Entoloma subgenus Leptonia in boreal-temperate Eurasia: Towards a phylogenetic species concept

Abstract

This study reveals the concordance, or lack thereof, between morphological and phylogenetic species concepts within Entoloma subg. Leptonia in boreal-temperate Eurasia, combining a critical morphological examination with a multigene phylogeny based on nrITS, nrLSU and mtSSU sequences. A total of 16 taxa was investigated. Emended concepts of subg. Leptonia and sect. Leptonia as well as the new sect. Dichroi are presented. Two species (Entoloma percoelestinum and E. sublaevisporum) and one variety (E. tjallingiorum var. laricinum) are described as new to science. On the basis of the morphological and phylogenetical evidence E. alnetorum is reduced to a variety of E. tjallingiorum, and E. venustum is considered a variety of E. callichroum. Accordingly, the new combinations E. tjallingiorum var. alnetorum and E. callichroum var. venustum are proposed. Entoloma lepidissimum var. pauciangulatum is now treated as a synonym of E. chytrophilum. Neotypes for E. dichroum, E. euchroum and E. lampropus are designated

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Persoonia32,2014:141–169
www.ingentaconnect.com/content/nhn/pimj http://dx.doi.org/10.3767/003158514X681774
RESEARCH ARTICLE
INTRODUCTION
The genus Entolomas.l.isveryspecies-richandmorpholo-
gicallydiverse.Itcontainsmore than 1500 species and oc-
cursworldwidefrom arctic to tropical habitats (e.g.,Largent
1977,1994, Romagnesi & Gilles 1979,Horak 1980, 2008,
Noordeloos1988, 1992, 2004, 2008, Gates & Noordeloos
2007,Noordeloos&Hausknecht2007,Vila&Caballero,2007,
2009,Noordeloos&Gates2009,2012,Morozovaetal.2012).
Recentmolecularlybasedphylogeneticstudieshaverevealed
that the genus is monophyletic and sister to the Clitopilus/
Rhodocybeclade(Moncalvoetal.2002,Mathenyetal.2006,
Co-Davidetal.2009,Baroni&Matheny2011).Besidesaga-
ricoid basidiocarp types, the genus also comprises gasteroid,
pleurotoidandcyphelloidforms(Co-Davidetal.2009,Baroni
&Matheny2011).Life-styleisequallyvaried,fromsaprotrophs
to parasites, or mycorrhizal symbionts. The classification of
Entolomatoidagaricstraditionallyfollowestwolines.Thefirst
group of authors interprets it as a single species-rich entity with
anelaborateinfragenericclassification(e.g.,Romagnesi1974,
1978,Noordeloos1992,2004),whilethesecondsegregates
Entoloma s.l.in upto13genera(e.g.,Largent&Baroni1988,
Orton1991a,b,Largent1994,Baroni&Matheny2011).De-
limitationof(sub)genericentities,haslongbeenbasedsolely
onmorphologicalcharacters(e.g.inRomagnesi&Gilles1979,
Noordeloos1981,2004).Asaresultofthephylogeneticstud-
ies mentioned above, it also became apparent that many of
the(sub)genericdivisionsappeartobeparaphyletic.Recently
Baronietal.(2011)describedthenewgenusEntocybe within
the Entolomatoid clade to accommodate species with basidio-
spore morphology intermediate between Entoloma and Rhodo-
cybe,supportedalsobythemoleculardata.Ongoingstudies,
however,suggestthatalsothisentityispolyphyletic(Morgado
etal.2013).Moreover,Baronietal.(2011)havedemonstrated
the paraphyly of the Entolomataceae.Continuedphylogenetic
studies, based on both morphological characters and molecular
markers(Heetal.2013,Morgadoetal.2013,Vilaetal.2013)
reveal more insight into the interrelation between morphological
and phylogenetic species concepts, as well as into the evolu-
tion of the Entolomataceae, that will result in future in a more
naturalclassification.
Also subg. Leptonia in the sense of Noordeloos (2004) is
polyphyletic.Sect. Leptonia of the subgenus belongs to the
Nolanea-Claudopus clade, and Cyanula and Griseorubida to the
Inocephalus-Cyanulaclade(Co-Davidetal.2009).Basedon
these data Cyanula recently has been raised to the subgenus
level(Noordeloos&Gates2012).
This paper is an attempt to clarify the phylogeny and species
concept of a morphologically distinct group within the genus
Entoloma,viz.subg.Leptonia sect.Leptoniaintheclassification
ofNoordeloos(2004).Despitethefactthatmostspeciesare
rare,someofthemweredescribedasfarbackasinthe19th
century.The protologues of these species were short and
incomplete, and the type specimens were not preserved.
The morphological variability of Leptonia species appears to
be very high and depending also on the age of basidiomata
andweatherconditions.Duetothesefactors,alargenumber
ofmisunderstandingsand incorrect identifications are found
in the literature. This paper aims to describe morphological
variability of each phylogenetic species resulting in emended
descriptionsandanidentificationtool,aswellastoreconstruct
aninfragenericclassificationofsubg.Leptonia.
MATERIALS AND METHODS
Taxa sampling
Toclarifythetaxonomicstatusof16taxaofEntolomasubg.Lep-
toniaaswellastheirpositionwithinthegenus,98specimens
of this subgenus or previously considered as belonging to this
subgenus were selected for morphological study and molecular
Entoloma subgenus Leptonia in boreal-temperate Eurasia:
towards a phylogenetic species concept
O.V.Morozova1,M.E.Noordeloos2,J.Vila3
1 KomarovBotanical Institute,197376, 2Prof. Popov Str.,St Petersburg,
Russia;correspondingauthore-mail:ovm.leptonia@gmail.com.
2 Naturalis Biodiversity Center, section Botany,P.O.Box 9517, 2300 RA
Leiden,TheNetherlands.
3 P.O.Box30041,08034Barcelona,Spain;
 e-mail:vilamicol@telefonica.net.
Key words
Entolomataceae
morphology
multiple gene phylogeny
neotypes
new species
Abstract This study reveals the concordance, or lack thereof, between morphological and phylogenetic species
concepts within Entolomasubg.Leptonia in boreal-temperate Eurasia, combining a critical morphological examina-
tionwithamultigenephylogenybasedonnrITS,nrLSUandmtSSUsequences.Atotalof16taxawasinvestigated.
Emendedconceptsofsubg.Leptoniaandsect.Leptoniaaswellasthenewsect.Dichroiarepresented.Twospecies
(Entoloma percoelestinum and E. sublaevisporum)andonevariety(E. tjallingiorumvar.laricinum)aredescribedas
newtoscience.OnthebasisofthemorphologicalandphylogeneticalevidenceE. alnetorum is reduced to a variety
of E. tjallingiorum, and E. venustum is considered a variety of E. callichroum. Accordingly, the new combinations
E. tjallingiorumvar.alnetorum and E. callichroumvar.venustumareproposed.Entoloma lepidissimumvar.pau-
ciangulatum is now treated as a synonym of E. chytrophilum.NeotypesforE. di chroum, E. euchroum and E. lam-
propusaredesignated.
Article infoReceived:22May2013;Accepted:9December2013:Published:1May2014.

142 Persoonia–Volume32,2014
Species Location Collector,vouchernumber GenBankaccessionno.
   nrITS mtSSU nrLSU
E. allochroum Austria K.F.Reinwald,A.Hausknecht(L9860) KC898370 – –
 Netherlands L.Bos(L,7-08-1993,asE. tjallingiorum) KC898368 – –
 Netherlands R.Chrispijn(L14-08-2000) KC898371 – –
 Netherlands KitsvanWaveren(L,29-07-1973,holotype) KC898372 – –
 Russia:Caucasus A.Kiyashko(LE262984) KC898375 – –
 Russia:Caucasus K.Potapov(LE254324) KC898374 – –
 Spain J.Vila,F.Caballero&A.Mayoral(JVG1060902-1) KC898376 KC898488 KC898522
 Spain F.Caballero(EFC1272008-137) KC898455 – –
 Spain F.Caballero(EFC2482008-148) KC898456 – –
 Switzerland G.Wölfel(L:E0809) KC898369 – –
 Germany G.Wölfel(L:E4884,asE. dichroum) KC898373 – –
E. callichroumvar.callichroum Switzerland E.Horak(ZT71/58,holotype) KC898350 – –
E. callichroum var. venustum Belarus P.Kolmakov(LE226909,asE. lepidissimum) KC898356 – –
 Germany G.Wölfel,F.Hampe(L,WöE17/10,holotypeE. venustum) KC898355 KC898490 KC898523
 Russia:Zhiguli E.Malysheva(LE227532,asE. lepidissimum) KC898357 – –
 Russia:Altaj V.Malysheva(LE254312) KC898351 KC898489 KC898521
 Russia:Novosibirsk T.Bulyonkova(LE254313) KC898352 – –
 Russia:Novosibirsk T.Bulyonkova(LE254314) KC898353 – –
 Russia:PrimorskyTerritory M.Nazarova(VLAM-20528,asRhodophyllus lampropus) KC898354 – –
E. chytrophilum Germany M.Enderle,(M,asE. lepidissimumvar.pauci- KC898435 – –
angulatum,holotype)
 Poland M.Wantoch-Rekowska,8Aug.2010 KC898425 – –
 Poland J.Soboń,27Aug.2010 KC898426 – –
 Poland M.Wantoch-Rekowski,3Sept.2010 KC898427 – –
 Russia:Altaj E.Malysheva(LE262994) KC898429 KC898480 KC898520
 Russia:Caucasus E.Malysheva(LE262993) KC898424 – –
 Russia:Caucasus K.Potapov(LE254337) KC898428 – –
 Russia:Novosibirsk T.Bulyonkova(LE254326) KC898430 – –
 Russia:MoscowRegion Yu.Rebriev(LE254325) KC898431 – –
 Russia:VologdaRegion O.Kirillova(LE235259) KC898432 – –
 Russia:NovgorodRegion S.Arslanov(LE254336) KC898433 – –
 Spain:CanaryIslands R.M.Dähncke(L855,holotype) KC898434 KC898479 KC898519
E. coelestinum  Russia:SverdlovskRegion L.Marina(LE258103) KC898362 KC898494 KC898524
E. dichroum Russia:Zhiguli E.Malysheva(LE227472,neotype) KC898440 – –
 Russia:Zhiguli E.Malysheva(LE234260) KC898442 KC898487 KC898528
 Spain J.Vila&F.Caballero(JVG1070821-4) KC898441 KC898486 KC898527
 Spain S.Català(SGC16-10-11) KC898454 – –
E. euchroum Germany L.Krieglsteiner(KR-M-0032474,neotype) KC898421 – –
 Germany L.Krieglsteiner(KR-M-0005673) KC898423 KC898485 –
 Germany M.Scholler(KR-M-0033332) KC898420 – –
 Germany L.Krieglsteiner(KR-M-0032221) KC898422 – –
 Netherlands C.Bas(L6502,asE. tjallingiorum) KC898415 – –
 Russia:Caucasus E.Popov(LE262995) KC898417 KC898483 KC898516
 Russia:LeningradRegion E.Popov(LE254334) KC898416 – –
 Russia:RyazanRegion E.Malysheva(LE254332) KC898419 KC898484 KC898517
 Russia:Tomsk N.Agafonova(LE254329) KC898418 – –
 Spain J.Vila(JVG1020827-2) KC898461 – –
 Spain:CanaryIslands D.Chávezetal.(JVG1091115A) KC898462 – –
E. eugenei Russia:PrimorskyTerritory E.Popov(LE253771holotype) KC898438 – KC898529
 Russia:PrimorskyTerritory T.Svetasheva(LE254347) KC898439 – KC898530
E. lampropus Austria F.Sucti(WU13198,asE. dichroum) KC898379 – –
 Austria A.Hausknecht(WU24148,asE. dichroum) KC898391 – –
 Austria A.Hausknecht(WU10092,asE. dichroum) KC898393 – –
 Germany G.Wölfel(L1509) KC898382 – –
 Russia:BryanskRegion A.Fedosova(LE254339) KC898392 – –
 Russia:Caucasus O.Morozova(LE254316) KC898390 – –
 Russia:KamchatkaRegion O.Morozova(LE254349) KC898389 – KC898507
 Russia:MurmanskRegion L.Mikhailovsky(LE9121,asLeptonia placida) KC898378 – –
 Russia:Novosibirsk T.Bulyonkova(LE262992) KC898388 – –
 Russia:OrenburgRegion O.Desyatova,(LE253584,asE. tjallingiorum) KC898387 – –
 Russia:TatarstanRepublic K.Potapov(LE262991) KC898384 KC898470 KC898505
 Russia:Tomsk N.Agafonova(LE262985) KC898383 – –
 Russia:SverdlovskRegion L.Marina(LE258111) KC898381 – –
 Russia:UdmurtiaRepublic V.Kapitonov(LE254315) KC898386 – –
 Russia:UdmurtiaRepublic V.Kapitonov(LE254338) KC898385 – –
 Russia:VologdaRegion O.Kirillova(LE235263) KC898380 – –
 Sweden T.Læssøe(UPS:BOT:F-176490,asE. placidum, KC898377 KC898471 KC898506
designated here as neotype)
 Spain S.Català(RM0855) KC898458 – –
E. lepidissimum CzechRepublic M.Svrček(PRM755801,holotype,asLeptonia lepidissima) KC898364 – KC898532
 Russia:NovgorodRegion E.Popov(LE234755) KC898365 KC898491 –
 Russia:NovgorodRegion E.Popov(LE254871) KC898363 KC898493 KC898531
 Russia:NovgorodRegion E.Popov(LE234751) KC898367 KC898492 KC898534
 Russia:PrimorskyTerritory E.Malysheva(LE254311) KC898366 – KC898533
Table 1SpecimensandGenBankaccessionnumbersofDNAsequencesusedinthemolecularanalyses.Entoloma subg.Leptonia.

