels, three transitions, and one

transversion).

ITS regions Cortinarius carabus clade II (including 5.8S re-

gion): 519 bases long (from a total of three sequences, Table

2, deposited in the public databases as Cortinarius aff. carabus),

all sequences are identical. The maximum pairwise distance

is 0. Although C. carabus clade I and II seem like genetically

different species based on Nordic material: more material

from the circum boreal zone may change the conclusion.

The difference compared with C. cicindela is at least ten evolu-

tionary events (four indels, five transitions and one

transversion).

Type material: The holotype includes three whole and two

half mature fruit bodies, and the isotype includes two whole

and one half mature fruit bodies and one pileus, all in good

condition.

Ecology and distribution: In sandy Pinus sylvestris heath for-

ests, often among low mosses or on paths. Known from

Northern Europe and considered common. An ectomycorrhizal

ITS sequence from Tsuga heterophylla roots from Western

Canada, Vancouver, was also identified as this species (Table 3).

Differential diagnosis: Cortinarius carabus is recognised by the

small, dark brown fruit body and weakly dacryoid spores. The

similar looking C. cicindela has broader, ovoid spores, and the

more reddish brown C. coleoptera subglobose spores. From

the small, Pinus forest species of the sect. Hydrocybe or Incrus-

tati, C. carabus differs by darker colours or sparse veil, and dif-

ferent spores. Based on the genetic studies our C. carabus

seems to include two very closely related species (clade I

and II). So far, no morphological or ecological differences

have been found, and therefore, both species are included in

the list of specimens examined. The Latin description in-

cludes only the sequenced collections of C. carabus clade I.

Specimens examined: Estonia: Hiiumaa: Tahkuna, 13 Sep

2001, I.K. & T.N.S (H). d Finland: Uusimaa: Hanko, Lappohja,

10 Oct 1999, I.K.S (H); loc. cit., 3 Sep 2004, I.K. & M.Toivonen,

IK04-032 D (H); Loppi, S of Luutasuo, 4 Oct 2001, I.K. & T.N.,

IK01-514b (H); Porvoo, Vessolandet, 22 Oct 1998, I.K. & T.N.S

(H); Tammisaaari, opposite to Skarlandet, 5 Oct 2004, I.K. (H).

Etela-Hame: Ruovesi, Visuvesi N, 18 Oct 2004, I.K. (H); Etela-

Savo: Leivonmaki, Selanpohja, 10 Aug 1996, I.K., IK96-460 S

(H). Pohjois-Savo: Nilsia, Valkeinen, 1 Sep 2001, I.K., IK01-026DS (H). Kainuu: Kuhmo, Viiksimo, 30 Aug 1999, I.K., IK99-707DS

(H); Suomussalmi, Hossa, 16 Sep 2002, T.N. et al., TN02-612

(H); Kylmaluoma, 12 Aug 2002, I.K. (H); Sarkka, 16 Sep 2002,

T.N. et al., TN02-668 (H). Koillismaa: Kuusamo, Kantojoki, 2

Sep 2002, K.L. & T.N., TN02-507 (H); loc. cit., 21 Sep 2002, I.K. &

T.N., TN02-1130 (H); Oulanka biological station, 31 Aug 2002,

K.L. & T.N., TN02-361 (H); loc. cit., 18 Sep 2002 T.N. et al., TN02-

697b (H); loc. cit., 23 Sep 2002, T.N. et al., TN02-1112 (TUR); loc.

cit., 17 Sep 2005, T.N. et al., TN05-140 DS (holotype, H); Salla,

Kaunisharju recreation area, 2 Sep 2002, K.L. & T.N., TN02-

467 (H); Naruska, 1 Sep 2002, K.L. & T.N., TN02-428 (TUR).

