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The species of Cortinarius, section Bovini, associated with conifers in northern Europe
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The species of Cortinarius, section Bovini, associated with conifers in northern Europe Tuula Niskanen 1 Department of Biosciences, Plant Biology, P.O. Box 65, FI-00014 University of Helsinki, Finland Ilkka Kyto ¨ vuori Botanical Museum, P.O Box 7, FI-00014 University of Helsinki, Finland Kare Liimatainen Department of Biosciences, Plant Biology, P.O. Box 65, FI-00014 University of Helsinki, Finland Ha ˚kan Lindstro ¨m O ¨ stansjo ¨ 150, SE-840 64 Ka ¨larne, Sweden Abstract: Cortinarius bovinus and morphologically similar conifer-associated species were studied using material mainly from northern Europe. To stabilize the nomenclature, relevant types were examined. Phylogenetic relationships and species limits were investigated with rDNA ITS and nuclear rpb2 sequences as well as morphological data. We recognize seven species: C. bovinus (neotypified) and six new species, C. anisochrous, C. bovinaster, C.bovinatus, C. fuscobovinus, C. fuscobovinaster and C. oulankae ¨nsis. Their taxonomy, ecology, distribution and relationships are discussed, and a key to species is provided. Based on our phylogeny and morphological data the species were placed in section Bovini. Key words: ITS, MrBayes, rpb2, taxonomy INTRODUCTION Cortinarius is the most species-rich mycorrhizal genus in the world occurring in a number of forest ecosystems. Recent molecular analyses have shown that Cortinarius is monophyletic, however they also show that many of the traditional infrageneric groups such as Phlegmacium are not monophyletic (Peintner et al. 2004, Garnica et al. 2005). Nevertheless the majority of the species traditionally placed in subge- nus Telamonia do form a monophyletic group referred to here as Telamonia s. str. (Peintner et al. 2004, Garnica et al. 2005, Harrower et al. 2011). Telamonia is the most species rich and taxonomi- cally challenging group, and particularly the brown, medium to large species are not well known. Molecular methods have provided new tools for understanding the diversity and relationships of Cortinarius species, consequently an increasing num- ber of studies on Cortinarius have been published in recent years (e.g. Frøslev et al. 2007; Ortega et al. 2008; Garnica et al. 2009; Sua ´rez-Santiago et al. 2009; Niskanen et al. 2011a, 2012a). The studies by Kyto ¨vuori et al. (2005a) and Niskanen et al. (2008, 2009, 2011b, 2012b) in particular have examined the brown Telamonia species. This study focuses on Cortinarius bovinus Fr. and morphologically similar species occurring in boreal coniferous forests in rich forest soils. For several decades the name C. bovinus has been applied to rather large, fleshy brown Telamonia species by several authors (Moser 1978, Soop 2004), but no generally accepted survey of C. bovinus and related species exists. Section Bovini was described by Moser in 1975 (Moser and Horak 1975). The section included the type species C. bovinus and several South American species associated with Nothofagus and Eucalyptus. In a treatment of European species Moser (1983) included C. bovinus, C. fuscoperonatus Ku ¨ hner and C. rusticus P. Karst. (syn. C. canabarba M.M. Moser) in section Bovini. Later Moser excluded C. rusticus from this section and described a new North American species, C. pseudobovinus in the Bovini (Moser et al. 1995). The treatment of section Bovini by Moser (2001) also included C. sytnikii M.M. Moser from Europe as well as species from the southern hemi- sphere. Bidaud et al. (2009) included C. ectypus J. Favre and C. subtigrinus Reumaux in the section Bovini but listed the species under section Brunnei as well. Furthermore, they present interpretations of the less known C. injucundus (Weinm.) Fr., C. nitrosus Cooke, and C. pardinipes Romagn. In addition, the dark brown medium-sized and more or less stout Telamonia species also have been included in sect. Sordescentes and Brunnei (Brandrud et al. 1989, 1992, 1994, 1998; Melot 1990; Bidaud et al. 2002, 2009). These classification systems are presented here (SUPPLEMENTARY TABLE I). In this study we examined the taxonomy, ecology and distribution of C. bovinus and morphologically similar, conifer-associated species as well as their relationships based on molecular data. We also studied available, relevant type specimens. This work is part of our larger ongoing study of the north temperate and boreal Telamonia species. Submitted 8 Sep 2012; accepted for publication 15 Jan 2013. 1 Corresponding author. E-mail: [email protected] Mycologia, 105(4), 2013, pp. 977–993. DOI: 10.3852/12-320 # 2013 by The Mycological Society of America, Lawrence, KS 66044-8897 977
Transcript of The species of Cortinarius, section Bovini, associated with conifers in northern Europe
The species of Cortinarius, section Bovini,associated with conifers in northern Europe
Tuula Niskanen1
Department of Biosciences, Plant Biology, P.O. Box 65,FI-00014 University of Helsinki, Finland
Ilkka KytovuoriBotanical Museum, P.O Box 7, FI-00014 University ofHelsinki, Finland
Kare LiimatainenDepartment of Biosciences, Plant Biology, P.O. Box 65,FI-00014 University of Helsinki, Finland
Hakan LindstromOstansjo 150, SE-840 64 Kalarne, Sweden
Abstract: Cortinarius bovinus and morphologicallysimilar conifer-associated species were studied usingmaterial mainly from northern Europe. To stabilizethe nomenclature, relevant types were examined.Phylogenetic relationships and species limits wereinvestigated with rDNA ITS and nuclear rpb2 sequencesas well as morphological data. We recognize sevenspecies: C. bovinus (neotypified) and six new species, C.anisochrous, C. bovinaster, C.bovinatus, C. fuscobovinus,C. fuscobovinaster and C. oulankaensis. Their taxonomy,ecology, distribution and relationships are discussed,and a key to species is provided. Based on ourphylogeny and morphological data the species wereplaced in section Bovini.
Key words: ITS, MrBayes, rpb2, taxonomy
INTRODUCTION
Cortinarius is the most species-rich mycorrhizal genusin the world occurring in a number of forestecosystems. Recent molecular analyses have shownthat Cortinarius is monophyletic, however they alsoshow that many of the traditional infrageneric groupssuch as Phlegmacium are not monophyletic (Peintneret al. 2004, Garnica et al. 2005). Nevertheless themajority of the species traditionally placed in subge-nus Telamonia do form a monophyletic groupreferred to here as Telamonia s. str. (Peintner et al.2004, Garnica et al. 2005, Harrower et al. 2011).
Telamonia is the most species rich and taxonomi-cally challenging group, and particularly the brown,medium to large species are not well known.Molecular methods have provided new tools for
understanding the diversity and relationships ofCortinarius species, consequently an increasing num-ber of studies on Cortinarius have been published inrecent years (e.g. Frøslev et al. 2007; Ortega et al.2008; Garnica et al. 2009; Suarez-Santiago et al. 2009;Niskanen et al. 2011a, 2012a). The studies byKytovuori et al. (2005a) and Niskanen et al. (2008,2009, 2011b, 2012b) in particular have examined thebrown Telamonia species.
This study focuses on Cortinarius bovinus Fr. andmorphologically similar species occurring in borealconiferous forests in rich forest soils. For severaldecades the name C. bovinus has been applied torather large, fleshy brown Telamonia species byseveral authors (Moser 1978, Soop 2004), but nogenerally accepted survey of C. bovinus and relatedspecies exists.
Section Bovini was described by Moser in 1975(Moser and Horak 1975). The section included thetype species C. bovinus and several South Americanspecies associated with Nothofagus and Eucalyptus. Ina treatment of European species Moser (1983)included C. bovinus, C. fuscoperonatus Kuhner andC. rusticus P. Karst. (syn. C. canabarba M.M. Moser) insection Bovini. Later Moser excluded C. rusticus fromthis section and described a new North Americanspecies, C. pseudobovinus in the Bovini (Moser et al.1995). The treatment of section Bovini by Moser(2001) also included C. sytnikii M.M. Moser fromEurope as well as species from the southern hemi-sphere. Bidaud et al. (2009) included C. ectypus J.Favre and C. subtigrinus Reumaux in the sectionBovini but listed the species under section Brunnei aswell. Furthermore, they present interpretations of theless known C. injucundus (Weinm.) Fr., C. nitrosusCooke, and C. pardinipes Romagn. In addition, thedark brown medium-sized and more or less stoutTelamonia species also have been included in sect.Sordescentes and Brunnei (Brandrud et al. 1989, 1992,1994, 1998; Melot 1990; Bidaud et al. 2002, 2009).These classification systems are presented here(SUPPLEMENTARY TABLE I).
In this study we examined the taxonomy, ecologyand distribution of C. bovinus and morphologicallysimilar, conifer-associated species as well as theirrelationships based on molecular data. We alsostudied available, relevant type specimens. This workis part of our larger ongoing study of the northtemperate and boreal Telamonia species.
Submitted 8 Sep 2012; accepted for publication 15 Jan 2013.1 Corresponding author. E-mail: [email protected]
Mycologia, 105(4), 2013, pp. 977–993. DOI: 10.3852/12-320# 2013 by The Mycological Society of America, Lawrence, KS 66044-8897
977
MATERIALS AND METHODS
Material.—A total of about 200 collections were studied.Most collections were gathered by us from different parts ofFennoscandia and some from Estonia and central Europe(deposited in H and S). We also examined the herbariummaterial from O and the type specimens mentioned in thebeginning of Molecular analysis. Herbarium acronymsfollow Thiers http://sweetgum.nybg.org/ih/ (continuouslyupdated). For biogeographical provinces in Nordic coun-tries see Knudsen and Vesterholt (2008); for the othercountries provinces are used. Collectors are abbreviated bythe initials Tuula Niskanen (TN), Ilkka Kytovuori (IK), KareLiimatainen (KL) and Pirjo Kytovuori (PK).
Molecular analysis.—Several collections (n 5 3–21) of eachspecies from different geographical areas were sequenced(TABLE I). Also type material of Cortinarius subbrunneusBidaud, Carteret & Reumaux PC0098115 (PC) and C.fratellus Bidaud, Carteret & Reumaux PC0098114 (PC) werestudied. The sequencing of the type material of C. sytnikiiM.M. Moser was not successful. In addition, 19 speciesrepresenting different sections of Telamonia s. str. and/ormorphologically similar species of which no publishedmaterial existed were sequenced (TABLE I).
DNA was extracted from dried material (a piece oflamella) with the NucleoSpin plant kit (Macherey-Nagel,Duren, Germany). Primers ITS 1F and ITS 4 (White et al.1990, Gardes and Bruns 1993) were used to amplify ITSregions and specific primers cort6F and b7.1R (Frøslev et al.2005) for the rpb2 region. The same primer pairs were usedin direct sequencing. PCR amplification and sequencingfollowed Niskanen et al. (2009). Sequences were assembledand edited with Sequencher 4.1 (Gene Codes Corp., AnnArbor, Michigan). A total of 76 new ITS sequences and 46rbp2 sequences were produced (TABLE I) including 15sequences from type specimens. Rpb2 was chosen as asecond molecular marker because Frøslev et al. (2005)indicated that the use of rpb2 increases the resolution andnodal support for phylogenies in Cortinarius.