143
O.V.Morozovaetal.:Entoloma subgenus Leptonia
Species Location Collector,vouchernumber GenBankaccessionno.
   nrITS mtSSU nrLSU
E. percoelestinum Russia:Novosibirsk T.Bulyonkova(LE254327) KC898359 KC898496 KC898526
 Russia:Novosibirsk N.Filippova(LE254341) KC898361 – –
 Russia:PenzaRegion A.Ivanov(LE18913,asE.lepidissimum) KC898358 KC898495 KC898525
 Spain J.Vila,X.Llimona(LE254390,holotype) KF745927 – KF745928
 Spain J.Vila,F.Caballero(JVG1061111-7,asE. coelestinum) KC898360 – –
E. placidum Russia:Caucasus O.Morozova(LE254335) KC898397 – –
 Spain J.Carreras,J.Vila,F.Caballero,A.Duran&A.Mayoral KC898395 KC898482 KC898515
  (JVG1060830-6)
 Spain J.Vila,F.Caballero&A.Mayoral(JVG1060902-2) KC898396 – –
 Spain F.Caballero(EFC2682008-151) KC898457 – –
 Sweden S.Lundell(5276)&G.Haglund(UPS:BOT:F-121714, KC898394 KC898481 KC898514
epitype, as Leptonia placida)
E. sublaevisporum Austria A.Hausknecht(MEN9858) KC898437 – –
 Spain J.Vila&F.Caballero(LIPJVG1070823T,holotype) KC898436 KC898478 KC898518
E. tjallingiorumvar.tjallingiorum
 Russia:MoscowRegion E.Lukashina(LE254320) KC898409 – –
 Russia:LeningradRegion R.Singer(LE9123) KC898410 – –
 Russia:UlyanovskRegion E.Ilyukhin(LE254319) KC898408 – –
 Russia:NovgorodRegion O.Morozova(LE254318) KC898411 KC898472 KC898510
 Russia:TatarstanRepublic K.Potapov(LE254317) KC898407 KC898475 KC898511
 Spain F.Caballero(SFC081019-01) KC898459 – –
 Spain F.Caballero(SFC081005-01) KC898460 – –
 Sweden S.Ryman(6124)(UPS:BOT:F-016378,holotype) KC898412 KC898474 KC898509
E. cf.tjallingiorumvar.tjallingiorum Russia:Zhiguli O.Morozova(LE227507) KC898404 – –
 Russia:Zhiguli E.Malysheva(LE227584,asE. placidum) KC898405 – –
 Russia:Zhiguli E.Malysheva(LE234285) KC898406 – –
E. tjallingiorumvar.alnetorum Russia:LeningradRegion E.Popov(LE254321) KC898398 – –
 Russia:TumenRegion E.Zvyagina(LE254322) KC898402 – –
 Russia:Zhiguli E.Malysheva(LE227527) KC898401 – –
 Russia:Zhiguli E.Malysheva(LE234287) KC898403 – –
 Switzerland O.Röllin(29-05-1994) KC898399 – –
 Switzerland O.Röllin(G00111402,holotype) KC898400 KC898473 KC898508
E. tjallingiorumvar.laricinum Russia:KamchatkaRegion E.Popov,O.Morozova(LE254343,holotype) KC898413 KC898477 KC898513
 Russia:KamchatkaRegion E.Popov,O.Morozova(LE254344). KC898414 KC898476 KC898512
Table 1(cont.)
sampling(Table1).ITS1-5.8S-ITS2sequenceswereobtained
forallofthem.Typematerial,ifpossible,wasincludedinthe
analysis.LSU sequences were obtainedfor 1– 3 collections
fromeachtaxon.SpeciesforwhichDNAextractionfromtype
specimens appeared to be impossible or unsuccessful and
wherenoadditionalreliablecollectionswereavailable(E. aus-
triacum, E. cedretorum, E. insidiosum, E. juniperinum, E. klo-
facianum, E. lidbergii, E. syringicolor and E. wynnei) arenot
consideredinthepresentwork.Theoutgroupchoiceandtaxa
sampling to determine the position of studied species in the
system were primarily based on the recent global study on
the phylogeny of the Entolomataceae(Co-Davidetal.2009).
Therefore, the representatives of the main subgenera of the
crown Entoloma clade – Nolanea and Claudopus(Nolanea-
Claudopus clade), Inocephalus and Cyanula(Inocephalus-
Cyanulaclade),aswellasEntoloma, Pouzarella, Alboleptonia,
and Trichopilus were included in the phylogenetic analyses
(Table2).Twospeciesofsubg.Entoloma(Prunuloides clade,
which occupies basal position towards the groups treated
above(Co-David et al. 2009)), wereselected as outgroup
inallanalyses.Atotalof114specimenswasincludedinthe
work.Mostofthesequenceswereobtainedfromthepresent
study.Additional8nrITS1-5.8S-ITS2,19nrLSUand19mtSSU
sequenceswereretrievedfromtheGenbank:withtheacronym
GQ–Co-Davidetal.2009;GU–Baronietal.2011;JQ–He
etal.2013;JX–Vilaetal.2013.Thegeographicoriginofthe
collections includes Europe, the Caucasus and extratropical
AsiafromtheUralstotheRussianFarEast.
Morphological analyses
The study was based both on recently collected material and
collectionskeptin European andAsianherbaria(KR, L, LE,
M,PRM,UPS,VLA,WU,ZT).Thespecimenswerecollected,
documentedandpreservedusingstandardmethods.Macro-
scopic descriptions are based on the study of the fresh material
aswellasonanalysisofthephotos.Thedriedmaterialwas
examined using standard microscopic techniques. Spores,
basidiaandcystidiawereobservedinsquashpreparationsof
smallpartsofthelamellaein5%KOHor1%CongoRedin
concentratedNH4OH.Thepileipelliswasexaminedinaprepa-
rationoftheradialsectionofthepileusin5%KOH.Microscopic
measurementsanddrawingsweremadewithAxioImagerA1
microscopes.Basidiosporedimensionsarebasedonobserving
20 spores, cystidia and basidia dimensions on observing at least
10structurespercollection.Basidiaweremeasuredwithout
sterigmata,andthesporeswithouthilum.Sporelengthtowidth
ratiosarereportedasQ.Thecollected materialisdeposited
intheNaturalisBiodiversityCenter,sectionBotany(L),inthe
MycologicalHerbariumoftheKomarovBotanicalInstitute(LE)
andinthecollectionofJ.Vila(JVG)andS.Català(SGC).The
holotype of E. sublaevisporumisdepositedinLIP(Lille,France).
DNA extraction, amplification and sequencing
DNA was extracted from herbarium material using a CTAB
extraction buffer technique with the following steps of con-
secutive addition of chloroform-isoamyl alcohol mixture, then
isopropylalcohol-3Msodiumacetatesolutionforprecipitation,
70% ethanol for washing and finally water for dissolution.
The alternative method of extraction DNA was using Axy Prep
MultisourseGenomicDNAMiniprepKit(AxygenBiosciences).
TheribosomalITS1-5.8S-ITS2regionwasamplifiedbyPCR
withthe fungal specific primers ITS1Fand ITS4B (Gardes
& Bruns 1993; http://www.biology.duke.edu/fungi /mycolab /