Pera-Pohjanmaa: Pello, Orajarvi, 17 Aug 2001, I.K. (H); Tornio,

Korkiamaa, 31 Aug 1998, I.K., IK98-201 (H); Ylitornio,

Palorommas nature reserve area, 2 Sep 2004, K.L. & T.N.,

TN04-682 (H). Kittilan Lappi: Kittila, Raattama, 10 Aug 2001,

I.K. (H); Rauhala, 10 Aug 2001, I.K.S (H); Kolari, 15 Aug 2001,

I.K., IK01-025D (H); loc. cit., 16 Aug 2001, I.K. (H). Muonio, Jeris,

11 Aug 2001, I.K. (H); Keimion maja, 9 Aug 2001, I.K. (H);

Katkasuvanto, 9 Aug 2001, I.K. (H); Olos S, 9 Aug 2001, I.K. (H).

Enontekion Lappi: Enontekio, Kuttanen, 8 Aug 2001, I.K.

(H); Muotkajarvi, 9 Aug 2001, I.K. (H); Vahaniva, 8 Aug 2001,

I.K. (H). Inarin Lappi: Inari, Keskikompsio, 14 Aug 1995, I.K.,

IK95-355DS (H). d Sweden: Jamtland: Bracke, Bodsjo, 4 Sep

1997, I.K., IK97-680D (H); Ratan, Handsjo 31 Aug 2003, T.N.

et al., TN03-937S (H); Rojan, 3 Sep 2003, T.N. et al., TN03-1121

(H). Lule Lappmark: Gallivare, Muorjevaara, 14 Aug 2001, I.K.

(H). Torne Lappmark: Jukkasjarvi, Paittasjarvi, 13 Aug 2001, I.K.

(H); W side of the village of Jukkasjarvi, I.K., IK98-828 (H); loc.

cit., 13 Aug 2001, I.K. (H); ca 3 km W of the village of Syvajarvi,

13 Aug 2001, I.K.S (H); Kiiruna, Svappavaara, 13 Aug 2001, I.K.,

IK01-023D (H). Norrbotten: Jerttalompolo, 14 Aug 2001, I.K. (H).

Junosuando, Karhakkavuoma, I.K.S (H); Muonionalusta,

Muodoslompolo, 14 Aug 1998, I.K., IK98-827 (H); loc. cit., I.K.,

IK98-828 (H); Pajala, SW of Aareavaara, 17 Aug 1998, I.K.,

IK98-1051 (H); Kaunisjoensuu, 15 Aug 2001, I.K.S (H); Tarendo,

Kalixalven, 12 Aug 2001, I.K. (H); Koijuniemi, 14 Aug 2001, I.K. (H).

Cortinarius cicindela Kytov., Niskanen & Liimat. sp. nov.

(Figs 1, 3D, 4 J, 6, 12)




Etym: cicindela (Latin), a beetle genus Cicindela belonging to

Coleoptera. The name refers to the closely related C. coleoptera.

Pileus 1–4 cm, obtuse conicus, tum explanatus, umbonatus,atroumbrinus, dein quasi nigricans, hygrophanus. Lamellae sub-distantae, brunneae vel umbrinae. Stipes 4–10� 0.2–0.4 cm, cylin-draceus, primo griseoalbide fibrillosus, dein obscure brunneusvel atrobrunneus, velo albido, sparsissimo. Caro atrobrunnea.Odor nullus. Exsiccata nigrescentia. Sporae 7.5–9� 5–6.5 mm, ovataevel obovatae, valde verrucosae. In silvis coniferis humidis velmediocriter humidis et rariore in silvis coniferis aridis etsemiaridis.

Typus: Norway: Troms: Bardu commune, Setermoen, 3.6 km S bythe road E6 from the Fosseng crossing, NNE slope, planted, mossy,mesic spruce forest (Picea abies), somewhat eutrophic, 8 Aug 2006,Liimatainen & Niskanen F06-036 (H d holotypus, NY d isotypus).GenBank no. EU266669.