To determine the phylogenetic placement of the species,we included representatives of different sections of subge-nus Telamonia s. str. in our analysis. Using a BLAST query ofthe public databases (GenBank: http://www.ncbi.nlm.nih.gov/ and UNITE: http://unite.ut.ee/) we also selected allthe sequences similar (, 5 evolutionary events difference)to studied species for the analysis. In addition, morpholog-ically similar species to which we have compared the studiedspecies in this paper or have been classified in sect. Boviniby different authors (SUPPLEMENTARY TABLE I) were includedin the analysis. Two species from subgenus Phlegmaciumwere chosen as outgroup species, as in Niskanen et al.(2011a).
The combined ITS and rpb2 alignment of 142 sequenceswas produced with the program Muscle (Edgar 2004) underdefault settings. The part including ITS sequences wasmanually adjusted in BioEdit (www.mbio.ncsu.edu/BioEdit/bioedit.html). The alignment of rpb2 sequenceswas unambiguous, and no adjustments were made. Theoriginal alignment comprised 1356 nucleotides (includinggaps), and after the exclusion of areas with ambiguous
alignment (positions 443–467) 1331 positions were used foranalysis. Alignment is available at TreeBASE under S13118(http://www.treebase.org/treebase-web/home.html).
Bayesian inference (BI) was performed with MrBayes3.1.1 (Ronquist and Huelsenbeck 2003). The best substitu-tion model for the alignment was estimated by both theAkaike information criterion and the Bayesian informationcriterion with jModelTest 0.1.1 (Posada 2008). GTR modelincluding a gamma shape parameter and estimating theproportion of invariable sites was chosen for both align-ments. Two independent runs with four chains in each wereperformed for 4 000 000 generations with sampling every100th generation. All trees sampled before stationarity werediscarded with a 25% safety margin (burn-in of 10 000 trees[1 000 000 generations]). Sampled trees from both runswere combined in a 50% majority rule consensus phylogramwith posterior probabilities (PP). The analysis was run withcomputer clusters of the CSC, IT Centre for Science, Espoo,Finland.
Morphological studies.—Morphological descriptions arebased on material collected by the authors includingspecimens in all stages of development. Microscopiccharacteristics were observed from dried material mountedin Melzer’s reagent (MLZ). Measurements were made inMLZ with an ocular micrometer with 1003 oil-immersionlens. Basidiospores were measured from the veil or top ofthe stipe, 20 spores from one basidiocarp. The length andwidth were measured for each spore, and their length/width ratios (Q value) were calculated. The lamellar tramaand basidia also were examined, and the pileipellisstructure was studied from both radial freehand sectionsand scalps from the pileus center. The measurements of theelements of pileipellis were made from scalps.
RESULTS
Molecular analysis.—Based on our phylogeneticanalysis (FIG. 1) the studied species were placed insection Bovini (PP 0.91). Four of them, C. bovinus, C.bovinaster, C. bovinatus and C. oulankaensis, formed awell supported clade (PP 1.00) inside sect. Bovini.The section also includes C. aleuriodor, C. sp. (TF-01-034) and a clade comprising C. anisatus, C.neofurvolaesus and C. sordidemaculatus (PP 0.92).
All the species have smaller genetic intraspecificvariation (0–3 evolutionary events) as compared withthe interspecific variation (at least 20 evolutionaryevents) in the ITS regions, except C. fuscobovinus andC. fuscobovinaster. In the ITS region they differ by twobase changes. In the rpb2 region no differences existwithin each species and in general all the sisterspecies pairs differ by fewer than 10 evolutionaryevents. The rpb2 sequences of C. fuscobovinus and C.fuscobovinaster are identical. Intragenomic polymor-phisms were observed in 17–75% of the sampleswithin species. Polymorphisms were not observed intwo species, C. bovinatus and C. bovinaster, but these
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TABLE I. Specimens included in DNA analysis
Species Voucher Herb. Localityb
GenBanka
ITS RPB2
Sect. Bovini
C. subbrunneus Bidaud,Carteret & Reumaux(holotype) (syn. C.aleuriodor Rob. Henry)
PC0098115 PC France, Haute-Savoie, SaintGermain-sur-Rhone
JX407326 JX407369
C. aleuriodor CFP869 S France, Jura JX407327 JX407370C. anisatus H. Lindstr.,
Kytov., NiskanenTN04-550 H Finland, PeP, Runteli DQ120754 JX407346
C. anisochrous Kytov.,Liimat., Niskanen & H.Lindstr. (holotype)
IK01-030 H, S, NY Estonia, Hiiumaa, Suuremoisa JX407297 JX407347
C. anisochrous IK09-1617 H Finland, V, Parainen JX407298 JX407348C. anisochrous IK94-1221 H Finland, PS, Kerimaki JX407299 —C. anisochrous 27.9.1996 H Germany, Baden-
Wurttenberg, FreiburgJX407300 JX407349
C. anisochrous IK10-005 H Norway, Oppl, Gran JX407301 JX407350C. anisochrous IK10-007 H Norway, Oppl, Gran JX407302 JX407351C. anisochrous IK10-004 H Norway, Hedm, Hamar JX407303 JX407352C. anisochrous IK10-003 H Norway, Hedm, Hamar JX407304 JX407353C. anisochrous IK10-002 H Norway, Hedm, Hamar JX407305 JX407354C. anisochrous IK10-011 H Norway, Hedm, Hamar JX407307 —C. anisochrous CFP1459 S Sweden, Gtl, Gothem JX407306 JX407355C. anisochrous IK04-042 H Sweden, Vg, Medelplana JX407308 JX407356C. bovinaster Niskanen,
Kytov. & Liimat.(holotype)
TN04-669 H, S, NY Finland, PeP, Ylitornio JX407264 JX407340
C. bovinaster IK07-951 H Finland, Ks, Kuusamo JX407265 —C. bovinaster CFP1656 S Sweden, Jmt, Froso JX407266 —C. bovinatus Kytov., Liimat.,
Niskanen & H. Lindstr.IK09-1520b H Finland, ES, Kerimaki JX407267 JX407341
C. bovinatus (holotype) IK07-616,H6000849
H, S, NY Finland, PeP, Tornio JX407268 —
C. bovinatus CFP640 S Sweden, Vg, Medelplana JX407269 —C. bovinus Fr. IK01-047 H Estonia, Hiiumaa, Lauka JX407270 —C. bovinus IK01-048 H Estonia, Hiiumaa, Lauka JX407271 —C. bovinus TN00-1066 H Finland, V, Kisko JX407272 —C. bovinus IK97-2278 H Finland, V, Lohja JX407273 —C. bovinus IK01-033 H Finland, V, Lohja JX407274 —C. bovinus IK01-034 H Finland, V, Sarkisalo JX407275 —C. bovinus (neotype) IK04-038 H, S, NY Finland, U, Kirkkonummi JX407276 —C. bovinus IK04-037a H Finland, U, Tammisaari JX407277 —C. bovinus IK04-037b H Finland, U, Tammisaari JX407278 —C. bovinus IK97-1544 H Finland, Kn, Puolanka JX407279 —C. bovinus IK07-473 H Finland, PeP, Tervola JX407280 JX407342C. bovinus CFP1118 S Germany, Baden-Wurtenberg,
Schwarzwald-Baar KreisJX407281 —
C. bovinus TUB011898 TUB AY669691 —C. bovinus IK10-006 H Norway, Oppl, Lunner JX407282 JX407343C. bovinus IK10-001 H Norway, Hedm, Hamar JX407283 —C. bovinus JB6243-08 H Spain JX407284 —C. bovinus IK11-001 H Sweden, Gtl, Alskog JX407285 —C. bovinus IK11-003 H Sweden, Gtl, Viklau JX407286 —C. bovinus IK11-002 H Sweden, Gtl, Vaskinde JX407287 —C. bovinus IK04-036 H Sweden, Vg, Medelplana JX407288 —
NISKANEN ET AL.: CORTINARIUS SECTION BOVINI 979
TABLE I. Continued
Species Voucher Herb. Localityb
GenBanka
ITS RPB2
C. bovinus KS-CO884 S Sweden, Upl, Alvkarleby JX407289 —C. bovinus TN03-1001 H Sweden, Jmt, Froso JX407290 —C. fuscobovinaster Kytov.,
Liimat., Niskanen & H.Lindstr.
IK01-031 H Estonia, Hiiumaa, Puhalepa JX407309 —
C. fuscobovinaster CFP1100 S France, Ain, Brenod JX407310 JX407357C. fuscobovinaster EB411-97 O Norway, Oppl, Gran JX407311 —C. fuscobovinaster IK10-009 H Norway, Oppl, Lunner JX407312 —C. fuscobovinaster IK11-005 H Norway, Oppl, Lunner JX407313 —C. fuscobovinaster IK09-360 H Norway, NTr, Steinkjer JX407314 JX407358C. fuscobovinaster IK09-361 H Norway, NTr, Steinkjer JX407315 —C. fuscobovinaster (holotype) IK09-537 H, S, NY Norway, NTr, Steinkjer JX407316 JX407359C. fuscobovinaster IK11-004 H Sweden, Gtl, Fleringe JX407317 —C. fuscobovinaster IK07-1468 H Sweden, Upl, Alvkarleby JX407318 —C. fuscobovinus Kytov.,
Niskanen & Liimat.TN04-403 H Finland, PeP, Rovaniemi JX407319 JX407360
C. fuscobovinus IK98-1244 H Finland, PeP, Tornio JX407320 JX407361C. fuscobovinus (holotype) IK07-877,
H6001110H, S, NY Finland, Ks, Kuusamo JX407321 JX407362
C. fuscobovinus IK10-008 H Norway, Oppl, Gran JX407322 JX407363C. fuscobovinus TN03-598 H Sweden, Mpd, Havero JX407323 JX407364C. fuscobovinus IK97-606 H Sweden, Jmt, Froso JX407324 JX407365C. fuscobovinus CFP1181 S Sweden, Jmt, Ragunda JX407325 JX407366C. neofurvolaesus Kytov.,
Niskanen, Liimat., H.Lindstr.
IK04-001 H Finland, U, Helsinki DQ139997 JX407367
C. oulankaensis Kytov.,Niskanen, Liimat. & H.Lindstr.
SMIA48 BC Canada, BC FJ039672 —
C. oulankaensis IK01-032 H Estonia, Hiiumaa, Lauka JX407291 —C. oulankaensis (holotype) TN05-169 H, S, NY Finland, Ks, Kuusamo JX407292 —C. oulankaensis TN02-835 H Finland, Ks, Kuusamo JX407293 —C. oulankaensis IK10-010 H Norway, Oppl, Gran JX407294 JX407344C. oulankaensis IK09-535 H Norway, NTr, Steinkjer JX407295 JX407345C. oulankaensis TN03-1002 H Sweden, Jmt, Froso JX407296 —C. sordidemaculatus Rob.