144 Persoonia–Volume32,2014
primers.htm).Sequences of nrLSU-rDNAweregenerated
usingprimersLR0RandLR5(Vilgalys&Hester1990),and
sequencesofmtSSU–usingprimersMS1andMS2(http://na-
ture.berkeley.edu/brunslab/tour/primers.html).PCRproducts
werevisualizedusingagarosegelelectrophoresisandGelRed
staining,andsubsequentlypurifiedwiththekitAxyPrepPCR
CleanupKit(AxygenBiosciences).Sequencingwasperformed
withABImodel 3130GeneticAnalyzer(AppliedBiosystems)
usingBigDyeTMTerminatorCycleSequencingReadyReac-
tionKit(AppliedBiosystems)withthesameprimers.Theraw
datawereprocessedusingSequencingAnalysis5.3.1(Applied
Biosystems).
Alignments and phylogenetic analysis
ThesequenceswerealignedwithMAFFTwebtool(http://align.
bmr.kyushu-u.ac.jp/mafft/online/server/)withQ-INS-Istrategy
anddefaultsettingsforotheroptions.Thefinalalignmentwas
correctedmanuallyusingMEGAversion5(Tamuraetal.2011).
Phylogenetic reconstructions were performed with maximum
likelihood(ML),maximumparsimony(MP)andBayesian(BA)
analyses.Representatives ofthebasalEntoloma clade(Co-
Davidetal.2009),E. turbidum and E. nitidum, were selected
asoutgroupforallanalyses.
MP analysis was performed using PAUP*4.0.b10(Swofford
2002).Onehundredheuristicsearcheswereconductedbystep-
wiseadditionwithrandomsequenceadditionandtreebisection-
reconnection (TBR) branch-swapping algorithm. One tree
washeldateachstepduringstepwiseaddition.Allcharacters
were treated as unordered and of equal weight. Parsimony
bootstrapvalueswerecalculatedfrom1000replicates.Only
cladeswithasupport≥50%wereretained.Gapsweretreated
asmissingdata.
TheMLanalysiswasrunintheRAxMLservers(http://phylobench.
vital-it.ch/raxml-bb/index.php; whichimplements the search
protocolofStamatakiset al.(2008)),underaGTR+Gmodel
withonehundredrapidbootstrapreplicates.
BayesiananalysiswasperformedusingMrBayes3.1(Ronquist
& Huelsenbeck 2003) for two independent runs, each with
2 000 000 generations with sampling every 100 generations,
withGTR+Gmodelandfourchains.Posteriorprobability(PP)
value≥0.95areconsideredsignificant.
Species were delineated on the base of phylogenetic species
conceptreferringtotheexamples from fungi in Tayloret al.
(2000).Monophyletic cladesarerecognizedasphylogenetic
specieswhentheyareconcordantlysupportedbythemajority
ofthereceivedphylogenetictrees.Additionally,ITSsequence
Species Location Collector,voucherNo GenBankaccessionno.
   nrITS mtSSU nrLSU
*Entoloma abortivum Canada H.denBakker(92) – GQ289290 GQ289150
*E. abortivum China:Jilin X.L.Heetal.(HMJAU1955) JQ281483 – –
*E. aprile Japan C.Takehashietal.(TNS:F-24626) AB520845 – –
*E. araneosum Belgium M.E.Noordeloos(200314) – GQ289293 GQ289153
E. cetratum Russia:LeningradRegion O.Morozova(LE235480) KC898450 – –
E. chalybeum Russia:LeningradRegion E.Morozova(LE254353) KC898445 KC898465 KC898500
E. clypeatum Russia:StavropolRegion I.Ukhanova(LE254350) KC898349 KC898497 KC898535
*E. cocles  Finland J.Vauras(9770F) – GQ289299 GQ289159
*E. conferendum Belgium M.E.Noordeloos(200313) – GQ289330 GQ289160
*E. crassicystidiatum China:Guangdong X.L.Heetal.(GDGM27357) – JQ993056 JQ291569
*E. furfuraceum China:Jilin X.L.Heetal.(GDGM28818) – JQ993062 JQ993094
E. griseocyaneum Russia:Caucasus O.Morozova(LE254351) KC898444 KC898463 KC898498
*E. hebes Netherlands C.Hartman(1992-10-28) – GQ289310 GQ289170
E. indutoides Russia:LeningradRegion O.Morozova(LE254354) KC898451 KC898468 KC898503
*E. infula Spain J.Vilaetal.(JVG1080907-13) JX454837 – –
E. inocephalum Vietnam O.Morozova(LE262922) KC898449 – –
E. insidiosum Norway M.E.Noordeloos(L376) KC898443 – –
*E. minutum Spain J.Vilaetal.(LIPPAM00072307) JX454829 – –
E. mougeotii Russia:Caucasus K.Potapov(LE254352) KC898446 KC898464 KC898499
*E. murrayi  V.Hofstetter(VHAs02.02) – GU384590 GU384620
*E. nitidum Italy E.Campo(287) JF907989 – –
*E. nitidum Slovakia M.E.Noordeloos(200426) – GQ289315 GQ289175
*E. parasiticum Belgium M.E.Noordeloos(200330) – GQ289317 GQ289177
*E. porphyrescens Tasmania:Australia M.E.Noordeloos(2004113) – GQ289322 GQ289182
*E. prunuloides Slovakia M.E.Noordeloos(200340) – GQ289324 GQ289184
E. quadratum Russia:PrimorskyTerritory E.Malysheva(LE254355) KC898452 KC898469 KC898504
*E. sericatum Slovakia M.E.Noordeloos(200328) – GQ289329 GQ289189
E. sericellum Russia:Caucasus O.Morozova(LE254362) KC898453 – –
*E. sericellum Belgium M.E.Noordeloos(200315) – GQ289330 GQ289190
*E. sericeum Slovakia M.E.Noordeloos(200329) – GQ289331 GQ289191
E. serrulatum Russia:Caucasus O.Morozova(LE254361) KC898447 KC898466 KC898501
*E. sinuatum Netherlands J.Wisman(2003-09-19) – GQ289333 GQ289193
*E. sordidulum Belgium Co-David(2003) – GQ289334 GQ289194
E. tectonicola India P.Manimohan(741,holotype) – GQ289336 GQ289196
*E. turbidum Italy E.Campo(16176) JF908005 – –
*E. turbidum Slovakia M.E.Noordeloos(200351) – GQ289341 GQ289201
*E. undatum Belgium M.E.Noordeloos(200327) – GQ289342 GQ289202
*E. undatum Italy E.Bizio,E.Campo(16854) JF908007 – –
*E. valdeumbonatum Germany M.Meusers(E4565,holotype) – GQ289343 GQ289203
*E. violaceovillosum India:Kerala P.Manomohan(645,holotype) – GQ289345 GQ289205
E. violaceozonatum Estonia V.Liiv(L275,holotype) KC898448 KC898467 KC898502
Table 2SpecimensandGenBankaccessionnumbersofDNAsequencesusedinthemolecularanalyses.Entoloma subg.Claudopus, Cyanula, Entoloma,
Inocephalus, Nolanea, Pouzarella. Thesymbol*isplacedbeforethenamesofthespeciesforwhichsequenceswastakenfromtheGenbank.

145
O.V.Morozovaetal.:Entoloma subgenus Leptonia
differencesweretakenintoaccount.Geneticdistancesbetween
ITS sequences were estimated using PAUP*4.0.b10 (Swof-
ford2002).Weconsiderap-distancegreaterthan3%tobea
criterionthatwewillusetorecognizenewspecies,following
Petersenetal.(2008)andHughesetal.(2009).Thisapproach
was based on the data for within-species variation and hetero-
zygosityfromtheGreatSmokyMountainsNationalParkinthe
UnitedStates.Accordingtothesedata,aproximatelly2 – 3%
sequencedivergenceusuallyrepresentsdifferentspeciesfor
Basidiomycotina.Morphologicalcriteriawerealsotakenintoac-
count.Inthecasewherethemorphologicaldifferencesbetween
separate monophyletic clades are not evident or p-distance is
lessthan3%weprefertoconsidertheseasvarietieswithina
phylogeneticspecies.
RESULTS AND DISCUSSION
Phylogenetic analysis
Analysis of the nrITS1-5.8S-ITS2 dataset
Thefullalignmentcontained122ITSsequenceswith1047cha-
racters.Wefirstexcludedfromthealignmenttwolargeam-
biguouslyaligned regions inITS1.The first region (282bp)
corresponded to a presumptive insertion characteristic for the
Entoloma dichroumgroup. In addition, E. eugenei from this
grouphadanotherinsertionof51bpthat wasalsoexcluded
fromthefullanalysis.
Theanalyseddatasetincluded720characters(gapsincluded),
ofwhich327wereparsimony-informative.IntheMPanalysis,
the100mostparsimonioustrees(MPTs)weresaved(length=
1738,CI=0.4177,HI=0.5823,RI=0.8402,RC=0.3510).The
ML,MPandBAanalysesrevealednearlyidenticaltopologies.
First of all, the analyses show a well-supported Leptonia clade
(1.0/100  – BA/ MP – hereinafter), separated from both the
Nolanea-Claudopus and the Inocephalus-Cyanulaclades(Fig.
1).As wasshown by Co-Davidet al. (2009)and confirmed
bythepresentwork,subg.Leptonia in the traditional sense
(Noordeloos 1992, 2004) is not monophyletic and must be
considered without sections Cyanula and Griseorubida.Sec-
tion Cyanula recently has been raised to the subgenus level
(Noordeloos & Gates 2012). Representatives of sect. Gri-
seorubida(E. indutoides)arenestedwithin the Inocephalus
subclade, and the exact taxonomic position of the section must
beclarified.Therefore,theavailabledataallowedtreatingsubg.
Leptoniainthe strict sense,correspondingtosect.Leptonia
inthesenseofNoordeloos(1992,2004)andLargent(1994).
The holotype of E. violaceozonatum is grouped together with
E. inocephalum (typespeciesofsubg.Inocephalus)withsig-
nificantsupportofBAvalue(0.95/51)intheInocephalus-Griseo-
rubidum clade;itmustthereforebeexcludedfromsubg.Lep-
tonia.Itis a similarsituationwithE. insidiosum.Aspecimen
identifiedasE. insidiosum is nested within the Cyanula clade
(0.99/ 57), hence itcannot belong tosubg. Leptonia. Unfor-
tunately, we were not successful in DNA isolation from the
holotypeofthisspecies.
TwohighlysupportedsubcladesarerecognizedintheLeptonia
clade(Fig.1,2)correspondingtotwodifferenttaxonomicsec-
tions–sect.Leptonia (1.0/85) and sect.Dichroi(1.0/100),newly
recognizedhere. Entoloma callichroum, E. coelestinum, E. le-
pidissimum and E. allochroum form independent subclades.
Insect.Leptonia twosubcladesarerecognized(Fig.2).First
ofthem–stirpsLeptonia–iscomposedofE. euchroum(type
speciesofsubg.Leptonia), E. lampropus, E. placidum and E.
tjallingiorum. Morphologicallytheweakangledandthinwalled
basidiosporesarecharacteristicforspecieslistedabove.Ento-
loma lampropus, E. placidum and E. tjallingiorum possess also
similarcolours–grey-brownpileusandbluishstipe.Despite
the rather high morphological variability expressed by the
numerouscasesofmisidentificationsofthesetaxainseveral
herbaria, E. euchroum, E. lampropus and E. placidum show
genetic similarity since the internal topology for each species is
unresolvedasthesequencesarealmost100%identical(Fig.
2).NeotypesforE. euchroum and E. lampropus are designated
inthepresentwork.Specimenscollectedascloseaspossibleto
the type locality were chosen: E. euchroumKR-M-0032474from
GermanyandE. lampropusUPS: BOT: F-176490fromSweden,
as well as an epitype for E. placidum–UPS :BOT:F-121714,
collectedclosetotheneotypelocality(asDNAextractionfrom
theneotypewasimpossible).
TheITSsequences for taxaintheE. tjallingiorum clade are
highlyvariable.Evenwiththisextremevariability,itispossibleto
identifyfivewell-supportedsubcladesthatarearrangedinthree
groups(Fig.2).Thefirstgroup(subcladeIandII)includesthe
holotype of the E. alnetorumanditiswellsupported(1.0/ 88).
SubcladeIunitesspecimensmorphologicallycorresponding
to E. alnetorum–withapalecolouredpileus,fruitinginAlnus
spp.forests early in the season(May-June, rarely July(in
northerntaiga), except LE227527, collected inAugust. The
holotype of E. alnetorum isnestedhere.However,subcladeII
includes three specimens from one locality morphologically simi-
lar to E. tjallingiorum.Thesecondgroup(subcladeIIIandIV)
unites specimens morphologically corresponding to E. tjallingi-
orum(withgrey-brownpileus,growinginAugustonsoiloron
wood of various decidious trees), including the holotype of
E. tjallingiorumand itisalsowellsupported(1.0/72).Asthe
p-distance between the holotypes of E. alnetorum and E. tjal-
lingiorumisonly1.3%,weconsiderE. alnetorum a variety of
E. tjallingiorum.The divergence between the sequences as
explained by the geographical distribution can be also recog-
nizedwithin the E. tjallingiorum clade.Subclade III (1.0 /87)
unitesspecimenswith more northern distribution. Itinvolves
holotype of E. tjallingiorum from Sweden, as well as specimens
fromtheLeningrad,NovgorodandMoscowregionsofRussia.
InsubcladeIV(0.99/ 72)specimensarefoundwithsouthern
origin–TatarstanRepublicandUlyanovskRegionofRussia,
andSpain.Thethirdgroup,involvingsubcladeV(1.0/98)only,
takes a rather distant position within the E. tjallingiorumclade.It
canbeexplainedbythegeographicreasonalso–theisolation
betweenEuropeanandFarEasternpopulations(p-distance
betweensequences–3.3%).Morphologicallythisdistinction
is supported by the presence of a sterile lamella edge made up
of abundant, septate terminal elements of the hymenophoral
trama.Alsothehabitatisdifferent,asitisgrowingonconiferous
wood.Inspiteofthep-distancebetweensubcladeVsequences
and holotype of E. tjallingiorumismorethan3%,wedecided
torecognizeanewvarietyE. tjallingiorumvar.laricinum only,
asthemorphologicaldifferencesareonlysmall.
The Entoloma chytrophilum–E. sublaevisporum subclade also
nested within the section Leptonia-clade, despite the rather
isolatedposition.Thesespeciesarecharacterizedbythenodu-
lose or almost nodulose basidiospores with extremely weak
andnumerousangles.Theydefinitelybelongthereforetosect.
Leptonia of subg. Leptonia. The holotype of E. lepidissimum
var.pauciangulatum is nested within the E. chytrophilum-
clade,sothesenamesmustbeconsideredsynonymous.This
evidenceisconfirmedbythemorphologicalfeatures–almost
nodulosesporesandbluecolourofthewholebasidiomata.It
is noteworthy that E. sublaevisporum macromorphologically
is similar to species with differently coloured pileus and stipe,
such as E. placidum, E. lampropus, E. callichroum, and at
firstitwasconsideredcloselyrelatedto E. callichroum. The
phylogenetically informative feature in this case is the shape
ofbasidiospores.Therefore,E. sublaevisporum isrecognized