Pileus: 1–4 cm, conical to hemispherical, very soon almost

plane with an acute umbo, up to 1/2 pellucid-striate, very

weakly innately fibrillose, dark brown, hygrophanous. Lamel-

lae: medium spaced to fairly distant, strongly to weakly emar-

ginate, medium thick, moderately broad to broad, dark brown,

edge concolorous or whitish. Stipe: 4–10� 0.2–0.4 cm, cylindri-

cal, whitish fibrillose, soon brown to dark reddish brown. Uni-

versal veil: whitish, very sparse. Mycelium: white, sparse.

Context: dark brown, darkening with age, often first at the

base of the stem. Smell: indistinct. Exsiccata: greyish black.

Spores: 7.5–9� 5–6.5 mm, Q¼ 1.34–1.56, �X¼ 8–8.4� 5.5–

5.8 mm, �XQ¼ 1.41–1.47 (18 collections, Fig 4), weakly ovoid

to weakly obovoid, strongly, coarsely verrucose, very

strongly at the apex, weakly to moderately dextrinoid,

dark, in KOH brown and dark-coloured. Lamellar trama hy-

phae: in MLZ and KOH olivaceous brownish, strongly zebra-

striped, and somewhat spot-like encrusted. Lamellar edge:

fertile, marginal cells clavate, 11–25� 5–11 mm, almost con-

colorous to dark umber in MLZ, pigment mostly intracellular

or in cell walls, seldom extra-cellular; in KOH fairly hyaline

to olivaceous brown. Basidia: 4–spored, 30–44� 7–10 mm, in

MLZ olivaceous brown, in KOH almost hyaline to olivaceous

brown, with granulose content. Pileipellis: with a fairly thin

epicutis of 3–6 mm wide, granulose, spot-like encrusted, in

MLZ and KOH sepia, upper hyphae, and wider (>10 mm),

very finely scabrous, paler sepia lower ones. The transition

to the hypoderm is obscure. Hypoderm present, but fairly

poorly differentiated, hyphae of the hypoderm 35–60� 15–

25 mm, in MLZ and KOH light coloured. Hyphae of the context

6–18 mm wide, sometimes zebra-striped encrusted and/or

with sepia spots. Stipitipellis hyphae: at the stipe top pale oli-

vaceous to pale sepia, weakly to strongly encrusted. Clamp

connections: present.


(based on a total of eight sequences, including the type,

Table 2). There are four infraspesific variation sites in the se-

quences, including two base and one length polymorphisms

sites that are intragenomic and three infraspecific polymor-

phism sites. The biggest pairwise distance is three changes.

The amount of the variation is quite high for one taxon

and the studied collections can be divided into two, or

even three separate groups based on the variable sites.

Thus it is possible that the studied collections of the

Cortinarius cicindela included several taxa, but more studies

would be needed for further conclusions. We have also three

unpublished partial ITS sequences, which support these sub-

groups. We did not find any sequences of this species in the

public databases. The difference compared with C. carabus

clade I is at least eight evolutionary events (four indels, three

transitions and one transversion) and to C. carabus clade II at

least ten evolutionary events (four indels, five transitions

and one transversion).

Type material: The holotype includes four whole and two

half fruit bodies, and the isotype includes three fruit bodies,

all in good condition.

Ecology and distribution: In Picea or Pinus bogs among Sphag-

num, in swampy depressions of Picea forests, but also in mesic

Picea forests and dry Pinus heath forests, often scanty. The

fruiting period is very long, often beginning as early as July.

Known in Northern Europe, occasional.

Differential diagnosis: Typical for Cortinarius cicindela is the

very slender, dark brown fruit body, weakly innately fibril-

lose pileus, somewhat ovoid, strongly verrucose spores,

and the very strongly encrusted lamellar trama hyphae.

When growing early in the season in paludified depressions

in spruce forests among Sphagnum, this species can be

easily recognised, but also easily overlooked because of

the solitary growing habit. In dry forests it can be mixed

with C. carabus or C. coleoptera, but the former has narrower

spores, and the latter is more reddish brown, often some-

what bigger, and has subglobose spores. Based on the

genetic studies C. cicindela may include a few taxa. So far,

no morphological or ecological characteristics have been

found to support these clades and they are treated here

as one species.