HenryIK04-003 H Finland, U, Kirkkonummi DQ139991 JX407368
C. sp. IB86/172 IB Austria, Tirol DQ139983 —C. sp. TF-01-034 C Denmark, Farum Norreskov AJ889942 —
Other Telamonia species
C. pseudorubricosusReumaux (holotype)(syn. C. adustorimosus Rob.Henry)
GK16165 G France, Foret de Belval EU266709 —
C. albogaudis Kytov.,Niskanen, Liimat.
TN02-1089 H Finland, Ks, Kuusamo EU266633 JX407371
C. bivelus (Fr.) Fr. TUB011897 TUB AY669682 —C. brunneus (Pers. : Fr.)Fr. TN04-932 H Finland, U, Kirkkonummi EU266638 JX407372C. fratellus Bidaud, Carteret
& Reumaux (holotype)(syn. C. brunneus)
PC0098114 PC France, Mouilles, Sainte-Agnes
JX407328 —
C. colymbadinus Fr. CFP1130 S Sweden, Jmt, Ragunda JX127302 —C. disjungendus P. Karst. TN03-1701 H Slovakia, Liptovsky Mikulats JX407329 JX407373C. ectypus J. Favre (holotype) GK13318 G Switzerland, foret de Sauaidas EU266689 —
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two species were represented by the lowest number ofsequenced specimens.
TAXONOMY
Cortinarius sect. Bovini M.M. Moser, Beihefte zurNova Hedwigia 52:430 (1975) em. Liimat., Niska-nen & Kytov.Type species: Cortinarius bovinus Fr.Species medium to large. Basidiomes brown to dark
brown. Stipe often clavate to bulbose, rarely cylindri-cal. Context 6 brown, with age becoming dark brownat least at the base of the stipe. Universal veil white,brownish white or grayish white, in some speciesbecoming grayish brown with age. Mycelium whitish.Odor in most species indistinct or slightly raphanoid.Exsiccatae with dark brown to blackish brown pileusand pale grayish brown to grayish brown stipe. Spores6 amygdaloid to ellipsoid. In coniferous and decid-uous forests. Many of the species either calcicolous orcalciphilous.
Cortinarius disjungendus and some species of sect.Brunnei and Uracei resemble Bovini species, but they
all have blackish exsiccatae (both the stipe andpileus). In addition, the species in sect. Brunnei areoften entirely dark brown, have subglobose toellipsoid spores, and several species have stronglyencrusted lamellar trama hyphae. The species ofsection Uracei have metallic or greenish tints in freshbasidiomes, vinaceous or metallic tints in the blackexsiccatae, and spores that are fairly strongly verru-cose, amygdaloid, or are large (11–12 mm long) andobovoid-ellipsoid. Cortinarius suberi Soop and C.malachius (Fr.) Fr. have dark exsiccatae as well, butthe former has small spores (7.5–9 3 5–5.5 mm) and adistinctly innately fibrillose pileus. The latter has large(9.5–11 3 6–7 mm), weakly amygdaloid and finelyverrucose spores and in fresh condition it is so palethat it would not be confused with species in sect.Bovini.
Cortinarius bovinus Fr., Epicr. Syst. mycol.: 297(1838). FIGS. 2A, 3, 5A, 6APileus: 4–10 cm, hemispherical at first, then low
convex to almost plane, sometimes with a low, broadumbo, weakly fibrillose when young, later more
TABLE I. Continued
Species Voucher Herb. Localityb
GenBanka
ITS RPB2
C. fuscoperonatus Kuhner CFP1470 S France, Ain JX407330 JX407374C. impennoides Bidaud,
Moenne-Locc. & ReumauxTN10-084 H Canada, NF JQ746618 —
C. ionosmus M.M. Moser,Nespiak & Schwobel
TN04-590 H Finland, PeP, Tornio JX407331 JX407375
C. malachius (Fr.) Fr. IK98-1298 H Finland, PeP, Tervola JX407332 JX407376C. pseudobovinus M.M.
Moser & McKnight(holotype)
IB1989/0300 IB USA, Wyoming, Teton Natl. F. DQ499465 JX407377
C. raphanoides (Pers.) Fr. IK00-003 H Finland, St, Aetsa JX407333 JX407378C. rubrovioleipes Bendiksen
& K. BendiksenIK04-031 H Finland, PH, Hankasalmi DQ497191 JX407379
C. rusticus P. Karst.(syn. C. canabarba)
TN02-228 H Finland, Ks, Kuusamo JX407334 JX407380
C. subbalaustinus Rob.Henry
TN02-834 H Finland, Ks, Kuusamo JX407335 JX407381
C. suberi Soop TN04-155 H Finland, EH, Ruovesi JX407336 JX407382C. torvus (Fr.) Fr. IK98-1973 H Denmark, Nordjyllands amt,
Fjerritslev og BrovstJX407337 JX407383
C. traganus (Fr.) Fr. JV16638 TURA Finland, A, Jomala JX407338 JX407384C. uraceus Fr. TEB92-03 H Norway, Busk JX407339 JX407385
Outgroup
C. barbarorum Bidaud,Moenne-Locc. & Reumaux
TSJ2000-069 C Italy DQ083773 DQ083881
C. sodagnitus Rob. Henry TF2001-094 C Denmark DQ083812 DQ083920
a Sequences produced in this study in boldface.b For acronyms of biological provinces see for example Knudsen and Vesterholt, eds. 2008. Funga Nordica. p 32–35.
NISKANEN ET AL.: CORTINARIUS SECTION BOVINI 981
FIG. 1. The Bayesian 50% majority rule consensus tree inferred from ITS and rpb2 regions. PP . 0.50 are indicatedabove branches.
982 MYCOLOGIA
apparent only on the margin, somewhat waxy-glossywhen moist, with some veil remnants on the margin;reddish brown to dark reddish brown rarely withblackish spots; hygrophanous, soon drying from thecenter like Kuehneromyces mutabilis to lighter andmore reddish brown, in dry condition lighter ochra-ceous brown, up to 3 mm pellucid-striate when moist.Lamellae: medium-spaced to distant, strongly toweakly emarginate, fairly broad to broad, pale grayishbrown, later dark reddish brown or chocolate brown;edge somewhat lighter colored. Stipe: 4–10 3 0.9–1.5 cm at apex, 1.5–4.0 cm wide at base, clavate tobroadly bulbous at base, grayish white fibrillose, soonwith age and with handling darkening to grayishbrown. Universal veil: thin to moderately thick,grayish white to ochraceous white, darkening tograyish brown, forming a girdle and thin sheath orincomplete girdles on stipe, almost completely lostwith age. Mycelium: white. Context: grayish brown to
chocolate brown, later (blackish) brown in stipe base,marbled hygrophanous. Odor: indistinct or slightlyraphanoid. Exsiccatae: pileus sordid brown to darkgrayish brown; stipe brownish gray.
Spores: 8.6–10.4 3 5.7–6.6 mm, Q 5 1.44–1.59, x 5
9.3–9.7(–10.1) 3 6.1–6.4 mm, Qav. 5 1.46–1.53(–1.60)(300 spores, 15 specimens, FIGS. 2A, 3), weaklyamygdaloid to ellipsoid, with fairly rounded apex,dark brown, fairly strongly dextrinoid, moderatelyverrucose, fairly strongly to strongly so at the apex.Lamellar trama hyphae: pale olivaceous yellowish toolivaceous brownish, often with granulose to guttulatecontents, smooth (to finely scabrous), sometimes withsepia spots. Lamellar edge: more or less concolorous,sometimes somewhat darkened, fertile to fairly fertile,with basidia and almost filamentous to lageniform toclavate to pyriform marginal cells (FIG. 6A), 10–65 3
4–15 mm, not projecting (302 marginal cells, 14specimens). Narrow marginal cells often somewhatdark-walled and with intracellular olivaceous sub-stance at the apex. Basidia: four-spored, 31–45 3 8–10 mm (90 basidia, nine specimens), almost concolor-ous with the background to olivaceous brownish.Pileipellis: Epicutis hyphae 3–10 mm wide, paleolivaceous brownish, with fairly distinct wall, mostlysmooth, the lower ones somewhat wider than theupper ones, often finely, densely scabrous (parietalpigment). Hyphae in the transition to hypoderm paleolivaceous brownish, somewhat to distinctly encrusted(spotted to zebra-striped). Hypoderm present, notdistinctly differentiated, cells up to 25 3 30 mm wide.Clamp connections: present.
Ecology and distribution: In mesic to dryish conifer-ous forests with Picea, on rich, calcareous soil. One
FIG. 2. Spores of (A) Cortinarius bovinus IK 04-038 (H), (B) C. bovinatus IK 07-616 (H), (C) C. bovinaster KL & TN 04-669(H), (D) C. oulankaensis TN et al. 05-169, (E) C. anisochrous TN & IK 01-030, (F) C. fuscobovinus IK 07-877 and (G) C.fuscobovinaster IK 09-537, in Melzer’s reagent. Drawings by I. Kytovuori and T. Niskanen.
FIG. 3. Spore size of Cortinarius bovinus, C. bovinatus, C.bovinaster and C. oulankaensis. The lines are drawn on thebasis of scatter diagrams and contain 95% of the sporemeasurements of each species.
NISKANEN ET AL.: CORTINARIUS SECTION BOVINI 983
collection from Spain, however, is from Pinus forest.In young to old forests, also in disturbed vegetation,restricted to calcareous ground, in Finland oftenaround limestone quarries. Known from hemiborealto middle boreal zone in northern Europe butbecoming rarer northward (SUPPLEMENTARY FIG. 1).Also known from mountainous areas of central andsouthern Europe. Considered rare. Fruiting from lateAugust to early October.
Type: FINLAND. UUSIMAA: Kirkkonummi, Meiko-trasket, Picea dominated forest, small lime quarry,gregarious, 23 Sep 2004, Kytovuori 04-038 (H,neotype, here designated; S and NY, isoneotype.).GenBank JX407276.
Illustrations: Brandrud et al. (2012: pl. E20), Soop(2004: pl. 20).
Differential diagnosis: Cortinarius bovinus is charac-terized by a broadly clavate stipe, somewhat waxy-glossyand hygrophanous pileus (two concentric zones withreddish center, like Kuehneromyces mutabilis), universalveil becoming brownish gray, and darkening basi-diomes. The spores are weakly amygdaloid to ellipsoidand fairly strongly dextrinoid. The closely related C.bovinatus, C. oulankaensis and C. bovinaster have amore grayish brown pileus. In addition, C. bovinatushas amygdaloid and coarsely and sharply verrucosespores and C. oulankaensis and C. bovinaster have fairlydistant lamellae and relatively narrower spores. Corti-narius fuscobovinus and C. fuscobovinaster differ fromC. bovinus by having a slightly innately fibrillose pileus,pale brown to brown exsiccatae, and somewhat largerand less verrucose spores. Cortinarius anisochrous haspaler and grayer basidiomess when young, especially inthe context, smaller spores, and exsiccatae withbicolored pileus. Cortinarius impennoides Bidaud,Moenne-Locc. & Reumaux has somewhat paler, moregrayish or yellowish brown pileus, narrower spores,7.5–9 3 5–5.5(6) mm, and somewhat encrustedlamellar trama hyphae. Cortinarius subbalaustinus
Rob. Henry also can resemble C. bovinus, but theformer is usually more slender, has a vivid red-brownpileus, paler context, narrower spores, paler exsiccataeand is associated with Betula.
