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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: tuula.niskanen@cortinarius.fi
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
Author's personal copy
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
Author's personal copy
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.
Author's personal copy
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)
Cortinarius sect. Brunnei in North Europe 185
Author's personal copy
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.
186 T. Niskanen et al.
Author's personal copy
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
Cortinarius sect. Brunnei in North Europe 187
Author's personal copy
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.
188 T. Niskanen et al.
Author's personal copy
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.
Cortinarius sect. Brunnei in North Europe 189
Author's personal copy
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
C. brunneus Scotland, Moray, Culbin
Forest
P. sylvestris and P. nigra,
few Betula pubescens
AMG DG25 0 UDB001541
C. brunneus Scotland, Moray, Culbin
Forest
P. sylvestris and P. nigra,
few Betula pubescens
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
P. sylvestris and P. nigra,
few Betula pubescens
AMG 2006/217 0 UDB002396
C. umbrinolens Scotland, Moray, Culbin
Forest
P. sylvestris and P. nigra,
few Betula pubescens
AMG DG05-64 0 UDB001539
C. umbrinolens Scotland, Moray, Culbin
Forest
P. sylvestris and P. nigra,
few Betula pubescens
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.
190 T. Niskanen et al.
Author's personal copy
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.
Cortinarius sect. Brunnei in North Europe 191
Author's personal copy
(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.
192 T. Niskanen et al.
Author's personal copy
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);
Cortinarius sect. Brunnei in North Europe 193
Author's personal copy
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)
MycoBank no.: MB 511425
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.
ITS regions (including 5.8S region): 526–527 bases long
(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.
194 T. Niskanen et al.
Author's personal copy
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.
ITS regions (including 5.8S region): 532 bases long (based on
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.
Cortinarius sect. Brunnei in North Europe 195
Author's personal copy
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.
196 T. Niskanen et al.
Author's personal copy
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.
ITS regions (including 5.8S region): 538 bases long (based on
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.
ITS regions (including 5.8S region): 512 bases long (based on
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.
Cortinarius sect. Brunnei in North Europe 197
Author's personal copy
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.
ITS regions (including 5.8S region): 533–534 bases long
(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.
198 T. Niskanen et al.
Author's personal copy
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)
MycoBank no.: MB 511422
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
according to the material examined.
Cortinarius sect. Brunnei in North Europe 199
Author's personal copy
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)
MycoBank no.: MB 511423
200 T. Niskanen et al.
Author's personal copy
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.
ITS regions (including 5.8S region): 530–534 bases long
(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
according to the material examined.
Cortinarius sect. Brunnei in North Europe 201
Author's personal copy
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.
ITS regions (including 5.8S region): 527–530 bases long
(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).
202 T. Niskanen et al.
Author's personal copy
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
Cortinarius sect. Brunnei in North Europe 203
Author's personal copy
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
204 T. Niskanen et al.
Author's personal copy
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.
r e f e r e n c e s
Aanen DK, Kuyper TW, Boekhout T, Hoekstra RF, 2000. Phyloge-netic relationships in the genus Hebeloma based on ITS1 and 2sequences, with special emphasis on the Hebeloma crustulini-forme complex. Mycologia 92: 269–281.
Ammirati J, Garnica S, Halling RE, Mata M, Mueller GM,Carranza J, 2007. New Cortinarius species associated withQuercus and Comarostaphylis in Costa Rica. Canadian Journal ofBotany 85: 794–812.
Bidaud A, Moenne-Loccoz P, Reumaux P, 1995. Atlas des Cortin-aires. Pars VII. Editions Federation mycologique Dauphine-Sa-voie, La Roche-sur-Foron.
Bidaud A, Moenne-Loccoz P, Reumaux P, 1997. Atlas des CortinairesHors-Serien no. In: Les Cortinaires Hinnuloides, 1. Editions Fed-eration mycologique Dauphine-Savoie, La Roche-sur-Foron.
Brandrud TE, Lindstrom H, Marklund H, Melot J, Muskos S, 1989.Cortinarius Flora Photographica I (Swedish version). CortinariusHB, Matfors.
Brandrud TE, Lindstrom H, Marklund H, Melot J, Muskos S, 1992.Cortinarius Flora Photographica II (Swedish version). CortinariusHB, Matfors.
Brandrud TE, Lindstrom H, Marklund H, Melot J, Muskos S, 1994.Cortinarius Flora Photographica III (Swedish version). CortinariusHB, Matfors.
Brandrud TE, Lindstrom H, Marklund H, Melot J, Muskos S, 1998.Cortinarius Flora Photographica IV (Swedish version). CortinariusHB, Matfors.
Consiglio G, Antonini D, Antonini M, 2004. Il genere Cortinarius inItalia II. Associazione Micologica Bresadola, Luglio.
Consiglio G, Antonini D, Antonini M, 2006. Il genere Cortinarius inItalia IV. Associazione Micologica Bresadola, Luglio.