Fig 12 – Distribution of Cortinarius cicindela in North Europe




Specimens examined: Finland: Uusimaa: Kyrkslatt, Porkkala,

R. Skyten, 3122S (H). Satakunta: Ikaalinen, Seitseminen National

Park, 8 Sep 2005, K.L. & T.N., TN05-070DS (H). Etela-Hame: Juupa-

joki, Hyytiala, 6 Sep 2005, H. TuovilaS (H); Lammi, Evo, 5 Sep

1979, M. Korhonen & R. TuomikoskiS (H); Virrat, Monoskyla, 20

Jul 1995, P.K & I. K., IK95-037DS (H); loc. cit., P.K & I. K., IK95-043

(H). Pohjois-Hame: Konnevesi, Varuskunta, 7 Sep 2002, I.K.S

(H); Uurainen, Murtopera, 10 Jul 1998, I.K., IK98-042S (H); Virrat,

Hauhuu, 16 Jul 1996, I.K., IK96-111S (H); loc. cit., 19 Jul 1996, I.K.,

IK96-133 (H); loc. cit., 21 Jul 1996 I.K., IK96-184 (H). Pohjois-

Karjala: Ilomantsi, Niemijarvi, 24 Sep 1997, I.K., IK97-1780 DS

(H). Kainuu: Hyrynsalmi, Paljakka, 28 Aug 1999, I.K. (H);

Puolanka, Hepokongas, 9 Aug 2003, T.N. et al., TN03-173DS

(H); Pihlajavaara, 15 Sep 2005, K.L. & T.N., TN05-118S (H);

Suomussalmi, Suolijarvi, 14 Sep 1997, I.K., IK97-1424S (H). Koil-

lismaa: Kuusamo, Oulanka biological station,18 Sep 2002, T.N.

et al., TN02-789 (H). Pera-Pohjanmaa: Pello, Orajarvi, 17 Aug

2001, I.K.S (H); Rovaniemi, Kaihua, 23 Aug 1999, I.K., IK99-

592DS (H); Ylitornio, Meltosjarvi, 21 Aug 1999, I.K., IK99-463

(H). Kittilan Lappi: Kittila, Aakenustunturi N, 29 Jul 2001, I.K.

et al.S (H); Muonio, Akasjarvi, 29 Jul 2001, I.K. et al. (H). Sompion

Lappi: Sodankyla, ca 1 km NE of the river Tankajoki and Tank-

apirtti, 13 Aug 1995, I.K., IK95-303DS (H). d Norway: Troms:

Bardu, Setermoen, 5 Aug 2006, K.L. & T.N., TN06-011S (H); loc.

cit., 8 Aug 2006, K.L. & T.N., TN06-036DS (holotype, H). d Slova-

kia: Liptovska kotlina basin, Va�zek, Va�zecke luky, 29 Sep 2003,

T.N. et al., TN03-1650S (H).

Cortinarius gentilis (Fr.) Fr., Epicr. Syst. mycol.: 297 (1838).

(Figs 1, 4 K, 5)

Basionym: Agaricus gentilis Fr., Syst. mycol. 1: 212 (1821).

Type: Picture 159 in Icones selectae, Fries 1867–84.

Illustrations: Brandrud et al. (1992, pl. B31), Consiglio et al.

(2004: pl. B67), Nylen (2000: 439), Ryman & Holmasen (1992:

492), Salo et al. (2006: 201).

Pileus: 3–6 cm, conical to hemispherical, later low convex to

plane, often with a small umbo, saturated reddish brown to

yellow brown, soon yellow due to drying, strongly hygropha-

nous. Lamellae: distant, strongly to weakly emarginate, fairly

thick, broad, yellow brown to red brown. Stipe: 5–12� 0.3–

0.8 cm, cylindrical, base sometimes somewhat rooting, yellow

to yellow brown, darker towards the base. Universal veil: yel-

low, forming distinct girdles on stipe. Mycelium: white. Context:

red brown. Smell: raphanoid, sometimes rubber-like. Exsiccata:

brownish yellow.