Cortinarius bovinus was described by Fries (1838) asa fleshy, robust species with a smooth and glossy,hygrophanous, cinnamon-brown pileus drying toyellowish brown. The lamellae were noted as thickand broad and the stipe having a grayish zone butwhite above. The fungus is rare in forests aroundUppsala, Sweden. In his later description Fries (1863)mentions that C. bovinus grows in coniferous forest.No published or unpublished illustrations of thisspecies by Fries exist. Our species fits well with Fries’sdecription: the fleshy shape, and the smooth andglossy, hygrophanous pileus with reddish brown tints.The other, closely related species usually have dullergrayish pileus. Our species also has been found inconiferous forest in the Uppsala area. Furthermore,C. bovinus ss. Soop (2004) and a sequence of C.bovinus (AY669691) published by Garnica et al.(2005) from Germany represent this species. Thephoto collection in Moser and Julich (1990) repre-sents another unknown species in this group, but thatspecies so far is known only from central Europe.Based on this we suggest that the name C. bovinusshould be used for the species presented here and wepropose a specimen-rich and representative collec-tion Kytovuori 04-038 as a neotype of the species. Inearlier literature the name C. bovinus has been usedfor several species, for example the material of C.bovinus in the ecological catalogues of Sweden andFinland (Hallingback and Aronsson 1998, Kytovuoriet al. 2005b) also includes other species of sectionBovini s. str.
Specimens studied: Specimens sequenced (TABLE I) andspecimens examined are included herein (SUPPLEMENTARY
TABLE II).
Cortinarius bovinatus Kytov., Liimat., Niskanen & H.Lindstr. sp. nov. FIGS. 2B, 3, 5B, 6G
MycoBank MB802593.Pileus: 3.5–8 cm, hemispherical at first, then low
convex to almost plane, with a low and broad umbo,with age often with concentric depression aroundumbo; weakly fibrillose when young, later moreapparent only on the margin; brown to grayishbrown, with blackish spots; hygrophanous, in dryconditions yellowish brown. Lamellae: medium-spaced, emarginate, fairly broad, at first pale grayishbrown, later dark reddish brown, edge somewhatlighter. Stipe: 6–14 long 3 0.8–1.5 cm thick at apex,2.0–4.0 cm at base, clavate, grayish white fibrillose,soon and when bruised darkening to grayish brown.Universal veil: thin to moderately thick, whitish,
FIG. 4. The spore size of C. anisochrous, C. fuscobovinusand C. fuscobovinaster. The lines are drawn on the basis ofscatter diagrams and contain 95% of the spore measure-ments of each species.
984 MYCOLOGIA
FIG. 5. A. Cortinarius bovinus IK 04-038 (H). B. C. bovinatus IK 07-616 (H). C. C. bovinaster KL & TN 04-669 (H). D. C.oulankaensis TN et al. 05-169 (H). E. C. anisochrous TN & IK 01-030. F. C. fuscobovinus IK 07-877 (H). G. C. fuscobovinaster IK11-004 (H). A, B, E, F, G by I. Kytovuori; C, D by K. Liimatainen.
NISKANEN ET AL.: CORTINARIUS SECTION BOVINI 985
forming a girdle and thin sheath or incompletegirdles on stipe, almost absent with age. Context:brown to grayish brown, later (blackish) brown fromstipe base upward, marbled hygrophanous. Odor: notrecorded. Exsiccatae: pileus dark brown, stipe palegrayish brown.
Spores: 8.4–10.4 3 5.7–6.6 mm, Q 5 1.44–1.67, av.5 8.7–10.0 3 5.9–6.3 mm, Qav. 5 1.48–1.59 (150spores, seven basidiocarps, three specimens, FIGS. 2B,3), amygdaloid (to somewhat fusoid), thick-walled,strongly, fairly coarsely and sharply verrucose, onlysomewhat more strongly so at the apex, stronglydextrinoid. Lamellar trama hyphae: pale olivaceousgreenish to olivaceous brownish, smooth to finelyscabrous. Lamellar edge: 6 concolorous with thelamellar trama, fertile, with basidia and clavate-filamentose (or narrowly sphaeropedunculate) mar-ginal cells (FIG. 6G), 15–35 3 4–9.5 mm (51 marginalcells, three specimens), narrow cells few, not project-ing. Basidia: four-spored, 33–45 3 8–9.5 mm (57basidia, three specimens), concolorous with thebackground to olivaceous brownish. Pileipellis: Epi-cutis hyphae 2–10 mm wide, thin-walled, almosthyaline to pale olivaceous brownish, smooth to finely,densely scabrous. Hypoderm present, not distinct,cells up to 25 3 45 mm, pale to yellowish. Transition totrama fairly light-colored (sometimes dark brown),with some obscure, olivaceous (to brown) encrusta-tions. Clamp connections: present.
Ecology and distribution: In mesic to dryish conifer-ous forests with Picea or Pinus, on calcareous soil. Onecollection from dry, pine-covered esker with lime dust,one from a dryish spruce forest with some pines andone from a mesic spruce forest with few pines. Rare inhemiboreal and boreal zones in northern Europe andknown only from Sweden and Finland (SUPPLEMENTARY
FIG. 1). Fruiting from late August to late September.Type: FINLAND. PERA-POHJANMAA: Tornio, Kor-
kiamaa, Runteli, rich, partly swampy grass-herb spruceforest with Betula, Populus tremula, Juniperus andPinus, on calcareous soil, 22 Aug 2007, Kytovuori 07-616, H6000849 (H, holotype; S and NY, isotype).GenBank JX407268.
Etymology: The name refers to affinity to C. bovinus.
Differential diagnosis: Cortinarius bovinatus hasbrown to grayish brown pileus, dark brown exsiccatae,and amygdaloid, strongly verrucose spores. It resemblesC. sordidemaculatus, but C. sordidemaculatus hassmaller (8–9 3 5–5.5 mm), thin-walled, weakly tomoderately dextrinoid spores. Similar are also C.bovinus, C. oulankaensis and C. bovinaster, but C.bovinus has a reddish brown to dark reddish brownpileus and more ellipsoid, moderately verrucose spores.Cortinarius oulankaensis and C. bovinaster have distantlamellae and relatively narrower spores. In addition theshape is different: narrowly ellipsoid to weakly amyg-daloid in C. oulankaensis and narrowly ellipsoid tonarrowly obovoidly ellipsoid in C. bovinaster.
Specimens studied: TABLE I.
FIG. 6. Marginal cells of (A) Cortinarius bovinus IK04-037 (H), (B) C. bovinaster CFP 1654 (S), (C) C. oulankaensis IK07-879 (H), (D) C. fuscobovinus IK 07-877 (H), (E) C. anisochrous IK 15 Sep 2001 (H), (F) C. fuscobovinaster TN & IK 01-031(H) and (G) C. bovinatus IK 07-616 (H), in Melzer’s reagent. Drawings by I. Kytovuori.
986 MYCOLOGIA
Cortinarius bovinaster Niskanen, Kytov. & Liimat., sp.nov. FIGS. 2C, 3, 5C, 6B
MycoBank MB802594.Pileus: 1.5–8 cm, broadly conical to hemispherical
when young, then low convex to almost plane with alow umbo, often narrowly pellucid-striate, surfacesomewhat fibrillose, margin often whitish especiallywhen young, brown to grayish brown when young,later dark brown, sometimes with dark strings orspots, hygrophanous, in dry condition yellowishbrown. Lamellae: medium-spaced to distant, stronglyto weakly emarginate, moderately broad to broad,pale yellowish brown, later dark brown, edge con-colorous or whitish. Stipe: 3.5–9 long 3 0.4–1 cmthick at apex, 1.3–2.5 cm at base, almost cylindrical toclavate, grayish white fibrillose, later grayish brown.Universal veil: white to grayish white, often forming athin sheath over the lower half of stipe. Mycelium:white. Context: pale grayish brown, dark browntoward the base, with age darkening entirely, marbledhygrophanous. Odor: indistinct. Exsiccatae: pileusdark brown; stipe grayish brown.
Spores: 8.8–10.7 3 5.4–6.3 mm, Q 5 1.54–1.70,av. 5 9.2–9.9 3 5.7–6.0 mm, Qav. 5 1.61–1.65 (200spores, 10 specimens, FIGS. 2C, 3), narrowly ellipsoidto narrowly obovoid-ellipsoid (or less commonlyweakly narrow ovoid-ellipsoid), with rounded apexand tapering base, sometimes with a low suprahilardepression, dark, moderately to fairly strongly dex-trinoid, fairly finely (to moderately), densely, evenlyverrucose, hardly at all more strongly so at the apex.Lamellar trama hyphae: pale olivaceous yellowish toolivaceous brownish, smooth tofinely, densely sca-brous. Lamellar edge: more or less concolorous,mostly fertile, with basidia and fairly wide, clavate toalmost globose marginal cells (FIGS. 6B) 10–33 3 5–14 mm, narrow cells rare, not projecting (155 marginalcells, 10 specimens). Basidia: four-spored, 30–40(–45)3 7–10 mm (90 basidia, nine specimens), almostconcolorous with the background to olivaceousbrown, mostly wide at the base. Pileipellis: Epicutishyphae 3–12 mm wide, pigmented, thin-walled, theupper ones mostly smooth, with sepia large spots, thelower ones somewhat wider, often distinctly zebra-striped (parietal pigment). Hypoderm fairly distinct,with roundish cells up to 30 3 45 mm wide. Hyphae inthe transition to trama (and trama) olivaceous tosepia spots more frequent than in the other species inthis group. Clamp connections: present.
Ecology and distribution: In mesic to fairly dampconiferous forests with Picea, on calcareous soil. Rarein the middle and northern boreal zones in northernEurope. So far known only from Finland and Sweden(SUPPLEMENTARY FIG. 1). Basidiocarps occur from mid-August to early September.
Type: FINLAND. PERA-POHJANMAA: Ylitornio,Palorommas, Kuusikkorompaat, Picea abies-dominat-ed forest on calcareous ground, 2 Sep 2004,Liimatainen & Niskanen 04-669 (H, holotype; Sand NY, isotype). GenBank JX407264, JX407340.
Etymology: The name indicates the resemblance to C.bovinus.
Differential diagnosis: Cortinarius bovinaster is rec-ognized by the often small basidiomes, the brown tograyish brown pileus, distant lamellae, and thenarrowly ellipsoid to narrowly obovoid-ellipsoid,finely, densely and evenly verrucose spores. Thespecies seems to have a northern distribution. Theother six species included here are usually larger. Thesister species, C. oulankaensis, is usually stouter andthe spores are larger and coarsely verrucose. Cortinar-ius bovinatus and C. sordidemaculatus differ byamygdaloid spores and denser lamellae. In addition,the spores of C. bovinatus are relatively broader andthick-walled and those of C. sordidemaculatus distinct-ly smaller. Cortinarius bovinus is distinguished bymore reddish brown, waxy-glossy pileus, and thespores are relatively broader.
Specimens studied: We have provided specimens sequenced(TABLE I) and specimens examined (SUPPLEMENTARY TABLE
III).