146 Persoonia–Volume32,2014
E. lampropus
E. placidum
E. tjallingiorum var. alnetorum
E. tjallingiorum var. tjallingiorum
E. tjallingiorum var. laricinum
E. euchroum
E. chytrophilum
E. sublaevisporum
E. callichroum var. callichroum
E. callichroum var. venustum
E. percoelestinum
E. coelestinum
E. allochroum
E. lepidissimum
E. eugenei
E. dichroum
E. insidiosum L 376
E. turbidum JF908005
E. nitidum JF907989
E. chalybeum LE 254353
E. mougeotii LE 254352
E. griseocyaneum LE 254351
E. serrulatum LE 254361
E. sericellum LE 254362
E. quadratum LE 254355
E. inocephalum LE 262922
E. violaceozonatum L 275
E. indutoides LE 254354
E. infula JX454837
E. minutum JX454829
E. cetratum LE 235480
E. undatum JF908007
E. abortivum JQ281483
E. aprile AB520845
E. clypeatum LE 254350
outgroup
Claudopus clade
Inocephalus-Cyanula clade
Inocephalus -
Griseorubidum
clade
Cyanula clade
Section Leptonia clade
Subgenus Leptonia clade
Stirps with uncertain
position
Section
Dichroi clade
Nolanea clade
Rhodopolioid clade
1.0/100
1.0/100
1.0/100
1.0/100
1.0/100
1.0/100
1.0/85
0.92/64
0.95/51
0.95/61
0.99/57
1.0/78
Alboleptonia
- spores nodulose or
almost nodulose
- spores weak angled
- spores moderately angled
- spores with sharp angles
Fig. 1PhylogenetictreederivedfromBayesiananalysis,basedonnrITS1-5.8S-ITS2data.1.Branchesofsubg.Leptoniasubtreecollapsed.Posteriorprob-
ability(PP)valuesfromtheBayesiananalysisfollowedbybootstrapvaluesfromtheMaximumParsimony(BS,%)analysis areaddedto theleftofanode
(PP/BS).PPvalues≥0.95andBSvalues>50%areshown.

147
O.V.Morozovaetal.:Entoloma subgenus Leptonia
/ /
WU 13198 Austria
UPS:BOT:F-176490 Sweden
WU 24148 Austria
WU 10092 Austria
L 1509 Germany
LE 254339 Russia
LE 254316 Russia
LE 254349 Russia
LE 9121 Russia
LE 262992 Russia
LE 253584 Russia
LE 258111Russia
LE 254315 Russia
LE 254338 Russia
LE 235263 Russia
LE 262985 Russia
RM0855 Spain
LE 262991 Russia
UPS:BOT:F-121714 Sweden
JVG 1060830-6 Spain
JVG 1060902-2 Spain
EFC 2682008-151 Spain
LE 254335 Russia
G 00111402 Switzerland
Röllin 29-05-1994 Switzerland
LE 254322 Russia
LE 227527 Russia
LE 234287 Russia
LE 254321 Russia
LE 227507 Russia
LE 227584 Russia
LE 234285 Russia
UPS:BOT:F-016378 Sweden
LE 254318 Russia
LE 254320 Russia
LE 9123 Russia
LE 254319 Russia
LE 254317 Russia
SFC 081019-01 Spain
SFC 08005-01 Spain
LE 254343 Russia
LE 254344 Russia
KR-M-0005673 Germany
KR-M-0033332 Germany
KR-M-0032474 Germany
KR-M-0032221 Germany
L 6502 Netherlands
JVG 1020827-2 Spain
JVG 1091115A Spain
LE 262995 Russia
LE 254334 Russia
LE 254332 Russia
LE 254329 Russia
L 855 Spain
LE 254336 Russia
LE 235259 Russia
LE 254325 Russia
LE 254326 Russia
LE 254337 Russia
LE 262993 Russia
LE 262994 Russia
03-09-2010 Poland
27-08-2010 Poland
08-08-2010 Poland
E. lepidissimum var. pauciangulatum Germany
LIP JVG 1070823T Spain
L MEN 9858 Austria
ZT 71/58 Switzerland
LE 226909 Belarus
L Wö E17/10 Germany
LE 254312 Russia
LE 254313 Russia
LE 254314 Russia
VLA M -20528 Russia
LE 227532 Russia
LE 254327 Russia
LE 254341 Russia
LE 18913 Russia
LE 254390 Spain
LE 258103 Russia
L 29-07-1973 Netherlands
L 9860 Austria
L 07-08-1993 Netherlands
L 14-08-2000 Netherlands
JVG 1060902-1 Spain
EFC 1272008-137 Spain
EFC 2482008-148 Spain
L: E0809 Switzerland
L: E4884 Germany
LE 262984 Russia
LE 254324 Russia
PRM 755801 Czech Republic
LE 234755 Russia
LE 254871 Russia
LE 234751 Russia
LE 254311 Russia LE 253771 Russia
LE 254347 Russia
LE 227472 Russia
SGC 16-10-11 Spain
LE 234260 Russia
JVG 1070821-4 Spain
Inocephalus-Cyanula clade
Nolanea clade
Claudopus clade
Rhodopolioid clade
Subgenus Leptonia clade
Section Leptonia clade
Section
Dichroi clade
Stirps with uncertain position
E. nitidum JF907989
E. turbidum JF908005 outgroup
E. lampropus
E. placidum
I
II
III
IV
V
E. tjallingiorum var. alnetorum
E. tjallingiorum var. tjallingiorum
E. tjallingiorum var. laricinum
E. euchroum
E. chytrophilum
E. sublaevisporum
E. callichroum var. callichroum
E. callichroum var. venustum
E. coelestinum
E. percoelestinum
E. allochroum
E. lepidissimum
E. eugenei
E. dichroum
1.0/100
1.0/100
1.0/100
1.0/99
1.0/100
1.0/85
1.0/100
1.0/96
1.0/98
1.0/98
1.0/98
1.0/88
0.95/63
1.0/72
1.0/100
1.0/100
1.0/100
1.0/100
1.0/100
1.0/100
/69
1.0/100
1.0/100
1.0/100
1.0/100
1.0/100
JVG 1061111-7 Spain
Fig. 2Phylogenetictree derivedfromBayesian analysis,based onnrITS1-5.8S-ITS2data. 2.Mainsubtree –subg.Leptonia clade.Posteriorprobability
(PP)valuesfromtheBayesiananalysisfollowedbybootstrapvaluesfromtheMaximumParsimony(BS,%)analysisareaddedtotheleftofanode(PP/BS).
Typespecimensaremarkedwithredpoints.