Spores: 7.5–9.5� 6–7.5 mm, Q¼ 1.20–1.34, �X¼ 7.9–8.8� 6.3–

6.8, �XQ¼ 1.25–1.30 (five collections, Fig. 4), subglobose to

somewhat elongated, thin-walled, weakly, densely, sharply

verrucose, weakly dextrinoid, in KOH brown. Lamellar trama

hyphae: in MLZ and KOH yellow to orange yellow, densely sca-

brous, small-spotted. Lamellar edge: quite fertile, marginal cells

clavate, 18–28� 5–12 mm, concolorous to more orange in MLZ

and KOH. Basidia: 4–spored, 34–50� 8–11 mm, in MLZ yellowish

to brownish orange, in KOH yellow, often granulose. Pileipellis:

with fairly thin epicutis of 3–7 mm wide, smooth to very finely

scabrous, somewhat granulose, in MLZ pale yellowish to yel-

lowish brownish, in KOH pale yellow to yellow upper hyphae,

lower ones a bit wider (10 mm), very finely to zebra striped

encrusted. Hypoderm fairly well-developed, with somewhat

smaller, 35–50� 15–20 mm, hyphae than in other Brunnei spe-

cies. Hyphae of the context 6–15 mm wide, often zebra-striped

encrusted. Stipitipellis hyphae: brownish yellow to orange yel-

low, strongly encrusted (zebra-stripedþ small-spotted), veil

hyphae also encrusted. Clamp connections: present.


(based on a total of three sequences, Table 2). The IK99-

396 collection has three intragenomic length polymorphism

sites and the collection TN06-250 (clade II) has three base

changes compared with CFP’s photo collection. Infraspecific

variation was observed at seven sites and the maximum

pairwise distance between the sequences was four evolu-

tionary events. From the public databases we found four

other sequences belonging to this species group (Table 3).

When comparing all the sequences together they seemed

to form three groups, consistently separated by three evo-

lutionary events. This indicates that the data include sev-

eral close sister species, but more specimens would be

needed to study this complex more in detail. The closest

relatives for Cortinarius gentilis are C. pseudorubricosus and

C. ectypus, which differ by at least 28 evolutionary events.

The difference compared with C. limonius, the type species

of the sect. Limonii, in which C. gentilis was previously clas-

sified, is more than 50 evolutionary events and the ITS re-

gion of C. limonius is 80 base pairs longer than the one of

C. gentilis.

Ecology and distribution: In dry to mesic coniferous forests

with Pinus and Picea, very common. Known from Northern

Europe. The ITS sequences deposited in the public databases

originate from UK, Russia, and USA and were also assigned

to this species group (Table 3).

Differential diagnosis: Cortinarius gentilis is easily recognised

by the slender, vivid yellow brown to reddish brown fruit

body, rooting stipe, and fairly big, subglobose spores. Based

on our own and GenBank sequences, C. gentilis seems to in-

clude some closely related species, but more studies are

needed for further conclusions. Traditionally, C. gentilis is

placed near the other brightly coloured species in the subge-

nus Cortinarius or Leprocybe, in the sect. Limoni (Brandrud

et al. 1992; Moser 1983); however, recent phylogenetic studies

have shown that it belongs to the subgenus Telamonia, pre-

sumably near C. brunneus (e.g. Høiland & Holst-Jensen 2000;

Peintner et al. 2004, Fig 1). Morphologically the hygrophanous

pileus, subglobose spores, raphanoid, sometimes rubber-like

smell, strongly encrusted lamellar trama hyphae and the

structure of pileipellis are common with some Brunnei species,

but more thorough morphological studies of telamonioid spe-

cies are needed, to identify all morphological charactersistics,

which may support this classification.