Cortinarius oulankaensis Kytov., Niskanen, Liimat. &H. Lindstr., sp. nov. FIGS. 2D, 3, 5D, 6C
MycoBank MB802887.Cortinarius oulankaensis Kytov., Niskanen, Liimat.
& H. Lindstr., Funga Nordica: 745 (2008) (invalid).Pileus: 4.5–10 cm, hemispherical, then low convex
to almost plane with a low and broad umbo, surfaceoften grayish white fibrillose, brown with some grayishtints when young, later dark brown, sometimes withblackish spots, margin whitish fibrillose; hygropha-nous, in dry condition yellowish brown. Lamellae:medium-spaced to distant, weakly emarginate, mod-erately broad to broad, pale grayish brown, later darkbrown, edge concolorous or whitish. Stipe: 5.5–10long 3 0.9–1.3 cm thick at apex, 1.5–2.5 cm at base,relatively stout and straight, almost cylindrical toclavate, grayish white fibrillose, later brownish. Uni-versal veil: grayish white, often forming a thin sheathover the lower half of stipe, sometimes forming a ringon the upper part of stipe. Mycelium: white. Context:pale grayish brown, chocolate brown toward the base,with age darkening entirely, marbled hygrophanous.Odor: indistinct or slightly raphanoid. Exsiccatae:pileus brown to dark brown pileus; stipe pale grayishbrown.
Spores: 9.3–11.1 3 6.1–6.8 mm, Q 5 1.51–1.70, av.5 9.9–10.6 3 6.3–6.6 mm, Qav. 5 1.56–1.62 (280spores, 14 specimens, FIGS. 2D, 3), narrowly ellipsoid
NISKANEN ET AL.: CORTINARIUS SECTION BOVINI 987
to weakly amygdaloid, dark, slightly (to moderately)dextrinoid, strongly and fairly coarsely verrucose,strongest at the apex. The large, dark verruculae onthe pale background of the spores give them avariegated appearance emphasizing the coarsenessof the ornamentation. Lamellar trama hyphae: paleolivaceous (greenish), mostly smooth, sometimes withgranulose contents. Lamellar edge: concolorous toweak sepia, fairly fertile, with basidia and clavate topyriform to sphaeropedunculate marginal cells(FIG. 6C), 11–45 3 7–15 um, narrow cells few, notprojecting (194 marginal cells, 11 specimens). Bluishgray to turquoise granulose contents present in someto many hyphae at lamellar edge of dried basidio-carps, at least in young basidiomes dried in goodcondition. Basidia: four-spored, 34–45 3 8–10 mm (95basidia, nine specimens), concolorous with thebackground to olivaceous brownish to olivaceousgreenish. Pileipellis: Epicutis hyphae fairly narrow,5–11 mm wide, thin-walled, almost smooth to distinctlyscabrous (zebra-striped to crust-like), sepia, withabundant small to large encrusted spots. Hypodermpresent, fairly thin-walled, cells up to 40 3 55 mmwide. Hyphae in the transition to sepia trama, withspot-like encrustations. Pileipellis light to dark sepiawhile olivaceous brownish in the other species of thegroup. Clamp connections: present.
Ecology and distribution: In mesic to fairly dampconiferous forests with Picea, on calcareous soil. Rarein the hemiboreal and boreal zones in northernEurope, although it can be abundant locally, as inOulanka National Park in Finland and in Steinkjer,Skrattasen, in Norway. Most records are from north-ern part of the middle boreal zone and southern partof the northern boreal zone (SUPPLEMENTARY FIG. 1).An ITS sequence from an ectomycorrhizal root tip,collected in Vancouver, Canada, also was identified asthis species.
Type: FINLAND. KOILLISMAA: Kuusamo, Ou-lanka, Puukkorinne, Picea abies dominated forest,on damp, calcareous ground, 19 Sep 2005, Niskanenet al. 05-169 (H, holotype; S and NY, isotype).GenBank JX407292.
Etymology: The species was first found in Oulanka,Finland.
Differential diagnosis: Typical for C. oulankaensisare grayish brown, somewhat fibrillose pileus, distantlamellae, dark brown exsiccatae, and fairly large andnarrowly ellipsoid, coarsely verrucose spores. Thesister species, C. bovinaster, differs by usually havingsmaller and more slender basidiomes, and smallerand finely, evenly verrucose spores. Cortinariusbovinatus has amygdaloid, strongly dextrinoid andrelatively broader spores and C. bovinus has morereddish brown, waxy-glossy pileus, and the spores are
smaller, relatively broader, and more dextrinoid.Cortinarius anisochrous has paler basidiomess whenyoung, exsiccatae with bicolored pileus, and smallerspores. Cortinarius fuscobovinus has distinctly innatelyfibrillose pileus, brown exsiccatae, and broadlyamygdaloid, fairly finely verrucose spores.
Specimens studied: We included specimens sequenced(TABLE I) and specimens examined (SUPPLEMENTARY TABLE
IV).
Cortinarius anisochrous Kytov., Liimat., Niskanen &H. Lindstr., sp. nov. FIGS. 2E, 4, 5E, 6E
MycoBank MB802888.Cortinarius anisochrous Kytov., Liimat., Niskanen & H.
Lindstr., Funga Nordica: 746 (2008) (invalid).
Pileus: 4–10(–12) cm, hemispherical when young,then low convex to plane, with a low and broad umbo,somewhat waxy-glossy when moist, surface oftenwhitish fibrillose, margin whitish silky-fibrillose, fairlypale grayish brown when young then later brown todark reddish brown, sometimes with dark reddishbrown to blackish dots, hygrophanous, often firstdrying from the center to lighter and more yellowishbrown then drying radially, in dry condition lightochraceous brown to light grayish brown. Lamellae:medium-spaced, emarginate, fairly broad, pale grayishbrown to yellowish brown, later dark brown, edgesometimes whitish. Stipe: 5–12(–15) 3 0.8–1.5 cm atapex, 2.0–3.5 cm wide at base, clavate to bulbous,sometimes stout, whitish fibrillose, later grayish brownto brownish. Universal veil: white, forming a thin, silkysheath or incomplete girdles on stipe, typically lostwith age or by handling. Mycelium: whitish. Context:pale grayish brown, later brownish, marbled hygro-phanous. Odor: indistinct. Exsiccatae: pileus usuallywith dark center and pale brown to brown margin;stipe pale grayish to brownish. Basidiomes in poorcondition or poorly dried specimens sometimesdrying blackish in the pileus center.
Spores: 8.2–9.7 3 5.4–6.3 mm, Q 5 1.42–1.61, av. 5
8.6–9.3 3 5.7–6.1 mm, Qav. 5 1.48–1.56 (260 spores,13 specimens, FIGS. 2E, 4), fusiformly ellipsoid withsomewhat tapering blunt apex (as in C. parvannula-tus), dark-colored, moderately to fairly strongly dex-trinoid, fairly strongly, sharply verrucose, strongest atthe apex. Lamellar trama hyphae: yellowish to paleolivaceous yellowish, smooth, with granulose contents.Lamellar edge: concolorous, fertile, with basidia andmostly fairly small, filamentose to pyriform marginalcells (FIG. 6E), 11–40(–65) 3 3.5–11(–15) mm, narrowcells fairly frequent, not projecting (203 marginalcells, 11 specimens). Basidia: four-spored, 31–45 3
7.5–9.5 mm (100 basidia, 10 specimens), almost hyalineto very pale olivaceous brownish. Pileipellis: Epicutishyphae 4–20 mm wide, thin-walled, light (pale
988 MYCOLOGIA
olivaceous to pale brownish), pellucid, poorly visible,smooth to finely, densely scabrous (zebra-striped,parietal pigment). Hypoderm present but fairly poorlydifferentiated, cells up to 25 3 30 mm wide. Hyphae inthe transition to trama somewhat encrusted, with veryfew large pigment spots and dark walls. Clampconnections: present.
Ecology and distribution: In mesic to dryish conifer-ous forests with Picea abies or Pinus sylvestris, oncalcareous soil. Known from the hemiboreal andsouthern boreal zones in northern Europe andconsidered rare (SUPPLEMENTARY FIG. 2). Also knownfrom mountaneous areas of central Europe. Basidio-carps occur from mid-September to mid-October.
Type: ESTONIA. HIIUMAA: Suuremoisa, Kallastepank, mesic Picea abies forest on calcareous soil, 14Sep 2001, Niskanen & Kytovuori 01-030 (H, holotype;S and NY, isotype). GenBank no. JX407297,JX407347.
Etymology: The name refers to the color of the exsiccatae,in which the pileus usually has a dark center and the marginis pale brown to brown.
Differential diagnosis: Cortinarius anisochrous hasthe palest basidiomes and context of this group whenyoung. It is a fairly stout species and typical has fairlysmall spores and exsiccatae with bicolored pileus. Ithas a southern distribution. Cortinarius bovinus, C.bovinatus, C. fuscobovinaster, C. oulankaensis and C.sordidemaculatus can resemble C. anischrous, buttheir context is usually darker, at least in the base ofthe stipe, and the pileus of exsiccatae is areconcolorous. In addition, C. bovinus has morereddish brown pileus and somewhat larger spores,C. bovinatus amygdaloid, thick-walled spores, C.fuscobovinaster slightly innately fibrillose pileus andlarger spores, C. sordidemaculatus distinctly smaller,amygdaloid and less dextrinoid spores, and C.oulankaensis distant lamellae and larger spores.
Specimens studied: Specimens sequenced (TABLE I) andspecimens examined are provided (SUPPLEMENTARY TABLE V).
Cortinarius fuscobovinus Kytov., Niskanen & Liimat.,sp. nov. FIGS. 2F, 4, 5F, 6D
MycoBank MB802889.Cortinarius fuscobovinus Kytov, Niskanen & Liimat.,
Funga Nordica: 744 (2008) (invalid).Pileus: (3.5)5–10 cm, when young hemispherical
with incurved margin, then low convex to plane,sometimes with an umbo, innately fibrillose, some-times with veil remnants on the margin, grayish yellowbrown to fairly dark brown, sometimes blackening inlarge spots; hygrophanous, with hygrophanous streaksand patches, in dry condition rather light brownishyellow to yellowish brown, somewhat darker towardthe margin. Lamellae: medium-spaced, emarginate,
fairly broad, pale brown, soon chocolate brown, edgeconcolorous or whitish. Stipe: 4–11 long 3 0.7–1.5 cmthick at apex, 2.0–3.5 cm at base, almost cylindrical toclavate at base, grayish white fibrillose, soon brownish.Universal veil: fairly abundant, grayish white, oftenforming a girdle and sheath or incomplete girdles onstipe. Mycelium: whitish. Context: brownish, darkertoward the base, at base blackish brown when old.Odor: weakly raphanoid. Exsiccatae: pileus unifor-mely pale brown to brown; stipe pale grayish orbrownish. Basidiomes in poor condition or poorlydried can be blackish in the pileus center.