148 Persoonia–Volume32,2014
here as a new species due to genetical evidence and spore
morphology(viz.noduloseform)combinedwiththedifference
incolourbetweenthepileusandstipe(seediscussioninthe
taxonomicpart).
Althoughsect. Dichroi has few species, it, however, forms a
clearlydistinctgroup.Morphologicallyitincludesspecieswith
very pronounced, sharply angled spores such as E. dichroum
and E. eugenei.Entoloma allochroum, of uncertain phylogenetic
position,istheonlyotherspecieswithsimilarspores.Section
Dichroi is also distinguished genetically by the presence of large
insertionsintheITS1region(282bp).ITSregioninE. dichroum
is rather variable, but the p-distance between the specimens
studieddoes not exceed 2%. Morphologicalintraspecific
variability in E. dichroumisalsopronounced(asthevariation
incolourofthepileus)(Fig.17a–c).Entoloma dichroum was
describedbyPersoonin1801.Astheholotypedoesnotexist
and the type locality is unknown we selected a neotype from the
availablematerial.Westudied8collectionspreviouslyidentified
as E. dichroum, and only 4 of them correspond to the current
conceptof this species. We selected thereforeLE227472
(Zhiguli,Russia)asitfitsbesttheprotologue.
The E. callichroum-cladeconsistsoftwobranches.One:the
holotype of E. callichroum only and the other including the
holotype of E. venustumand7specimenswhichareconspe-
ciphicwithit.Asthedifferencebetweenthe holotypesislow
(p-distance1.8%bpdifference)andmorphologicalcharacters
vary within the range of genetically identical specimens of
E. venustum(basidiosporesize,presenceandamountofthe
cheilocystidia,theintensityofthebluishtingeinlamellae–see
discussioninthetaxonomicpart)sothatsomespecimensare
indistinguishable from the E. callichroum, we propose to treat
E. venustum as a variety of E. callichroum.
Two clades corresponding to the morphological concept of
E. coelestinum were recovered in all analyses. Specimens
previouslyidentifiedasE. coelestinum form a well-supported
clade, which, however, consists of two sister clades that can be
distinguishedmorphologically.Onecollectioncharacterizedby
more pronounced angled, not nodulose spores and a polished
stipefitswellwiththeprotologueofthetype,andthecurrent
concept of E. coelestinum(Noordeloos2004).Thelargerclade
(5.3%divergencewithprevious)isrepresentedbyspecimens
characterizedbyvaguelyangled,almostnodulosesporesand
alongitudinally fibrillose-striate stipe.Anewspecies, E. per-
coelestinum, is proposed for this taxon below.The E. allo-
chroum clade involves almost identical collections, including
the holotype, and only the Caucasian one slightly stands out
amongthem.The E. lepidissimum clade is also highly sup-
ported,homogeneous,andincludestheholotypespecimen.
The position of E. callichroum, E. coelestinum, E. lepidissimum
and E. allochroumwithin subg. Leptonia clade is uncertain.
IntheBA,MLandMPanalysestheynestinthesectionLep-
toniaclade with lowsupport. Morphologically thesespecies
occupyan intermediatepositionbetweensect. Leptonia and
sect.Dichroi.Thefirstthreespecieshavebasidiosporeswith
moderately developed angles, but E. allochroum has sharply
angledbasidiospores.Itcanbeconcludedthatthedegreeof
development of angles in basidiospores for members of the
Entolomataceae generally is a phylogenetically informative
feature,butwithsomerestrictions.Similarlyshapedbasidio-
sporeshave developedindependentlyinthesisterclades.A
widersamplingofspeciesassignedtosubg.Leptonia from other
geographic regions will be necessary to resolve the phylogeny
withinsubg.Leptonia.
Analysis of the nrLSU and mtSSU datasets
Toaccommodatesubg.Leptonia in an understanding of the
EntolomataceaetreeproducedbyCo-Davidetal.(2009),analy-
sesoftheLSU,mtSSUandcombinedLSU-mtSSUdatasets
were performed on almost all taxa of Leptonia examined in this
study.ThePCRwithMS1andMS2primerswasunsuccess-
ful for E. callichroumvar.callichroum and for E. eugenei.The
analyseswereperformedwithoutthesespecies.
Analysis of the nrLSU dataset
Thedatasetcontained57LSUsequenceswith775characters
(gaps included), of which 162 were parsimony-informative.
The100equallymostparsimonioustrees(MPTs)weresaved
(length=854,CI=0.3255, HI=0.6745, RI=0.4043,RC=
0.1316).Thereweresometopologicaldifferencesandcontra-
dictionsamongtheMPTs,thestrictconsensus treefromthe
MPanalysisthebesttreefromtheMLanalysis,andthe50%
majorityrule consensus treefromtheBA.Therewasahigh
numberofcladesineachofthesetreeswithlowsupport.
Intotal,LSUtreesarelessinformativethannrITSandmtSSU
ones.TopologicalrelationsamongthespeciesofLeptonia and
other subgenera of Entolomawerenotconfirmedbysufficient
supportbecauseoflowdifferencebetweenLSUsequences.
So,wedonotpresentanyLSUtreesinthisworkasambiguous.
Analysis of the mtSSU dataset
Thedatasetcontained 54 mtSSU sequences that represent
the main subgenera of the Entolomas.l.taxacomprisingthe
main branch of the Entolomataceae as well as representatives
of almost all species of Leptonia considered in the present
work. The analysed dataset included 658 characters (gaps
included),ofwhich175wereparsimony-informative.Thesaved
100 most parsimonious trees had the following characteristics:
length=612,CI=0.4776, HI=0.5224, RI=0.7245,RC=
0.3788.TheML,MPandBAanalysesrevealednearlyidenti-
caltopologies.
The main clades of the crown part of the Entolomataceae tree
indicated by Co-David et al. (2009) were mostly confirmed
(Fig.3).TheRhodopolioid(1.0/100),Nolanea(1.0/78),Clau-
dopus(0.98 /69),Pouzarella(0.99/73),Inocephalus-Cyanula
(1.0/ 55)andtheLeptoniaclades(1.0/57)receivedthehighest
support,especiallyinBA.TheholotypeofE. violaceozonatum
was nested in the Inocephalus-Cyanula clade, therefore this
species must be excluded from subg. Leptonia. Inside the
Inocephalus-Cyanula clade three subclades can be revealed:
the highly supported Cyanula and Alboleptonia-Trichopilus
clades and the lowly supported Inocephalus-Griseorubidum
clade.So,the position of theE. violaceozonatum within this
cladeisratheruncertain.
TheanalysesperformedconfirmtheresultsofCo-Davidetal.
(2009),thatsubg.LeptoniainthesenseofNoordeloos(2004)is
polyphyletic and sections Leptonia, Cyanula and Griseorubidum
donotformamonophyleticclade(Fig.3).Cyanula, contain-
ing most known species of former subg. Leptonia, recently
hasbeen raisedtothe subgenuslevel(Noordeloos&Gates
2012).Itischaracterizedbythepresenceofbrilliantgranules,
absenceofclamp-connectionsinalltissuesandafibrilloseto
polishedstipe.WesuggestthattheGriseorubidum must be also
excludedfrom subg.Leptoniabasedon phylogeneticaffinity
and morphological resemblance to Inocephalus species, like
the presence of brilliant granules and well differentiated cheilo-
cystidia.SectionLeptonia(1.0/ 89)clearlystandsoutwithinthe
claderepresented bysubg.Leptonia,andit ischaracterized
morphologically by the presence of clamp-connections, ab-
senceofbrilliantgranulesandmoreorlessfibrillosetosqua-
mulosestipe.
TheanalysesofthemtSSUdatasetconfirmtheseparationof
the new species E. percoelestinum and E. sublaevisporum and
the variety E. tjallingiorumvar.laricinum.

149
O.V.Morozovaetal.:Entoloma subgenus Leptonia
/ /
/ /
E. turbidum GQ289341
E. nitidum GQ289315
E. hebes GQ289310
E. valdeumbonatum GQ289343
E. conferendum GQ289330
E. abortivum GQ289290
E. undatum GQ289342
E. parasiticum GQ289317
E. sinuatum GQ289333
E. sordidulum GQ289334
E. sericatum GQ289329
E. clypeatum LE 254350
E. furfuraceum JQ993062
E. araneosum GQ289293
E. violaceovillosum GQ289345
E. chalybeum LE 254353
E. mougeotii LE 254352
E. griseocyaneum LE 254351
E. serrulatum LE 254361
E. indutoides LE 254354
E. violaceozonatum L 275
E. cocles GQ289299
E. tectonicola GQ289336
E. murrayi GU384590
E. quadratum LE 254355
E. sericellum GQ289330
E. porphyrescens GQ289322
UPS:BOT:F-176490 Sweden
LE 262991 Russia
UPS:BOT:F-016378 Sweden
LE 254317 Russia
LE 254318 Russia
LE 254343 Russia
LE 254344 Russia
UPS:BOT:F-121714 Sweden
JVG 1060830-6 Spain
LE 262995 Russia
LE 254332 Russia
KR-M-0032474 Germany
L 855 Spain
LE 262994 Russia
LIP JVG 1070823T Spain
LE 258103 Russia
LE 254327 Russia
LE 18913 Russia
L Wö E17/10 Germany
LE 254312 Russia
LE 234755 Russia
LE 254871 Russia
LE 234751 Russia
JVG 1060902-1 Spain
LE 234260 Russia
JVG 1070821-4 Spain
outgroup
Nolanea clade
Claudopus clade
Rhodopolioid clade
E. lampropus
E. tjallingiorum var. alnetorum
G 00111402 Switzerland
E. tjallingiorum var. tjallingiorum
E. tjallingiorum var. laricinum
E. placidum
E. euchroum
E. chytrophilum
E. sublaevisporum
E. percoelestinum
E. coelestinum
E. callichroum var. venustum
E. lepidissimum
E. allochroum
E. dichroum
Subgenus Leptonia clade
Pouzarella clade
Inocephalus-Cyanula clade
Alboleptonia-Trichopilus clade
Inocephalus -
Griseorubidum
clade
Cyanula clade
0.99/82
0.99/87
1.0/89
0.96/59
1.0/100
1.0/55
1.0/57
1.0/89
1.0/100
1.0/78
0.98/69
1.0/100
0.98/
0.95/76
1.0/83
1.0/82
1.0/86
/56
0.98/85
1.0/98
1.0/99
1.0/83
0.98/57
1.0/98
1.0/100
0.99/73
Fig. 3PhylogenetictreederivedfromMaximumLikelihood,basedonmtSSUdata.Posteriorprobability(PP)valuesfollowedbybootstrapvaluesfromthe
MaximumParsimony(BS,%)analysisareaddedtotheleftofanode(PP/BS).
Analysis of the combined mtSSU-LSU dataset
The dataset contained 52 sequences as in the previous
analyses. The analysed dataset included 1433 characters
(gapsincluded),ofwhich334wereparsimony-informative.In
theMPanalysis,93mostparsimonioustreeswererecovered
(length=1313, CI=0.4501, HI=0.6113,RI=0.6204, RC=
0.2793).TheML,MPandBAanalysesrevealednearlyidentical
topologies,althoughtheMPbootstrapvalueswereratherlow.
The topology revealed by the analysis of the combined data
issimilartotheresultsoftheanalysisofthemtSSUdata(Fig.
4).Themain subgenera ofEntoloma form their clades with
high support: Rhodopolioidclade(1.0 /99),Nolanea(1.0/97),
Claudopus(1.0/91), Pouzarella (1.0 /57) and Inocephalus-
Cyanulaclade(1.0/),supportedonlybytheBA.Subg.Lepto-
niaclade receivedslightlyhighersupportintheMPanalysis
(1.0/74).Sect.Leptonia(1.0/91)isalsowellrevealed.Etoloma