Den Bakker HC, Gravendeel B, Kuyper TW, 2004. An ITS phy-logeny of Leccinum and an analysis of the evolution ofminisatellite-like sequences within ITS1. Mycologia 96:102–118.
Dahncke RM, 1993. 1200 Pilze. AT Verlag, Stuttgart.Edgar RC, 2004. MUSCLE: multiple sequence alignment with high
accuracy and high throughput. Nucleic Acids Research 32:1792–1797.
Favre J, 1960. Catalogue descriptif des champignons super-ieurs de la zone subalpine du Parc National Suisse. Ergebr.wiss. Untersuchungen schweiz. Nationalpark NF 6, 42: 513and 588.
Fries EM, 1821. Systema mycologicum I. Lundae.Fries EM, 1838. Epicrisis systematis mycologici. Upsaliae.Fries EM, 1863. Monographia Hymenomycetum Sueciae. Upsaliae.
Fries EM, 1874. Hymenomycetes Europaei. Upsaliae.Frøslev T, Matheny P, Hibbett D, 2005. Lower level relationships in
the mushroom genus Cortinarius (Basidiomycota, Agaricales):a comparison of RBP1, RPB2 and ITS phylogenies. MolecularPhylogenetics and Evolution 37: 602–618.
Frøslev TG, Jeppesen T, Laessøe T, 2006. Seven new calochroidand fulvoid species of Cortinarius. Mycological Research 110:1046–1058.
Frøslev TG, Jeppesen TS, Læssoe T, Kjøller R, 2007. Molecularphylogenetics and delimitation of species in Cortinarius sec-tion Calochroi (Basidiomycota, Agaricales) in Europe. MolecularPhylogenetics and Evolution 44: 217–227.
Garnica S, Weiß M, Oberwinkler F, 2003. Morphological and mo-lecular phylogenetic studies in South American Cortinariusspecies. Mycological Research 107: 1143–1156.
Garnica S, Weiß M, Oertel B, Oberwinkler F, 2005. A framework fora phylogenetic classification in the genus Cortinarius (Basidio-mycota, Agaricales) derived from morphological and moleculardata. Canadian Journal of Botany 83: 1457–1477.
Gardes M, Bruns TD, 1993. ITS primers with enhanced specifityfor basidiomycetes application to the identification of mycor-rhizae and rusts. Molecular Ecology 2: 113–118.
Gladstein D, Wheeler W, 2001. POY (Phylogeny Reconstruction viaDirect Optimization of DNA data), version 2.7. American Museumof Natural History, NY.
Goloboff PS, Farris S, Nixon K, 2000. TNT (Tree analysis using NewTechnology) (BETA) ver. 1.0. Published by the authors Tucuman,Argentina.
Hansen L, Knudsen H (eds) 1992. Nordic Macromycetes, Vol. 2 (Pol-yporales, Boletales, Agaricales, Russulales). Nordsvamp,Copenhagen.
Hibbett DS, Fukumasa-Nakai Y, Tsuneda S, Donoghue MJ, 1995.Phylogenetic diversity in shiitake inferred from nuclear ribo-somal DNA sequences. Mycologia 87: 618–638.
Holmgren PK, Holmgren NH, Barnett LC, 1990. Index herbariorum1. In: The herbaria of the world, 8th edn. Regnum Vegetabile, 120:1–693.
Huelsenbeck JP, Ronquist F, 2003. MRBAYES 3: Bayesianphylogenetic inference under mixed models. Bioinformatics 19:1572–1574.
Høiland K, Holst-Jensen A, 2000. Cortinarius phylogeny and pos-sible taxonomic implications of ITS rDNA sequences.Mycologia 92: 694–710.
Kuyper TW, 1988. Specific and intraspecific delimitation. In:Bas C, Kuyper TW, Noordeloos WE, Vellinga EC (eds), FloraAgaricina Neerlandica I AA Balkema, Rotterdam, pp. 30–37.
Kuhner R, Romagnesi H, 1953. Flore analytique des champignonssuperieurs. Masson et Cie, Paris.
Kytovuori I, Niskanen T, Liimatainen K, Lindstrom H, 2005.Cortinarius sordidemaculatus and two new related species, C.anisatus and C. neofurvolaesus, in Fennoscandia (Basidiomycota,Agaricales). Karstenia 45: 33–49.
Larsson E, Jacobsson S, 2004. Controversy over Hygrophorus cossussettled using ITS sequence data from 200 year-old typematerial. Mycological Research 108: 781–786.
Lindstrom H, Soop K, 1999. Quelques petits Telamonia calciphiles.Journal des J.E.C. 2: 40–60.
Lindstrom H, Bendiksen E, Bendiksen K, Larsson E, 2008. Studiesof the Cortinarius saniosus (Fr. : Fr.) Fr. complex and a newclosely related species, C. aureovelatus (Basidiomycota, Agari-cales). Sommerfeltia 31: 139–159.