Spores: 9.1–11.1 3 6.1–7.0 mm, Q 5 1.44–1.65,av. 5 9.4–10.8 3 6.3–6.7 mm, Qav. 5 1.48–1.59 (280spores, 14 specimens, FIGS. 2F, 4), broadly amygda-loid to ovoid, with low suprahilar depression, moder-ately dextrinoid, fairly finely, densely and evenlyverrucose, hardly at all more strongly so at the apex.Lamellar trama hyphae: pale olivaceous yellowish,smooth, sometimes with granulose or guttulatecontents. Lamellar edge: concolorous (to a bitdarker), moderately fertile, with basidia and mostlywide sphaeropedunculate, globose, ventricose toclavate marginal cells (FIG. 6D), 11–45 3 5–22 mm,narrow cells few, not projecting (244 marginal cells,12 specimens). Basidia: four-spored, 32–48 3 8–10 mm(95 basidia, nine specimens), concolorous with thebackground to olivaceous brownish. Pileipellis: Epi-cutis a thin layer, hyphae 5–15(–20) mm wide, thin-walled, light (pale brownish), mostly smooth (tofinely, densely scabrous). Hypoderm present, distinct,cells up to 40 3 50 mm wide, almost hyaline. Hyphaein the transition to trama fairly light, hardly encrust-ed, often with fairly large lactifers. Clamp connec-tions: present.
Ecology and distribution: In mesic coniferous forestswith Picea sometimes with Pinus, on calcareous soil. Itseems to favor well developed natural forests, most ofthe known occurrences are in nature reserves. Rare inboreal zone in northern Europe. The distribution isstrongly concentrated in the northern part of themiddle boreal zone and the southern part of thenorthern boreal zone (SUPPLEMENTARY FIG. 2). Fruit-ing from mid-August to mid-September.
Type: FINLAND. KOILLISMAA: Kuusamo com-mune, Oulanka National Park, between the Campingplace and E part of Puukkorinne, rich grass-herbPicea abies forest with some Pinus sylvestris, Betula andPopulus tremula, on calcareous ground, 27 Aug 2007,Kytovuori 07-877, H6001110 (H, holotype; S and NY,isotype). GenBank JX407321.
Etymology: The name refers to the resemblance to C.bovinus and the uniformly brown exsiccatae, which is oftenbrown unlike the almost blackish exsiccatae of C. bovinus.
NISKANEN ET AL.: CORTINARIUS SECTION BOVINI 989
Differential diagnosis: Typical for Cortinarius fusco-bovinus is innately fibrillose pileus, with persistentlyincurved margin, exsiccatae with uniformly palebrown to brown pileus, and fairly large, broadlyamygdaloid to ovoid spores. The sister species C.fuscobovinaster often has darker exsiccatae, narrowerspores and a more southern distribution. Cortinariusdisjungendus sometimes can be difficult to distuin-guish from C. fuscobovinus in the field, but C.disjungendus has almost blackish exsiccatae and thespores are obovoid. The related C. anisochrous hassmaller spores, the pileus is less innately fibrillose andin exsiccatae bicolored. Cortinarius bovinus, C.bovinatus and C. oulankaensis have darker exsiccataeand the pileus is less fibrillose.
Specimens studied: We included specimens sequenced(TABLE I) and specimens examined (SUPPLEMENTARY TABLE
VI).
Cortinarius fuscobovinaster Kytov., Liimat., Niskanen& H. Lindstr., sp. nov. FIGS. 2G, 4, 5G, 6F
MycoBank MB802890.Pileus: 2.5–7 cm, when young hemispherical, with
an incurved margin, later low convex to plane,sometimes with a low umbo; slightly innately fibril-lose, with thin veil patches on the margin, grayishbrown; hygrophanous and fading in streaks, dryingsomewhat yellowish brown. Lamellae: mediumspaced, emarginate, rather thin, at first pale brown,soon dark reddish brown, somewhat darkening withhandling, edges light. Stipe: 3–13 long 3 0.7–1.3 cmthick at apex, 1.5–3 cm at base, almost cylindrical toclavate, grayish white fibrillose, soon brownish.Universal veil: at first whitish, later distinctly brown,forming a wooly zone on stipe. Mycelium: whitish.Context: pale grayish brown, darker toward the baseof stipe, at base blackish brown when old, marbledhygrophanous. Odor: indistinct. Exsiccatae: pileusbrown to dark brown, stipe pale grayish or brownish.Basidiomes in poor condition or poorly dried can beblackish in the pileus center.
Spores: 8.8–10.7 3 5.4–6.3 mm, Q 5 1.52–1.73, av.5 9.4–10.1 3 5.8–6.2 mm, Qav. 5 1.57–1.64 (220spores, 11 basidiomes, nine specimens, FIGS. 2G, 4),narrowly ellipsoid to narrowly lacrymoid, with 6
rounded apex and a distinct suprahilar depression(sometimes the apex weakly tapering), fairly thin- tothick-walled, fairly finely to moderately, denselyverrucose, most strongly so at the apex (in shapeand size like those of C. bovinaster), moderately tostrongly dextrinoid. Lamellar trama hyphae paleolivaceous yellowish to olivaceous brownish, 6
smooth, sometimes with some granulose to guttulatecontents. Lamellar edge: concolorous, fairly fertile tosomewhat sterile, with basidia and sphaeropeduncu-
late to clavate (to filamentose) marginal cells(FIG. 6F), 16–36 3 4–15 mm, narrow cells notprojecting (82 marginal cells, two specimens). Basid-ia: four-spored, 33–45 3 7.5–9 mm (90 basidia, ninespecimens), almost concolorous with the backgroundto olivacous brownish. Pileipellis: Epicutis a thin layer,hyphae 6–15 mm wide, thin-walled, light (palebrownish), smooth to finely, densely scabrous (ze-bra-striped). Hypoderm present, distinct, cells up to35 3 50 mm, almost hyaline. Transition to trama light,not encrusted, with few small olivaceous pigmentglobules. Clamp connections present.
Ecology and distribution: In mesic to dryish conifer-ous forests with Picea abies, Pinus sylvestris or Abiesalba, on calcareous soil. Of the seven records two arefrom mesic spruce-dominated, grass-herb forests,three from submesic to dryish spruce forests withabundant pines and two from dryish karst ridges withspruce and pine. In the hemiboreal and southernboreal zones. Known from central and northernEurope (SUPPLEMENTARY FIG. 2).
Type: NORWAY. NORD-TRØNDELAG: Steinkjer,Skrattasen, mesic to dryish coniferous forest (Picea,Pinus), on karst, 5 Sep 2009, Kytovuori 09-537 (H,holotype; S and NY, isotype). GenBank JX407316.
Etymology: The name refers to affinity to C. fuscobovinus.
Differential diagnosis: Cortinarius fuscobovinaster is amedium-sized, grayish brown species with narrowlyellipsoid to narrowly lacrymoid, fairly large spores, anda southern distribution. The sister species, C. fuscobo-vinus, is most easily distinguished from C. fuscobovi-naster by the spores: The spores of C. fuscobovinus arebroader, 6.1–7.0 mm wide, whereas those of C.fuscobovinaster are 5.4–6.3 mm wide (Fig. 4). Also theshape is different, the spores of C. fuscobovinus arebroadly amygdaloid to ovoid (Fig 2). In addition, C.fuscobovinus has larger basidiomes, paler exsiccatae,and more northern distribution; it is concentrated inthe boreal zone while C. fuscobovinaster is mostfrequent in the hemiboreal zone. The ITS sequencesof the two species do not differ much, only two sites inthe ITS region are different. Nonetheless C. fuscobo-vinaster formed a monophyletic group in our phylo-genetic analysis, however C. fuscobovinus did not,instead it stayed as a basal group to C. fuscobovinaster,so that two distinct groups were not formed. Based onthe correlation of morphological and distributionaldifferences however, we conclude that C. fuscobovinusand C. fuscobovinaster represent two different species.
Cortinarius bovinaster and C. anisochrous can beconfused with C. fuscobovinaster. However, C. bovina-ster has a less fibrillose pileus and darker exsiccataeand C. anisochrous a less fibrillose pileus, palercontext, and somewhat smaller and broader spores.
990 MYCOLOGIA
Specimens studied: We include specimens sequenced(TABLE I) and specimens examined (SUPPLEMENTARY TABLE
VII).
KEY TO STUDIED SPECIES
1. Spores subglobose to broadly ellipsoid, . 7.3 mmlong; pileus and stipe in exsiccatae black to grayishblack . . . . . . . . sect. Brunnei p.p. (C. brunneus,
C. glandicolor, C. clarobrunneus)1.9 Spores amygdaloid to ellipsoid or if subglobose
then , 7.3 mm long; pileus in exsiccatae palebrown to dark brown to blackish, stipe (pale)grayish brown to black . . . . . . . . . . . . . . . . . . . . 22. Dried basidiomes entirely black(ish) . . . . . . 32. 9 Pileus in exsiccatae brown to blackish brown,
stipe (pale) grayish brown . . . . . . . . . . . . . . 63. Spores , 5.4 mm wide . . . . . . . . . . . . . . . . . . . . . 43.9 Spores . 5.4 mm wide . . . . . . . . . . . . . . . . . . . . . 5
4. Spores subglobose . . . . . . . . . . . . . . . . . . . . . . . .sect. Brunnei p.p. (C. albogaudis, C. ectypus)
4.9 Spores ellipsoid. . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . sect. Brunnei p.p. (C. pseudorubricosus)
5. Spores . 10 mm long . . . C. disjungendus, C. crassifolius5.9 Spores , 10 mm long . . . . . . . . . . . . . . C. uraceus
6. Spores , 5.5 mm wide . . . . . C. anisatus, C.neofurvolaesus, C. sordidemaculatus
6.9 Spores . 5.5 mm wide . . . . . . . . . . . . . . . . . 77. Pileus in exsiccatae brown all over; universal veil
fairly abundant to abundant . . . . . . . . . . . . . . . . 87.9 Pileus in exsiccatae blackish brown or with blackish
brown center; universal veil thin to moderatelythick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98. Large, northern species; spores 6.1–7.0 mm
wide . . . . . . . . . . . . . . . . . . 6. C. fuscobovinus8.9 Medium-sized, southern species; spores 5.4–
6.3 mm wide . . . . . . . . . . 7. C. fuscobovinaster9. Pileus in exsiccatae with warm reddish brown margin
and blackish brown center . . . . . . . . 5. C. anisochrous9.9 Pileus in exsiccatae blackish brown to olivaceous
brown all over . . . . . . . . . . . . . . . . . . . . . . . . . . 1010. Fresh pileus dark reddish brown, drying
bicolored with pale ochraceous brown centerand dark reddish brown marginal zone, likeKuehneromyces mutabilis; spores with 6 round-ed strongly verrucose apex . . . . . 1. C. bovinus
10.9 Fresh pileus grayish brown, often first dryingfrom the center and soon radially in otherparts or fading in streaks or spots; spores moreevenly verrucose . . . . . . . . . . . . . . . . . . . . 11
11. Spores narrowly lacrymoid, finely to moderately,densely verrucose . . . . . . . . . . . . . 3. C. bovinaster
11.9 Spores amygdaloid to ellipsoid, strongly, 6 coarse-ly verrucose . . . . . . . . . . . . . . . . . . . . . . . . . . . 1212. Lamellar edge of pileus (exsiccatae) with
grayish bluish contents; spores 9.3–11.1 mmlong, at most moderately dextrinoid. . . . . . . . .