150 Persoonia–Volume32,2014
allochroum, E. callichroum, E. coelestinum-percoelestinum
clade, E. dichroum, E. lepidissimum occupy isolated uncertain
positionswithinsubg.Leptonia.TheCyanulaclade(1.0/64)and
the Inocephalus-Griseorubidumclade(1.0/)arewellindicated
within the Inocephalus-Cyanulaclade.Etoloma violaceozona-
tum also nests within this clade, but groups together with E. seri-
cellum.Itstaxonomicpositionneedstobecarefullystudiedin
thefuture.
TheanalysesofthecombinedmtSSU-LSUdatasetalsocon-
firmtheseparationofnew species E. percoelestinum and
E. sublaevisporum and the variety E. tjallingiorumvar.lari-
cinum.
/ /
/ /
E. turbidum Slovakia
E. nitidum Slovakia
E. hebes Netherlands
E. valdeumbonatum Germany
E. conferendum Belgium
E. abortivum Canada
E. undatum Belgium
E. parasiticum Belgium
E. sinuatum Netherlands
E. sordidulum Belgium
E. sericatum Slovakia
E. clypeatum LE 254350
UPS:BOT:F-176490 Sweden
LE 262991 Russia
G 00111402 Switzerland
UPS:BOT:F-016378 Sweden
LE 254317 Russia
LE 254318 Russia
LE 254343 Russia
LE 254344 Russia
UPS:BOT:F-121714 Sweden
JVG 1060830-6 Spain
E. tjallingiorum var. laricinum
E. placidum
E. euchroum
L 855 Spain
LE 262994 Russia
LIP JVG 1070823T Spain
LE 234260 Russia
JVG 1070821-4 Spain
L Wö E17/10 Germany
LE 254312 Russia
LE 254327 Russia
LE 18913 Russia
LE 258103 Russia
JVG 1060902-1 Spain
LE 254871 Russia
LE 234751 Russia
E. griseocyaneum LE 254351
E. mougeotii LE 254352
E. chalybeum LE 254353
E. serrulatum LE 254361
E. violaceozonatum L 275
E. sericellum Belgium
E. indutoides LE 254354
E. cocles Finland
E. murrayi VHAs02.02
E. quadratum LE 254355
E. tectonicola India
E. porphyrescens Tasmania
E. violaceovillosum India
E. araneosum Belgium
E. furfuraceum GDGM 28818
outgroup
Nolanea clade
Claudopus clade
Rhodopolioid clade
E. lampropus
E. tjallingiorum var. alnetorum
E. tjallingiorum var. tjallingiorum
E. chytrophilum
E. sublaevisporum
E. dichroum
E. callichroum var. venustum
E. lepidissimum
E. allochroum
E. percoelestinum
E. coelestinum
Cyanula clade
Inocephalus -
Griseorubidum
clade
Pouzarella clade
Subgenus Leptonia clade
1.0/100
1.0/74
1.0/91
/91
0.98/50
1.0/76
1.0/100
1.0/87
1.0/96
Inocephalus-Cyanula clade
1.0/64
1.0/86
1.0/77
1.0/57
0.99/
1.0/
0.95/
1.0/100
1.0/
1.0/97
0.97/97
1.0/74
1.0/91
1.0/99
0.99/87
/89
1.0/67
1.0/100
1.0/100
1.0/100
1.0/100 LE 262995 Russia
LE 254332 Russia
1.0/100
1.0/100
1.0/100
1.0/100
1.0/100
Fig. 4PhylogenetictreederivedfromMaximumLikelihood,basedoncombinedmtSSU-nrLSUdata.Posteriorprobability(PP)valuesfollowedbybootstrap
valuesfromtheMaximumParsimony(BS,%)analysisareaddedtotheleftofanode(PP/BS).

151
O.V.Morozovaetal.:Entoloma subgenus Leptonia
TAXONOMIC PART
Entoloma subgenus Leptonia(Fr.:Fr.)Noordel.emend.O.V.
Morozova,Noordel.&Vila
Entolomasubg.Leptoniasect.Leptonia (Fr.:Fr.)Noordel.,Persoonia11:146.
1981.—Agaricustrib.LeptoniaFr.,Syst.Mycol.(Lundae)1:10,201.1821.
—Lectotype(Clements&Shear1931:349):Agaricus euchrousPers.:Fr.
Rhodophyllussect.LeptoniariiRomagn.,Bull.Soc.Mycol.France53:332.
1937 (nom. nud.; no Latin diagnosis). — Lectotype (Noordeloos 1981:
146):Agaricus euchrousPers.:Fr.
Leptoniasect.Lampropodae Konrad& Maubl.,LesAgaricales:259.1948
(nom.nud., noLatin diagnosis).—Lectotype (Noordeloos1982a: 453):
Agaricus lampropusPers.:Fr.
Rhodophyllussect.Lampropodes(Kühner&Romagn.,Fl.Anal.:208.1953)
exRomagn.,Bull.Mens.Soc.Linn.Lyon43:328.1974.—Holotype:R. lam-
propus(Pers.:Fr.)Quél.
Habit mycenoid, collybioid or tricholomatoid; pileus conico-
convex or plano-convex, rarely with papilla or umbo, sometimes
depressed; lamellae almost free to emarginate, sometimes
adnate,rarely with decurrent tooth; stipefibrillose-striate or
flocculose-scaly,rarelyalmostsmooth;lamellae edge sterile,
fertileorheterogeneous;cheilocystidiapresentorabsent,ba-
sidiospores with both weak or sharp angles, or almost nodulose,
pileipellis more or less a trichoderm of inflated elements with
intracellular,andoftenadditionallyencrusting,pigment;brilliant
granulesabsent;clamp-connectionspresentandoftenfrequent;
lignicolousorterrestrial,mostlyinforests.
Entolomasubg.Leptonia(Fr.:Fr.)Noordel.isemendedhere
by excluding sections Cyanula(Romagn.)Noordel.(thatnowis
consideredsubg.Cyanula(Romagn.)Noordel.(Noordeloos&
Gates2012)),Griseorubida(Romagn.)Noordel.andRhampho-
cystotae(Largent)Noordel.Thesubgenusnowischaracterized
byspecies previouslyconsideredinsect. Leptonia only, and
characterizedbythepresenceofclamp-connections,absence
ofbrilliantgranules,andmoreorlessfibrillosetosquamulose
stipe.
Synopsis of the phylogenetic species in subgenus
Leptonia from boreal-temperate Eurasia
Section Leptonia (Fr.:Fr.)Noordel.emend.O.V.Morozova,
Noordel.&Vila
 1. Entoloma chytrophilum Wölfel,Noordel.&Dähncke
 2. Entoloma euchroum(Pers.:Fr.)Donk
 3. Entoloma lampropus(Fr.:Fr.)Hesler
 4. Entoloma placidum(Fr.)Noordel.
 5. Entoloma sublaevisporumVila,Noordel.&O.V.Morozova
 6a. Entoloma tjallingiorum Noordel.var.tjallingiorum
 6b. Entoloma tjallingiorumvar.alnetorum(Monthoux&Röllin)
O.V.Morozova,Noordel.&Vila
 6c. Entoloma tjallingiorum var.laricinum O.V. Morozova,
Noordel.,Vila&E.S.Popov
Section Dichroi O.V.Morozova,Noordel.&Vila
 7. Entoloma dichroum(Pers.:Fr.)P.Kumm.
 8. Entoloma eugeneiNoordel.&О.V.Morozova
Species of uncertain position
Stirpe Allochroum
 9. Entoloma allochroumNoordel.
Stirpe Callichroum
10a. Entoloma callichroum E. Horak & Noordel. var.calli-
chroum
10b. Entoloma callichroumvar.venustum(Wölfel&F.Hampe)
O.V.Morozova,Noordel.&Vila
Stirpe Coelestinum
 11. Entoloma coelestinum(Fr.)Hesler
 12. Entoloma percoelestinumO.V.Morozova,Noordel.,Vila
& Bulyonkova
Stirpe Lepidissimum
 13. Entoloma lepidissimum(Svrček)Noordel.
Certainly, the list of potential phylogenetic species within
subg.Leptonia is much larger.Several morphospecies are
still in need of molecular analysis in order to determine their
phylogeneticposition.Thefollowingadditionalmorphospecies
are known from boreal and temperate Eurasia: E. austriacum
Courtec., E. cedretorum (Romagn.&Riousset)Noordel., E. in-
sidiosum Noordel., E. juniperinum Barkman&Noordel., E. klo-
facianum Noordel.,Wölfel&Hauskn., E. lidbergii Noordel. and
E. wynneae (Berk.&Broome)Sacc.(Noordeloos2004);E. sy-
ringicolorE.Ludw.&Noordel.,E. dichroumvar.leptosporum
E.Ludw.&Noordel. (Ludwig 2007); E. dichroumvar.corsi-
cum(Romagn.)Courtec.(Courtecuisse2008);E. legionense
Blanco-Dios(Blanco-Dios‘2012’2013).Inthepresentstudy,
the DNA extraction of these morphospecies failed, either due
totheageandconditionofthe(type)specimens,ormaterial
wasnotavailablefordestructivesampling.
Forasoundphylogenyofsubg.Leptonia, extra-European spe-
cies(someofwhicharelistedbelow)mustalsobetakenintoac-
count,viz.fromNorthAmerica:E. velatum Hesler(Hesler1967);
E. kauffmaniiMalloch (Malloch2010); E. cyaneum [Peck→]
Sacc.,Leptonia approximataLargent,L. carneaLargent,L. con-
vexaLargent,L. cyaneonitaLargent,L. insuetaLargent,L. occi-
dentalisMurrill,L. subeuchroa Kauffman, L. subgracilisLargent,
L. violaceonigraLargent,L. subcoelestinaLargent,L. violacea
(Kauffman)Largent,L. zanthophyllaLargent(Largent1994);
Leptoniella acericolaMurrill; from India: E. indoviolaceum
Manim.&Noordel.(Manimohanetal.2006);fromNewGuinea:
E. egregiumE.Horak(Horak1980);andfromAustralia:E. pan-
niculus (Berk.) Sacc., E. tomentosolilacinum G.M. Gates &
Noordel.,E. violascens G.M.Gates&Noordel.andE. endotum
Noordel.&G.M.Gates(Noordeloos&Gates2012).
Key to the species of subgenus Leptonia in
boreal and temperate Eurasia
For the convenience in identifying following morphospecies have been added
tothekey,onaccountoftheevidenceinNoordeloos(2004):E. austriacum,
E. cedretorum, E. insidiosum, E. juniperinum, E. klofacianum, E. lidbergii
and E. wynnei.AlsoE. violaceozonatumNoordel.&Liivisincluded,despite
its placement in the Inocephalus-Cyanulaclade.
1. Pileusandstipeblue,violaceousorpinkishlilaceous;lamel-
lae white or coloured when young ..................2
1. Pileusyellow-brown,brown,grey-brownorochraceous,some-
timeswith blue or violaceous tinge near thecap margin;
stipeblueorviolaceous;lamellaewhitewhenyoung.. 16
2. Lamellaecolouredwhenyoung .................... 3
2. Lamellaewhitewhenyoung......................7
3. Lamellaefirstlybrownishthenreddishbrownwithfimbriate
concolorous edge; pileus fibrillose-zonate, lamellae edge
entirelysterilewithdenseclustersofverylong(upto300μm)
subcylindrical cheilocystidia ........ E. violaceozonatum
3. Lamellaewithbluishorviolaceous-bluetingewhenyoung,
lamellae edge sterile or heterogeneous, cheilocystidia of
another type, smaller ............................4
4. Lamellae violaceous-blue with brownish violaceous edge
whenyoung;fruitbodiesentirelydeepviolaceous-blue;on
wood, preferably of Alnus.............2.E. euchroum
4. Lamellaepale tomoderately darkblue orviolaceous-blue
with concolorous or slightly paler edge when young....5
5. Pileusandstipewiththesamedeepbluecolour,lamellaeblu-
ishwithconcolorousorslightlypaleredge;spores7.5–11.5
×6.0–8.0μm ....................13.E. lepidissimum
5. Pileuspinkishlilaceoustoviolaceous,stipedarkblue. . 6