Specimens examined: Finland: Satakunta: Sakyla, Virttaan-

kangas, 18 Oct 2006, K.L. & T.N., TN06-250 DS (H). Pohjois-

Hame: Multia, Uittamonkoski, 12 Sep 2005, I.K.S (H). Pera-

Pohjanmaa: Rovaniemi, Savijoki, 19 Aug 1999, I.K., IK99-396 DS

(H). d Norway: Oppland: Dokka, Vest-Torpa, 15 Sep 1983, H.

Lindstrom et al., CFP178 DS (S). Sør-Trøndelag: Orkdal, 30 Jul

1974, P.K. & I. K., IK4138 (H).




Discussion

Brunnei

All the species presented above are morphologically well

delimited. The majority can most unambiguously be recog-

nised by their spores, although the size and colour of the fruit

body, the nature of the hygrophanous properties of the pileus,

the amount and colour of the veil, and ecology are also useful

characteristics in species identification.

Most of the species are common and widespread in north-

ern coniferous forests; however, Cortinarius albogaudis seems

to be very rare and has only been found in five areas from cal-

careous spruce forests. C. ectypus is also rare and C. pseudoru-

bricosus less common, but the ecological factors restricting

their distribution are not known.

Although many Brunnei species are common, sequences of

only six of the 11 species were found in the public databases

indicating a poor coverage of the group. In addition, only

35 % of the sequences were correctly named. Interestingly,

two species, C. pseudorubricosus and C. carabus, had identical

or nearly identical ectomycorrhizal sequences from Vancou-

ver, in Western North America. Matches for several other spe-

cies were also found from Scotland, Central Europe, and

Russia either from sequences from ectomycorrhizal root tips

or fruiting bodies. Sequences from different sources provide

a new and useful way to study the distribution and ecology

of the species in addition to traditional approaches.

The relationships of the species in the section Brunnei were

not solved with ITS data, but certain groups were stable in all

the analyses. The small species C. carabus, C. cicindela, and C.

coleoptera formed one group, and C. albogaudis and C. pseudor-

ubricosus another. Additional genetic data would be needed

to resolve the rest of the relationships.

The section Brunnei represents a typical example of a sec-

tion in subgenus Telamonia. The limits of the section Brunnei

have been obscure and the section has also previously in-

cluded members from other sections. It is evident that a revi-

sion of the sections is needed to gain a more natural

classification for the sections in the subgenus Telamonia. The

number of the new species also reflects the amount of un-

known species in Telamonia. When we also count the excluded

species, the percentage of new species is ca 50 %.

Key to the species of Cortinarius section Brunnei

1 Fruiting body vivid yellow reddish brown.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . gentilis

Fruiting body� brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2 (1) Most spores >5.5 mm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Most spores <5.5 mm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

3 (2) Stipe <4 mm, cylindrical.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cicindela

Stipe >4 mm, usually at least slightly clavate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4 (3) Most spores <6 mm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Most spores >6 mm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

5 (4) Mainly with Pinus; pileus pale brown to brown; stipe 8–15 thick; spores 7–8.5� 5.5–6.5 mm. . . . . . . . . . . . clarobrunneus

With Picea; pileus greyish brown to reddish brown, sometimes with bluish tints; stipe 4–8 mm thick; spores

7.5–9� 5–6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. caesiobrunneus

6 (4) With Picea; stipe 8–15 mm thick; universal veil usually forming a distinct girdle on the stipe; spores 8–9.5� 6–7 mm,

Q¼ 1.24–1.44 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . brunneus

Usually with Pinus; stipe 4-11 mm broad; universal veil often sparse, rarely forming a distinct girdle on the stipe; spores

8–10� 6–7 mm, Q¼ 1.3–1.54 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. glandicolor

7 (2) Pileus strongly hygrophanous; stipe 2–7 mm thick, cylindrical to tapering downwards or clavate . . . . . . . . . . . . . . . . ..8

Pileus faintly hygrophanous or with hygrophanous streaks; stipe 6–15 mm thick, clavate to cylindrical . . . . . . . . . . ..11