. . . . . . . . . . . . . . . . . . 4. C. oulankaensis
12.9 Lamellar edge of pileus (exsiccatae) notbluish; spores 8.4–10.4 mm long, stronglydextrinoid . . . . . . . . . . . . . . . . 2. C. bovinatus
DISCUSSION
Cortinarius bovinus and similar conifer-associatedspecies.—We studied the C. bovinus group andrecognized six additional conifer-associated speciesfrom northern Europe: C. anisochrous, C. bovinaster,C. bovinatus, C. fuscobovinus, C. fuscobovinaster andC. oulankaensis. The species are characterized bybrown to dark brown basidiomes and exsiccatae withdark brown to blackish brown pileus. The universalveil is white, brownish white or grayish white, in somespecies becoming grayish brown with age, and theodor is indistinct or slightly raphanoid. The sporesseemingly develop fairly slowly, so older basidiomesalways should be included in the exsiccatae. Thespores of young basidiocarps are often somewhatsmaller than those from older ones correlating withthe observation of Niskanen et al. (2011a) from sect.Armillati. The color of the exsiccatae is important foridentifying species. The basiodiomes, however,should be in fresh, good condition when driedbecause those dried in the field yield much lighterexsiccatae. The marginal cells of lamellae are some-what different among the species, but the intraspe-cific variation is fairly high making it difficult to usethem in identification.
Most of the species are rare and occur inhemiboreal boreal and oroboreal coniferous forestsand some species have mostly more northerndistribution. They are all calcicolous and could beused as indicators of valuable forest sites as can thePhlegmacium species (Vesterholt 1991, Hallingbackand Aronsson 1998).
Coverage of the studied species was poor in thepublic sequence databases, as it was for sectionsBrunnei and Armillati studied by Niskanen et al.(2009, 2011a), and only sequences of two species werefound. However, the sequence data can providevaluable information on the ecology and distributionof the species, especially in groups such as sect. Bovinithat include rare and morphologically difficultspecies. This is true particularly in the future whenlarger surveys from different parts on the world areaccomplished.
Species and section taxonomy.—Five of the sevenspecies we recognized fulfill the criteria of a morpho-genetic species (Niskanen et al. 2009). They havedistinctive morphological characters, and pairwiseintraspecific differences (0–3 evolutionary events) inthe ITS regions are smaller than interspecific ones (at
NISKANEN ET AL.: CORTINARIUS SECTION BOVINI 991
least 20 evolutionary events). The two remainingspecies, C. fuscobovinus and C. fuscobovinaster, havemorphological and distributional differences butdiffer only by two evolutionary events in the ITSregion. This is not a unique situation in Cortinariusand the fact that some morphological species haveidentical or almost identical ITS sequences has beenreported in several Cortinarius studies (Garnica 2003,Ammirati et al. 2007, Frøslev et al. 2007, Niskanenet al. 2011a). As stated by Niskanen et al. (2011a), it isimportant to realize that similar ITS sequences arenot necessarily in conflict with the idea of distincttaxa. We lack sufficient data to confirm that ITSregions can separate all Cortinarius species. Usingmore variable regions might resolve the problem.
Based on our phylogeny and morphological data,the studied species were placed in section Bovini. Thesection also includes C. anisatus, C. neofurvolaesus, C.sordidemaculatus, C. aleuriodor and C. sp. (TF-01-034).Our phylogeny also shows that historical groupings ofthe section Bovini are polyphyletic. For example,Cortinarius pseudobovinus belongs to sect. Boulder-enses (Niskanen et al. 2006) and C. ectypus to sect.Brunnei (Niskanen et al. 2009), others did not haveclose relatives in our dataset but were placed outsidesect. Bovini. The sequencing of the type material of C.sytnikii, a species placed in sect. Bovini by Moser(2001) was not successful. Cortinarius sytnikii hasbroadly ellipsoid to subglobose spores (6.5–8.5 3 4.9–5.9 mm). It does not resemble any of the speciescurrently placed in the section Bovini and most likelyhas its closest relatives somewhere else in subgenusTelamonia s. str.
ACKNOWLEDGMENTS
We thank the Cortinarius Flora Photographica team forcollecting and documenting material of the species treatedin this paper, Joe Ammirati for revision of English, HeinoVanska for revision of Latin names and reviewers forvaluable comments. We are grateful for the help of thecurators of these herbaria: G, H, IB, O, PC and S. We alsothank Josep Ballara and Karl Soop for lending us theirmaterial. This work was supported by the Ministry ofEnvironment, Finland (YM38/5512/2009), and the SwedishTaxonomy Initiative (dha 165/08 1.4).
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NISKANEN ET AL.: CORTINARIUS SECTION BOVINI 993
SUPPLEMENTARY TABLE I. Cortinarius species classified in the group Bovini by different authors. Bold names indicate species included in the
section Bovini in this study. * Indicates species from Southern Hemisphere.
Moser (1975) Moser (1978) Moser (1995) Moser (2001) Bidaud et al. (2009)
C. bovinus C. bovinus C. bovinus C. bovinus C. bovinus
C. arcabucensis* C. canabarba C. fuscoperonatus C. fuscoperonatus C. canabarba
C. brunneovelatus* C. fuscoperonatus C. pseudobovinus C. pseudobovinus C. ectypus
C. lasiospermus* C. sytnikii C. fuscoperonatus
C. limbatus* C. arcabucensis* C. injucundus Weinmann
C. rufobrunneus* C. brunneovelatus* C. nitrosus Cooke
C. siccus* C. lasiospermus* C. pardinipes Romagn.
C. trachysporus* C. limbatus* C. subbovinus ad. int.
C. veronabrunneus* C. rufobrunneus* C. subtigrinus var. ectypoides Bidaud
C. vinaceocinereus* C. siccus* C. subtigrinus var. Subtigrinus Remaux
C. trachysporus* C. sytnikii Moser
C. vinaceocinereus*
SUPPLEMENTARY TABLE II. Cortinarius bovinus, specimens examined.
Locality Voucher Date Herb.
ESTONIA: Hiiumaa:
Lauka, Kõpu Niskanen & Kytövuori 01-047 15 Sep 2001 H
Lauka, Ristna Niskanen & Kytövuori 01-048 15 Sep 2001 H
Pühalepa, Kallaste Pank Niskanen & Kytövuori 14 Sep 2001 H
FINLAND: Varsinais-Suomi:
Kisko, Viiramäki Niskanen 00-1066 19 Sep 2000 H
loc. cit. Niskanen 00-1063 19 Sep 2000 H
loc. cit. Niskanen 00-1100 3 Oct 2000 H
loc. cit. Niskanen 01-451 23 Sep 2001 H
Lohja, Hermala Kytövuori 98-1648 6 Sep 1998 H
loc. cit. Kytövuori 97-2278 9 Oct 1997 H
Lohja, Virkkala – Vappula Kytövuori 27 Aug 2000 H
loc. cit. Kytövuori 10 Oct 2001 H (4 ex.)
Parainen, Ersby Kytövuori 09-1611–09-1615 30 Sep 2009 H
loc. cit. 09-1632 2009 H
loc. cit. 09-1634 2009 H
Suomusjärvi – Kisko,
Lemulanrinne Kytövuori 98-2197 21 Sep 1998 H
Särkisalo, Förby Kytövuori et al. 01-034 25 Sep 2001 H
loc. cit. P. & I. Kytövuori 09-1645 1 Oct 2009 H
Västanfjärd, Illo Kytövuori et al. 25 Sep 2001 H
Uusimaa:
Kirkkonummi, Meikoträsket Kytövuori et al. 04-038 23 Sep 2004 NEOTYPE, H
loc. cit. Kytövuori et al. 23 Sep 2004 2 ex. H
loc. cit. Kytövuori 07-1720 27 Sep 2007 H
loc. cit. Kytövuori 07-1744 29 Sep 2007 H
Tammisaari, Skärlandet Kytövuori et Toivonen 04-037 4 Oct 2004 H
Kainuu:
Paltamo, Melalahti Kytövuori 08-1453 10 Sep 2008 H
Puolanka, Väyrylä Kytövuori 97-1544 15 Sep 1997 H
Perä-Pohjanmaa:
Tervola, Raemäki Kytövuori 07-473 21 Aug 2007 H
GERMANY: Baden-
Würtenberg:
Schwarzwald-Baar Kr., Wolt.
Stadt CFP 1118 12 Oct 1991 H
NORWAY: Akershus:
Oslo, Nordmarka E. Bendiksen 368/84 27 Sep 1984 O
Buskerud:
Røyken, Bøsnipa Kytövuori 13 Sep 2011 H
Oppland:
Lunner, Galtedalstjerna N Kytövuori et al. 10-006 12 Sep 2010 H
Lunner, Skøyenåsen Kytövuori 3 Sep 2011 H
Hedmark:
Hamar Kytövuori et al. 10-001 7 Sep 2010 H
SWEDEN: Gotland:
Alskog, Olljavs Kytövuori 27 Sep 2011 H
Gammelgarn, Uppstaig Kytövuori 27 Sep 2011 H
Viklau, Tjaukle Kytövuori 30 Sep 2011 H
Väskinde Brandrud 26 Sep 2011 H
loc. cit. Kytövuori 29 Sep 2011 H (2 ex.)
loc. cit. P. & I. Kytövuori 29 Sep 2011 H
loc. cit. Lundmark 29 Sep 2011 H
Västergötland:
Medelplana, Hjälmsäter Kytövuori et al. 27 Sep 2004 H
Sjöskogen Kytövuori et al. 04-036 28 Sep 2004 H
loc. cit. Kytövuori 2004 H
loc. cit. Niskanen et al. 04-1001 2004 H
Uppland:
Hållnäs, Hållen Toivonen & Kytövuori 07-2063 19 Sep 2007 H
Älvkarleby, Gårdskär Soop CO884 30 Sep 1997
Medelpad:
Borgsjö, Ensillre nature reserve Kytövuori 24 Aug 2010 H
Jämtland:
Frösö, Fillstabäcken Niskanen 03-1001 et al. 2 Sep 2003 H
loc. cit. Kytövuori 27 Sep 2010 H
Revsund, Ammerön Kytövuori 26 Aug 2010 H
Sunne, Andersön CFP 995 10 Sep 1990 S
SUPPLEMENTARY TABLE III. Cortinarius bovinaster, specimens examined.
Locality Voucher Date Herb.
FINLAND: Perä-Pohjanmaa:
Tervola, Peura, Kaitaharju Kytövuori 07-512 21 Aug 2007 H
Tornio, Runteli Kytövuori 98-1258a 20 Aug 1998 H
Ylitornio, Palorompaat Niskanen 04-669 2 Sep 2004 HOLOTYPE, H
Koillismaa:
Kuusamo, Oulanka Kytövuori 07-853 27 Aug 2007 H
loc. cit. Kytövuori 07-1090 1 Sep 2007 H
Kuusamo, Oulanka, Liikasenvaara
Liimatainen & Niskanen 02-
314 30 Aug 2001 H
loc. cit. Toivonen & Kytövuori 07-951 29 Aug 2007 H
SWEDEN: Jämtland:
Frösö, Fillstabäcken Kytövuori 97-601 2 Sep 1997 H
loc. cit. CFP 1653 28 Aug 2005 S
loc. cit. CFP 1654 28 Aug 2005 S
loc. cit. CFP 1655 28 Aug 2005 S
loc. cit. CFP1656 28 Aug 2005 S
loc. cit. Kytövuori 27 Sep 2010 H
SUPPLEMENTARY TABLE IV. Cortinarius oulankaënsis, specimens examined.