152 Persoonia–Volume32,2014
6. Sporeswithbluntangles,cheilocystidiaabsent .......
................10a.E. callichroum var.callichroum
6. Sporeswithmoderatelypronouncedangles,cheilocystidia
broadly clavate, intermixed with basidia.............
..................10b.E. callichroum var. venustum
7. Pileusandstipeblueorviolaceous-blue ............8
7. Pileusandstipeviolaceous,brown-violaceousorpinkishlila-
ceous ......................................14
8. Sporesisodiametrical,upto9μmdiam . E. klofacianum
8. Sporesheterodiametrical....................... 9
9. Spores 6.5 –8.5 (– 9.0) × 5.5– 6.5 μm, pileus conical
to campa nulate, hardly expanding, stipe smooth to
subfibrillose................................. 10
9. Sporeslarger,pileushemisphaerical,convextoapplanate,
stipefibrilloseorminutelysquamulose............ 11
10. Sporesmoderatelyangled,stipesmoothtosubfibrillose
...............................11.E. coelestinum
10. Sporesweaklyangled,stipefibrilloseorminutelysquamu-
lose........................12.E. percoelestinum
11. Sporesdistinctlynodulose,8.5–11.5×6.0–7.0μm,pileus
convex to applanate .............. 1.E. chytrophilum
11. Sporeswith5–7pronouncedangles .............. 12
12. Cheilocystidiaabsent................E. cedretorum
12. Cheilocystidiapresent.........................13
13. Fruitbodycollybioid..................7.E. dichroum
13. Fruitbodytricholomatoid...............8.E. eugenei
14. Sporeswith7–9 weakangles; 9.0–12.2 ×6.0 – 8.2 μm;
pileus a tender pinkish ochre with brown with lilac or vio-
laceous tint at centre.................E. austriacum
14. Sporeswith5–7pronouncedangles.............. 15
15. Fruitbodiesverysmall;pileusupto10mm,pink......
.....................................E. lidbergii
15. Fruitbodieslarger;pileus20–40mm,greyishbrownwith
lilaceous tinge.................... 9.E. allochroum
16. Spores 10.0 –16.0 (–18.0) × 7.0–10.0 μm; pileus warm
brown contrasting with the blue stipe........ E. wynnei
16. Sporessmaller.............................. 17
17. Sporeswith5–7pronouncedangles............. 18
17. Sporesnoduloseorwith6–9weakangles ......... 19
18. Stipeblue,flocculose-squamulose.................
........................ 7.E. dichroum(paleform)
18. Stipeviolaceous,longitudinallyfibrillose 9.E. allochroum
19. Sporesdistinctlynodulose...... 5.E. sublaevisporum
19. Sporeswith6–9weakangles..................20
20. Stipeglabrous,polished.......................21
20. Stipelongitudinallyfibrilloseorsquamulose........ 23
21. Sporesisodiametricaltosubisodiametrical.........22
21. Sporesheterodiametrical............. E. insidiosum
22. Spores8.0 –10.0×6.0–8.0μm;pileusdarkbrownwhen
young ........................... E. juniperinum
22. Spores7.5–9.0×6.5–8.5 μm; pileus blue when young
................................. E. klofacianum
23. Stipeflocculose-squamulose................... 24
23. Stipefibrillose-striate ..........................26
24. Pileuspale ochraceous;onwoodofAlnus, early in sea-
son ..............6b.E. tjallingiorum var.alnetorum
24. Pileusbrownorgrey-brown.................... 25
25. Lamellaeedgeheterogeneous,cheilocystidialageniform,
intermixedwithbasidia;onsoiloronwoodofdeciduous
trees ............6a.E. tjallingiorum var. tjallingiorum
25. Lamellaeedgesterile,cheilocystidiaasstrandsofterminal
elements of hyphae of trama hymenial..............
................... 6c.E. tjallingiorum var. laricinum
26. Pileipelliswithintracellularpigment;cheilocystidiaabsent;
smellfarinaceous;onwoodofdeciduoustrees,mainlyof
Fagus ........................... 4.E. placidum
26. Pileipelliswithbothintracellularand encrustingpigment;
cheilocystidiapresentorabsent;smellnone;onsoiloron
wood of coniferous trees............3.E. lampropus
Descriptions of the species
Below we provide descriptions of the species included in the
analysis(inthekeyindicatedwiththenumberthatcorresponds
withthedescriptivetextofthispaper).Mostofthemarerarely
collectedandexactandcompleteinformationwasdifficultto
find.Thereforethedescriptionsbelowcontainnewdataonecol-
ogy, substratum, colour variation, spore shape, and presence/
absenceofcheilocystidia.Onesection,twospeciesandone
varietyaredescribedasnewtoscience.
Section Leptonia (Fr.:Fr.)Noordel.emend.O.V.Morozova,
Noordel.&Vila
 Lectotype(Clements&Shear1931:349):Agaricus euchrousPers.:Fr.
Habitcollybioidtoalmosttricholomatoid;pileusconico-convex
or plano-convex, rarely with papilla or umbo, sometimes
depressed; lamellae almost free to emarginate, sometimes
adnate,rarely with decurrent tooth; stipefibrillose-striate or
flocculose-scaly; lamella edge sterile, fertile or heterogene-
ous;cheilocystidia present or absent, spores with poor angles
to almost nodulose, pileipellis more or less a trichoderm of
inflated elements with intracellular, and often additionally en-
crusting,pigment;pileitramaregular;brilliantgranulesabsent;
clamp-connectionspresentandoftenfrequent;lignicolousor
terrestrial,mostlyinforests.
Section Leptonia is emended here to unite only those species
with weakly angled, almost nodulose spores, which form a
well-supported monophyletic clade, as opposed to the new sec-
tion Dichroi which includes species with pronouncedly acutely
angled spores. Several species with uncertain phylogenetic
position(Entoloma allochroum, E. callichroum, E. coelestinum,
E. lepidissimum, E. percoelestinum)also arenotincludedin
sect.Leptonia.
1. Entoloma chytrophilum Wölfel,Noordel.&Dähncke,Öst.
 Z.Pilzk.10:190.2001.—Fig.5a,b,6
Syn.:Entoloma lepidissimumvar.pauciangulatumGminder&Enderle,Beitr.
Kenntn.PilzeMitteleur.10:60.1996.
Pileus 2 –15 mm broad, plano-convex to concave with de-
pressed centre, not hygrophanous, not translucently striate,
withstraightthendeflexedundulatingmargin,radiallyfibrillose,
incentresquamulose,darkblue,slightlydiscolouringtobluish
violaceus.Lamellae moderately distant, adnate-emarginate with
small decurrent tooth, ventricose, white, becoming pinkish, with
entireconcolorousedge.Stipe20 – 40×1–2mm,cylindrical,
fibrillose-striate,slightlysquamuloseintheupperpart,concol-
ourouswithpileusorgrey-blue,with white basal tomentum.
Smell strong, fungoid, reminiscent of Cantharellus cibarius or
indistinct,tastenotreported.Spores8.4–11.5×5.8 –7.0μm,
Q=1.3–1.8,nodulose,heterodiametrical.Basidia37.8–48.2×
10.2–12.3μm,1–4-spored,clavate,clamped.Lamellae edge
fertile.Cheilocystidiaabsent.Pileipellis a plagiotrichoderm of
cylindricaltoslightlyinflatedhyphae10–20μmwidewithblue
intracellularpigment.Clamp-connectionspresent.
 Habitat—TypespecimenonchipsofPinus bark in pot with
Cymbidium in garden. In natural conditions on rotten wood
(mostlyofconiferoustrees)inconiferousandmixedforests.
 Knowndistribution—Western,CentralandEasternEurope,
CanaryIslands(holotype),Caucasus,WesternSiberia.

153
O.V.Morozovaetal.:Entoloma subgenus Leptonia
Fig. 5 a, b: Entoloma chytrophilum.a.LE262994;b.LE254326.—c– f:E. euchroum.c,d.LE254329(photo’stakenatthesamelocalitybutatdifferenttimes);
e.JVG1091115A;f.LE262995.―Scalebars=1cm.—Photosby:a,f.O.Morozova;b.T.Bulyonkova;c.N.Agafonova;d.S.Gashkov;e.M.Á.Ribes.