8 (7) Spores obovoidly subglobose, 6–8� 5–6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .coleoptera

Spores ovoid, ellipsoid to weakly dacryoid, >7.5 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..9

9 (8) Stipe 4–8 mm thick, cylindrical to clavate; lamellar trama hyphae not or finely zebra striped encrusted. caesiobrunneus

Stipe 2–4 mm thick, cylindrical; lamellar trama hyphae very strongly zebra- and spot-like encrusted . . . . . . . . . . . . ..10

10 (9) Spores 7.5–9� 5–6.5 mm, weakly ovoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .cicindela

Spores 7–9� 4.5–5.5 mm, weakly dacryoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . carabus

11 (7) Spores ellipsoid to obovoid-ellipsoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pseudorubricosus

Spores subglobose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..12

12 (11) Stipe clavate; universal veil white, distinct, forming a girdle on the stipe; spores 6–7.5� 5–6 mm, obovoid-subglobose,

echinate-verrucose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. albogaudis

Stipe cylindrical; universal veil brownish, indistinct; spores 5.5–7� 4.5–6 mm, ovoid to ovoid-subglobose, finely

verrucose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .ectypus



Species level taxonomy

Based on our results, closely related species (C. brunneus and C.

glandicolor, C. carabus and C. cicindela, Fig 1) are not always rec-

ognised in phylogenetic analyses, although they have differ-

ences in ITS regions. Two main reasons for this are: (1) ITS

regions are not variable enough, and (2) information is lost

or overlooked during the aligning and/or analysis.

The first problem would be solved by using more variable

regions. In Cortinarius, RPB1 has been shown to have more

informative characters than ITS regions (Frøslev et al. 2005).

Another candidate could be IGS, which has been used, e.g. in

Armillaria (Terashima et al. 1998). The second problem is partly

due to the fact that ITS sequences of different species are of-

ten variable in length and may also contain highly divergent

regions. Therefore, alignments are seldom unambiguous and

information is lost when removing the ambiguous regions be-

fore the analyses. An attempt to resolve this problem has been

tried by developing another kind of approach in which phylo-

genetic analyses and alignment of DNA sequences are

performed simultaneously [e.g. POY (Gladstein & Wheeler

2001; Wheeler 1996)]. These methods have not been widely

used, partly because they require considerable computing

time. Another way, used in this study for C. brunneus and

C. glandicolor, is to analyse only very closely related species

together.

A further problem when choosing to align sequences is the

treatment of gaps. By using the missing data option, some in-

formation is overlooked, whereas treating the gaps as a fifth

characteristic, overemphasises their significance. Using the re-

sults from our studies, it is evident that the difference in

topology and support values is notable between these two

approaches, especially with closely related species where

treating the gaps as fifth characteristics gave results most con-

gruent with the morphological data. A compromise between

the methods would probably be more productive, and for this

reason different gap coding methods have been introduced

(e.g. Hibbett et al. 1995; Høiland & Holst-Jensen 2000), but

most of them are not very easy to use unambiguously with

large datasets and were, therefore, not used in this study.

Despite species not being recognised in phylogenetic anal-

yses for the reasons mentioned above, they may still be sup-

ported by the sequence data. In the 11 morpho-genetic

species recognised in this study, the pairwise infraspecific dif-

ferences are smaller than the interspecific ones, correlating

well with the results by Frøslev et al. (2007). For example, for

the two species pairs not separated in all the analyses (Fig 1)

the values are (1) C. brunneus and C. glandicolor one infraspe-

cific and five interspecific differences, and (2) C. carabus (clade

I) and C. cicindela three infraspecific and eight interspecific

differences and discontinuous groups are formed. For some

reason the support value of C. albogaudis was also low, al-

though there is no infraspecific variation and the difference

to C. pseudorubricosus is over 18 evolutionary events.

Based on the differences in the infraspecific variation of

the ITS regions we could preliminary classify the morpho-ge-

netic species of sect. Brunnei into four different categories, al-

though small sample size has to be taken into account. The

sequences from the public databases were not included.