Liu YJ, Rogers SO, Ammirati JF, 1997. Phylogenetic relationships inDermocybe and related Cortinarius taxa based on nuclear ribo-somal DNA internal transcribed spacers. Canadian Journal ofBotany 75: 519–532.
Marchand A, 1983. Champignons du nord et du midi 7. Soci-ete Mycologique des Pyrenees MediterraneennesPerpignan.
Cortinarius sect. Brunnei in North Europe 205
Author's personal copy
Matheny PB, Ammirati JF, 2006. Cortinarius lucorum (Fr.) Karst.,a Populus associate from North America. Pacific Nortwest Fungi1: 1–10.
Melot J, 1989. Combinationes et taxa nova. Documents Mycologiques20: 93–100.
Moenne-Loccoz P, Reumaux P, Henry R, 1990. Atlas desCortinaires I. Editions Federation Mycologique Dauphine-Savoie, Annecy.
Moser MM, 1983. Die Rohrlinge und Blatterpilze. In: Gams H (ed),Kleine Kryprogamenflora, 2b/2, 5th edn. Gustav Fischer Verlag,Stuttgart, pp. 1–534.
Moser MM, Peintner U, 2002a. The species complex Cortinariusscaurus–C. herpeticus based on morphological and moleculardata. Micologia e Vegetation Mediterranea 17: 3–17.
Moser MM, Peintner U, 2002b.Die phylogenetischenBeziehungen derCortinarius aureopulverulentus-Gruppe. Journal des J.E.C. 4: 28–38.
Niskanen T, 2007 [‘2008’]. Cortinarius Subgenus Telamonia p.p. inNorth Europe. Ph.D. thesis, University of Helsinki.
Niskanen T, Liimatainen K, Kytovuori I, 2006. Taxonomy, ecologyand distribution of Cortinarius rubrovioleipes and C. hinnuleoar-millatus (Basidiomycota, Agaricales) in Fennoscandia. Karstenia46: 1–12.
Niskanen T, Kytovuori I, Liimatainen K, 2007. Cortinarius sect.Brunnei (Basidiomycota, Agaricales) in North Europe. Manuscriptdistributed with Niskanen 2007: 38.
Nylen B, 2000. Svampar i Norden och Europa. Natur och Kultur/LTsforlag.
Peintner U, Ladurner H, Simonini G, 2003. Xerocomus cisalpinus sp.nov., and the delimitation of species in the Xerocomus chrys-enteron complex based on morphology and rDNA-LSU se-quences. Mycological Research 107: 659–679.
Peintner U, Moncalvo J-M, Vilgalys R, 2004. Towards a better un-derstanding of the infrageneric relationships in Cortinarius(Agaricales, Basidiomycota). Mycologia 96: 1042–1058.
Ryman S, Holmasen, 1992. Svampar en Falthandbok, TredjeUpplagan. Interpublishing, Stockholm.
Reumaux P, 1982. Etude de quelques Cortinaires de l’Ardennefrancaise. Bulletin Trimestriel de la Societe Mycologique de France98: 319–349.
Salo P, Niemela T, Salo U, 2006. Suomen sieniopas. WSOY, Helsinki.Seidl MT, 2000. Phylogenetic relationships within Cortinarius
subgenus Myxacium, sections Defibulati and. Myxacium.Mycologia 92: 1091–1102.
Soop K, 2004. Cortinarius in Sweden, 9th revised edn. EditionsScientrix, Mora.
Taylor JW, Jacobson DJ, Kroke S, Kasuga T, Geiser DM,Hibbett DS, Fisher MC, 2000. Phylogenetic species recognitionand species concepts in fungi. Fungal Genetics and Biology 31:21–32.
Terashima K, Cha JY, Yajima T, Igarashi T, Miura K, 1998. Phylo-genetic analysis of Japanese Armillaria based on the intergenicspacer (IGS) sequences of their ribosomal DNA. EuropeanJournal of Forest Pathology 28: 11–19.
Thompson JD, Higgins DG, Gibson TJ, 1994. Clustal W: improv-ing the sensitivity of progressive multiple sequence align-ment through sequence weighting, position specific gappenalties and weight matrix choice. Nucleic Acid Research 22:4673–4680.
Vellinga EC, de Kok RPJ, Bruns D, 2003. Phylogeny and taxonomyof Macrolepiota (Agaricaceae). Mycologia 95: 442–456.
Wheeler WC, 1996. Optimization alignment. The end of mul-tiple sequence alignment in phylogenetics? Cladistics 12:1–9.
White TJ, Bruns T, Lee S, Taylor J, 1990. Amplification and directsequencing of fungal ribosomal RNA genes for phylogenetics.In: Michael AJ, Gelfand DH, Sninsky JJ, White TJ (eds), PCRprotocols: a guide to the methods and applications Academic Press,New York, pp. 315–322.
206 T. Niskanen et al.