Locality Voucher Date Herb.
ESTONIA: Hiiumaa:
Lauka, Kõpu Kytövuori & Niskanen 01-032 15 Sep 2001 H
FINLAND: Koillismaa:
Kuusamo, Liikasenvaara Kytövuori & Toivonen 07-953 29 Aug 2007 H
Kuusamo, Oulanka, Ampumavaara Niskanen 02-835 19 Sep 2002 H
loc. cit. Niskanen 02-846 19 Sep 2002 H
loc. cit. Niskanen 02-857 19 Sep 2002 H
loc. cit. Niskanen 02-863 19 Sep 2002 H
loc. cit. Niskanen D05-139 17 Sep 2005 H
loc. cit. Niskanen 05-140 17 Sep 2005 H
loc. cit. Niskanen 05-153 17 Sep 2005 H
loc. cit. Kytövuori 07-856 27 Aug 2007 H
loc. cit. Kytövuori 17 Sep 2005 H (4 ex. )
loc. cit. Kytövuori 07-1114 2 Sep 2007 H
loc. cit. Kytövuori 07-1115 2 Sep 2007 H
Kuusamo, Oulanka, Kiutaköngäs Niskanen 02-883 18 Sep 2002 H
Kuusamo, Oulanka, Puukkorinne Kytövuori 07-1084 1 Sep 2007 H
loc. cit. Kytövuori 07-1092 1 Sep 2007 H
loc. cit. Kytövuori 19 Sep 2005 H (2 ex. )
loc. cit. Niskanen 02-995 22 Sep 2002 H
loc. cit. Niskanen D02-1006 22 Sep 2002 H
loc. cit. Niskanen 02-1079 23 Sep 2002 H
loc. cit. Niskanen D05-169 19 Sep 2005 HOLOTYPE, H
loc. cit. Niskanen 05-177 19 Sep 2005 H
Oulanka, S foot of Puukkorinne Kytövuori 07-879 27 Aug 2007 H
loc. cit. Kytövuori 07-880 27 Aug 2007 H
loc. cit. Kytövuori 07-881 27 Aug 2007 H
NORWAY: Oppland:
Gran K. Bendiksen IK10-010 12 Sep 2010 H, O
Nordre Land, Dokka naturreservat Brandrud 137-00 18 Aug 2000 O
Nord-Trøndelag:
Snåsa, Bergåsen E. Bendiksen 2 Sep 2009 O
Steinkjer, Kvam, Noem Kytövuori 09-362 2 Sep 2009 H
Steinkjer, Sunnan, Skrattåsen Kytövuori 09-535 5 Sep 2009 H
loc. cit. Kytövuori 09-536 5 Sep 2009 H, O, S
SWEDEN: Medelpad:
Borgsjö, Ensillre nature reserve Kytövuori 24 Aug 2010 H
Jämtland:
Frösö, Fillstabäcken Niskanen et al. 03-1002 2 Sep 2003 H
loc. cit. Kytövuori 27 Sep 2010 H
Frösövallen CFP 1452 13 Aug 2000 S
Revsund, Ammerön Kytövuori 26 Aug 2010 H
SUPPLEMENTARY TABLE V. Cortinarius anisochrous, specimens examined.
Locality Voucher Date Herb.
ESTONIA: Hiiumaa:
Lauka, Kõpu Niskanen & Kytövuori 15 Sep 2001 H (3 ex. )
Pühalepa, Kallaste pank Niskanen & Kytövuori 01-030 17 Sep 2001 HOLOTYPE, H
loc. cit. Vauras 17 Sep 2001 TUR
loc. cit. Kytövuori 14 Sep 2001 H
FINLAND: Varsinais-Suomi:
Parainen, Ersby Salo & Salo 10042 22 Sep 2004 H
loc. cit. Kytövuori 09-1617 30 Sep 2009 H
loc. cit. Kytövuori 09-1618 30 Sep 2009 H
loc. cit. Kytövuori 09-1633 30 Sep 2009 H
Pohjois-Savo:
Kerimäki, Louhi Kytövuori 94-1221 1 Oct 1994 H
loc. cit. Kytövuori 09-1517 23 Sep 2009 H
loc. cit. Kytövuori 09-1518 23 Sep 2009 H
GERMANY: Baden-
Würtenberg:
Freiburg, Hornberg Kytövuori 23 Sep 1996 H
NORWAY: Oppland:
Gran, Jøvika E Kytövuori et al. 10-005 12 Sep 2010 H
loc. cit. Kytövuori 10-007 12 Sep 2010 H
loc. cit. Kytövuori 12 Sep 2010 H
Hedmark:
Hamar Kytövuori et al. 10-004 7 Sep 2010 H
loc. cit. Kytövuori et al. 10-003 7 Sep 2010 H
loc. cit. Kytövuori et al. 10-002 7 Sep 2010 H
SWEDEN: Gotland:
Eksta, Ekstakusten, Skansudd Wasstorp 28 Sep 2011 H
Eksta, Vavle von Bonsdorff 28 Sep 2011 H
Gothem, Gothem klint CFP 1459 20 Sep 2000 S
Väskinde, Brucebo Brandrud 26 Aug 2011 GB, H
loc. cit. Kytövuori 29 Sep 2011 H
loc. cit. P. & I. Kytövuori 29 Sep 2011 H
Västergötland:
Medelplana, Sjöskogen Kytövuori et al. 28 Sep 2004 H (2 ex. )
Medelplana, Eriksberg CFP 1590 Sep 2004 S
Medelplana, Såtatorp Kytövuori et al. 27 Sep 2004 H (2 ex. )
loc. cit. Niskanen 04-991 27 Sep 2004 H
Södermanland:
Mörkö, Oaxen P. & I. K. 98-2240 27 Sep 1998 H
Uppland:
Älvkarleby, Rullsand CFP 9 Oct 2006 S
SUPPLEMENTARY TABLE VI. Cortinarius fuscobovinus, specimens examined.
Locality Voucher Date Herb.
FINLAND: Pohjois-Savo:
Kerimäki, Louhi Kytövuori 98-1746 8 Sep 1998 H
Kainuu:
Puolanka, Väyrylä Toivonen & Kytövuori 10 Aug 2002 H
Perä-Pohjanmaa:
Keminmaa, Törmä Kytövuori 08-068 25 Aug 2008 H
Rovaniemi, Louevaara Niskanen 04-403 28 Aug 2004 H
Tervola, Kaitaharju Kytövuori 07-511 21 Aug 2007 H
Tervola, Kirvesmaa Kytövuori 07-709 24 Aug 2007 H
loc. cit. Kytövuori 07-754 2007 H
Tervola, Raemäki Kytövuori 98-1299 21 Aug 1998 H
loc. cit. Kytövuori 07-742 21 Aug 2007 H
Tornio, Kalkkimaa Kytövuori 07-560 22 Aug 2007 H
loc. cit. Kytövuori 07-599 22 Aug 2007 H
loc. cit. Niskanen 04-600 1 Sep 2004 H
loc. cit. Niskanen 04-549 30 Aug 2004 H
Tornio, Runteli Kytövuori 98-1243 20 Aug 1998 H
loc. cit. Kytövuori 98-1244 20 Aug 1998 H
loc. cit. Niskanen 04-539 30 Aug 2004 H
Koillismaa:
Kuusamo, Oulanka Kytövuori 07-828 26 Aug 2007 H
loc. cit. Kytövuori 07-829 26 Aug 2007 H
loc. cit. Kytövuori 07-830 26 Aug 2007 H
loc. cit. Kytövuori 07-877 27 Aug 2007 HOLOTYPE, H
loc. cit. Kytövuori 07-878 27 Aug 2007 H
loc. cit. Kytövuori 07-876 27 Aug 2007 H
loc. cit. Kytövuori 07-855 27 Aug 2007 H
loc. cit. Kytövuori 07-1091 1 Sep 2007 H
loc. cit. Kytövuori 07-1113 2 Sep 2007 H
Kuusamo, Oulanka, Liikasenvaara Toivonen & Kytövuori 07-1005 29 Aug 2007 H
loc. cit. Toivonen & Kytövuori 07-952 29 Aug 2007 H
NORWAY: Oppland:
Gran, Jøvika E Kytövuori et al. 10-008 12 Sep 2010 H
loc. cit. E. Bendiksen 12 Sep 2010 H, O
Hedmark:
Hamar Kytövuori et al. 7 Sep 2010 H
Nord-Trøndelag:
Lierne, Sandsjöen Brandrud 29 Aug 2010 H
SWEDEN: Medelpad:
Haverö, Boflon O Niskanen et al. 03-598 27 Aug 2003 H
Jämtland:
Frösö, Fillstabäcken Kytövuori 97-606 2 Sep 1997 H
loc. cit. Kytövuori 27 Sep 2010 H (3 ex. )
Nyhem, Storselet Kytövuori 25 Aug 2010 H
Ragunda, Hoo CFP 1181 9 Aug 1993 S
Revsund, Ammerön Kytövuori 26 Aug 2010 H (2 ex. )
Sundsö, Väster-Andsjön M. A. 23 Aug 2010 H
Sunne, Andersön CFP 995 10 Sep 1990 S
Åre, Nordhallen
Toivonen & Kytövuori 06-
1639 23 Aug 2006 H
SUPPLEMENTARY TABLE VII. Cortinarius fuscobovinaster, specimens examined.
Locality Voucher Date Herb.
ESTONIA: Hiiumaa:
Lauka, Kõpu Niskanen & Kytövuori 15 Sep 2001 H
Pühalepa, Kallasta pank Niskanen & Kytövuori 01-031 17 Sep 2001 H
FRANCE: Ain:
Brenod, Jalinard CFP 1100 7 Oct 1991 S
NORWAY: Oppland:
Gran, Askimlandet E. Bendiksen 411/97 3 Oct 1997 O
Lunner, Galtedalstjerna N Kytövuori et al. 10-009 12 Sep 2010 H
Lunner, Skøyenåsen Kytövuori 11-005 3 Sep 2011 H
Nord-Trøndelag:
Steinkjer, Kvam, Noem Kytövuori 09-360 2 Sep 2009 H
loc. cit. Kytövuori 09-361 2 Sep 2009 H
Steinkjer, Sunnan, Skrattåsen Kytövuori 09-537 5 Sep 2009 HOLOTYPE, H
loc. cit. Kytövuori 09-538 5 Sep 2009 H
SWEDEN: Gotland:
Fleringe, Bästeträsk Kytövuori 11-004 26 Sep 2011 H (3 ex. )
Väskinde, Brucebo NE Lindström & Bendiksen 30 Sep 2011 H, O
Uppland:
Älvkarleby, Billudden Toivonen & Kytövuori 07-1468 18 Sep 2007 H