Some of them had their own unique nucleotide differences,

but we were not able to check the chromatographic data to

see whether the differences really existed.

The first category includes C. albogaudis and C. ectypus, and

is characterised by having no genetic variation in the ITS re-

gions. C. pseudorubricosus, C. brunneus, C. clarobrunneus, and

C. coleoptera form the second group, where all the infraspecific

variation is intragenomic polymorphism, i.e. no characteristic

sites exist where two sequences would have different charac-

ter states. In the third group [C. glandicolor and C. caesiobrun-

neus (clade I and II)] the infraspecific divergence has gone

further. Different sequences occur within the species, but in

all the characteristic sites that differ, intermediates (intrage-

nomic polymorphism) also appear, and we treated these as

one genetic species although further studies with more vari-

able DNA regions would be needed to clarify this. The differ-

ences did not seem to correlate with geographical distance.

Species in the last group [C. gentilis (clade I and II), C. carabus

(clade I and II) and C. cicindela] have one or more characteristic

sites with discontinuous variation (no intragenomic polymor-

phisms). This indicates that they might include several spe-

cies, and it would probably be best to refere them as species

groups. More studies would be needed to find out whether

morphological or ecological characteristics supporting these

groups exist. If the species in group 4 would be divided into

several species, the groups would be genetically more coher-

ent with other species, e.g. C. gentilis clade I (Fig 1) and C. car-

abus clade I would be in group 2, and C. carabus clade II in

group 1. Dividing C. cicindela is not so obvious and would

require more data.

All the genetic studies in the genus Cortinarius have so far

treated the amount of variation observed in group 4 to be in-

fraspecific (e.g. Kytovuori et al. 2005; Lindstrom et al. 2008;

Moser & Peintner 2002a). However, Garnica et al. (2003) and

Frøslev et al. (2007) found that morphologically accepted C.

atrovirens and C. ionochlorus had identical ITS sequences. Fur-

thermore, we already have evidence that, e.g. C. laniger and

C. solis-occasus (GenBank nos AF325591 and AY669696), only

have one discontinuous characteristic site in the ITS regions

to separate them. This indicates that ITS sequences are prob-

ably not variable enough to clearly separate all the Cortinarius

species. Therefore, it also seems that for some Cortinarius spe-

cies the ITS region is not suitable for species-level recognition

(barcoding).

In this study we decided to only describe those species that

can be delimited based on molecular and morphological data

(morpho-genetic species recognition); however, it seems that,

presumably, some of the species include several ITS species

(at least C. carabus, possibly also C. cicindela and C. caesiobrun-

neus). Our data do not support the broader morphological spe-

cies concept used in Brandrud et al. (1989). Taxa previously

regarded as varieties or subspecies, appeared to be genetically

and morphologically well delimited, and therefore, consid-

ered as species, like C. clarobrunneus. In the taxonomy of Corti-

narius, taxa below the species level have been widely used (e.g.

Brandrud et al. 1989, 1992, 1994, 1998; Bidaud et al. 1995, 1997).

In the extensive study by Frøslev et al. (2007), as well as in our

study, no infraspecific, morphologically distinct taxa were rec-

ognised. These results raise the question as to whether we are






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actually able to recognise taxa below the species level with the

charactersistics we have in use.

Acknowledgements

We thank Jarno Tuimala (CSC) for assistance with computer

programs, Heino Vanska for revision of the Latin descrip-

tions, Ernest Emmett for help with the English, Hakan

Lindstrom for lending his material and providing his field

notes of Cortinarius pseudorubricosus, Teuvo Ahti for helping

with the questions regarding nomenclature, and the referees

for valuable comments. This work was supported by the

Ministry of Environment, Finland (YM195/5512/2002-5 &

YM196/5512/2002-5) and the University of Helsinki Research

Foundation.

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Cortinarius sect. Brunnei ( Basidiomycota, Agaricales) in North Europe

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