154 Persoonia–Volume32,2014
Specimens examined.Germany,Baden-Württemberg,Langenau-Lindenau,
500mNN, MTB7426 /4, amongconiferneedles,9Aug.1985,M. Enderle,
(M,as Entoloma lepidissimum var.pauciangulatum, holotype). – Poland,
TrójmiejskiLandscapePark,Gdansk,Oliwadistricy,onPicea abies cones in
mixedforest(Fagus sylvatica, Pinus sylvestris, Picea abies, Betula pendula),
8Aug. 2010, M. Wantoch-Rekowska;ibid.,on Fagus cupules and woody
debrisinmixedforest(Fagus sylvatica, Pinus sylvestris, Picea abies, Betula
pendula),3 Sept. 2010, M. Wantoch-Rekowski; Sudety Mts, Kaczawskie
foothills, Mount Szeroka, on Pinus branches in Pinus sylvestris-Larixsp.
forest,27Aug.2010,J. Soboń. –russia,VologdaRegion, ‘Russky Sever’
National Park, Nilovetskoye forestry, on soil in Picea abies-Pinus sylvestris
forest,20Aug.2004,O. Kirillova(LE235259,asE. lepidissimum);Novgorod
Region,MalayaVisheraDistrict,vicinitiesofSyujskavillage,amongmosses
in Picea abies-Populus tremulaforest,7Aug.2012,S. Arslanov(LE254336);
MoscowRegion,ZvenigorodBiologicalStationofMoscowStateUniversity,on
rotten stump, 24 July 2012, Yu. Rebriev(LE254325);Karachaevo-Cherkesia
Republik,TeberdaBiosphereReserve,valleyofBadukRiver,onrottenstump
in Picea orientalis -Abies nordmanniana forest,8Aug. 2009,E. Malysheva
(LE262993); ibid., vicinities of Teberda, on rotten stump, K. Potapov, 22
Aug.2012(LE254337);AltajRepublic,AltajskyNatureReserve,Atkichu,on
rottenstumpsinfloodplainforest,18Aug.2008,E. Malysheva(LE262994);
Novosibirsk,Akademgorodok,onrottenbirch stumpnear theICG SBRAS
moselanimalbreedingfacility,15June2008,T. Bulyonkova(LE254348);ibid.,
onrottenpinestumpinmixedforest(Pinus sylvestris, Acer negundo, Betula
pendula)nearICGSBRASmodule,12Aug.2011,T. Bulyonkova(LE254326);
ibid.,onrottenmossystumpinyoungbirchgroveborderingmixedforest,with
old rotting stumps, 14 July 2010, T. Bulyonkova(LE254328).–sPain, Canary
Islands,LaPalma,inapotwithanorchid,31Aug.1994,R.M. Dähncke(855,
L,holotype).
 Notes—Thisspecieshasoriginallybeendescribedfrom
theislandofLaPalma (IslasCanarias,Spain), whereitwas
found on chips of Pinus bark in pot with Cymbidiuminagarden.
After that it was collected several times in different types of
naturalhabitatsinEurope,Caucasus,WesternSiberia.These
recordsfitthe original descriptionwell(Wölfel&Noordeloos
2001,Noordeloos2004),andmoleculardataconfirmthecon-
specifityofthenewrecordswiththeholotype.Entoloma chy-
trophilumcan berecognizedbythebright bluecolourofthe
fruitbodies, plano-convex shape of the pileus and lignicolous
habitonconiferouswood.Fromtheotherbluecolouredspe-
cies(E. coelestinum, E. dichroum, E. lepidissimum) itdiffers
firstofallbytheratherlarge,thin-walledandnodulosespores.
Entoloma lepidissimum var.pauciangulatum must be con-
sidered a synonym of E. chytrophilum due to morphological
(almostnodulosesporesandbluecolourofbothpileusand
stipe(Gminder&Enderle1996))andphylogeneticevidence.
Despitealaterpublication,theepithet‘chytrophilum’ has prior-
ityover‘pauciangulatum’attherankofspecies,becausethe
name‘pauciangulatum’haspriorityonlyintherankofvariety
inwhichitwaspublished(Art.11.2).
2. Entoloma euchroum (Pers.: Fr.)Donk, Bull. Bot. Gard.
Buitenzorg,ser.III,18:157.1949.—Fig.5c– f,7
Syn.:Agaricus euchrousPers.,Syn.Meth.Fung.(Göttingen)2:343.1801;
Agaricus euchrousPers.:Fr.,Syst.Mycol.1:203.1821;Leptonia euchroa
(Pers.:Fr.)P.Kumm.,FührerPilzk.(Zwickau):24,96.1871;Rhodophyl-
lus euchrous (Pers.: Fr.)Quél., Enchir. Fung.: 60. 1886; Hyporrhodius
euchrous(Pers.:Fr.)J.Schröt.inCohn,Krypt.-Fl.Schlesien(Breslau)3.1
(33– 40):615.1889.
Neotype (designated here). Germany, Baden-Württemberg, Schwäbisch-
FränkischerWald,Ks.Ostalbkreis,Durlangen,WTäferrot,Rottal,onAlnus
sp.,25Aug.2005,L. Krieglsteiner(KR-M-0032474).
Pileus 5 –40 mm broad, broadly conical, hemisphaerical or
campanulate, then convex, often with central depression, not
hygrophanous,not translucently striate, radially fibrilloseto
squamuloseall over, violaceous blue,discolouring dark or
palebrownwithviolaceousorpurplishtint.Lamellae adnate-
emarginate, to adnexed often with decurrent tooth, violaceous
blue, dark brownish violet, or sometimes discolouring to pale
blue, with irregular to serrulate brownish, violaceous or bluish
edge.Stipe20–70×1–6mm,cylindrical,slightlybroadenedto
baseordistinctlybulbose,solid,longitudinallyfibrillose,squa-
mulose in the upper part, violaceous blue, then brownish blue
withviolaceousorpurplishtint,basewhitelytomentose.Context
concolorouswithsurfaceorpaler.Smell sweet, aromatic like
violetflowers,orindistinct,tasteunpleasant.Spores9.0–11.5
×6.0– 8.0μm,Q=1.2–1.7,heterodiametricalwith6–8rather
bluntanglesinsideview.Basidia45.8– 48.3×10.1–12.8μm,
4-spored,clavate, clamped. Lamellae edge heterogeneous.
Cheilocystidia cylindrical, narrowly clavate or lageniform, some-
times septate, often thick walled in the upper part, hyaline or
Fig. 6 Entoloma chytrophilum.a.Basidium;b.spores(LE262994).―Scale
bars=10μm.
Fig. 7 Entoloma euchroum. a.Spores;b.basidium;c. cheilocystidia(KR
0032474,neotype).―Scalebars=10μm.

155
O.V.Morozovaetal.:Entoloma subgenus Leptonia
withviolaceousbrowncontent.Pileipellis a trichoderm in the
centre, plagiotrichoderm towards the pileus margin, made up
ofcylindricalhyphaewithinflatedterminalelements,8–15μm
wide with blue-violaceous or violaceous brown intracellular
pigment.Caulocystidia present as septate clamped hairs up
to300μm,colourlessorwithblue-violaceousintracellularpig-
ment.Hyphaeofthetramaofboththepileusandthestipewith
incrustedpigment.Clamp-connectionsabundant.
 Habitat—Ondeadandlivingdeciduoustrees(mostlyAlnus
but also Quercus, Sorbus, Corylus, Acer, Prunus),exceptionally
onconiferoustree(JVG 1091115A;Noordeloos1982a).
 Knowndistribution—WesternandEasternEurope,Canary
Islands,Caucasus,WesternSiberia,RussianFarEast.
Additional specimens examined. Germany, Brandenburg, Ks. Ucker-
mark, Gartz, NSG Fauler Ort, on Alnus sp., 22 Sept. 1990, M. Scholler
(KR-M-0033332);Bayern,Rhön,Gefäll,Seebachtal,Buntsandstein,450m,
MTB/ Q 5625/4, Carici remotae-Fraxinetum, on Alnus sp., 11Oct. 2003,
L. Krieglsteiner (KR-M-0005673); Baden-Württemberg, Lorch, N Wacht-
haus, Haselbachtal, Welzheimer Wald,on Quercus robur, 6 Sept. 2007,
L. Krieglsteiner(KR-M-0032221). – netherlands, Bloemendaal, Koningshof,
2Nov.1974,C. Bas(L6502,asE. tjallingiorum).–russia,LeningradRegion,
KingiseppDistrict,Kotelsky Sanctuary,Sept.2005,E. Popov (LE254334);
NovgorodRegion,vicinitiesoftheOpechenskyPasad,GornayaMsta,onthe
Alnus glutinosa stump, Yu. Rebriev(LE253879);Moscow Region,Taldom
District,BolshoyeStrashevo,27Sept.2010,C. Lukashin(LE254333);Ryazan
Region,OkskyNatureReserve,onAlnus glutinosa,2006,E. Malysheva
(LE254332);StavropolTerritory,onAcer platanoides (?)stump,25Aug.2009,
I. Ukhanova(LE254331); Karachaevo-Cherkesia Republic, Teberdinsky
NatureReserve,valleyofTeberdaRiver,onthebaseofAlnus glutinosa, 10
Aug.2009,E. Popov(LE262995);Tomsk,intheplantingofbirchandpine,
on Prunus padusstump,2Oct.2007,N. Agafonova(LE254329).–sPain,
Saga,Cerdanya, Girona,alt. 1050m,onAlnus glutinosa decaying wood,
27Aug.2002,J. Vila(JVG1020827-2); CanaryIslands,LaMalgarida,La
Palma,alt.1080m,ondecayingwood of Pinus canariensis,15Nov.2009,
D. Chávez, V. Escobio, J.F. López, J.I. Velaz, J.L. Lantigua and M.Á. Ribes
(JVG1091115A).
 Notes—Entoloma euchroum is a very distinctive species
distributedalloverEuropeandreportedalsofromSiberia.Usu-
allyitiseasytorecognizedue toitsentirelyblue-violaceous
basidiomes, lignicolous habitat and sweet, flowerlike smell.
Sometimes it can be rather variable in colour, depending on the
growing conditions, but violaceus blue tinges are always present
inallpartsofthebasidioma,especiallyinthelamellae.Itcan
be distinguished from the other taxa that can possess more or
lessviolaceous-bluelamellae(E. callichroumvar.venustum,
E. lepidissimum)bythepresenceofcheilocystidiawithathick-
walledupperpart, often filledwithviolaceousbrowncontent
cheilocystidia.CheilocystidiaareabsentinE. callichroumvar.
venustum, rare, and intermixed with basidia in E. lepidissimum.
Alsothesquamulosestemandlignicoloushabitathelptodis-
tinguish E. euchroum.
3. Entoloma lampropus(Fr.:Fr.)Hesler,Beih.NovaHedwigia
23:154.1967.—Fig.8,9a,b
Syn.:Agaricus lampropus Fr.,Observ.Mycol.1:19.1815; Agaricus lam-
propusFr.:Fr.,Syst.Mycol.1: 203. 1821;Leptonia lampropus(Fr.:Fr.)
Quél.,Mém.Soc.Émul.Montbéliard,sér.2,5:121.1872;Rhodophyllus
lampropus(Fr.:Fr.)Quél.,Enchir.Fung.:60.1886.
Excl.:Rhodophyllus lampropussensuJ.E.Lange,Fl.Agar.Dan.2,pl.76C.
1937[=?E. corvinum(Kühner)Noordel.];Leptonia lampropus sensuBres.,
Iconogr.Mycol.XII,pl.570-1.1929;P.D.Orton,Trans.Brit.Mycol.Soc.43,
Suppl.:105.1960;Entoloma lampropussensuHesler,Beih.NovaHedwigia
23:154.1967[=E. sodaleKühner&Romagn.exNoordel.].
Neotype(designatedhere):sweden,Medelpad,Liden,Sundsjöåsen,T. Læssøe
(asE. placidum),31Aug.1999(UPS:BOT:F-176490).
Pileus10–40mmbroad,conicalorhemisphericalexpanding
to plano-convex with incurved margin and usually slightly de-
pressed centre, not hygrophanous, not translucently striate,
entirelyradially fibrillosetosquamuloseatcentre,withsmall
darkgrey-brownsquamulesonpalerbackground,mostdense
at centre, varying from rather pale beige-grey to moderately
dark grey-brown, often darker, almost black in centre, some-
times with bluish or lilac tint especially near pileus margin but
notveryoften.Lamellae moderately distant, narrowly adnate,
emarginate or adnate with small decurrent tooth, sometimes
arcuate, whitish to cream becoming greyish or brownish pink,
with irregular concolorous edge. Stipe 25 –65 × 1– 3.5 mm,
cylindrical or slightly broadened towards base, often twisted,
steel blue, brownish or greyish blue, distinctly longitudinally
fibrillose-striatewith dark blue fibrils, usuallyglabrous but
sometimes flocculose with white or bluish floccules in upper
part,basewithwhitetomentum.Fleshwhitish,bluebeneaththe
stipesurface.Smellnotdistinctive,tastemildorslightlybitter.
Spores8.8–11.5×6.0–7.5μm,Q=1.3–1.7,heterodiametrical,
with6 – 9ratherbluntanglesinsideview.Basidia28.6–35.4
×9.3 –10.5μm,4-spored,narrowlyclavate,clamped.Lamel-
lae edgefertileorheterogeneous.Truecheilocystidia absent,
but terminal elements of tramal hyphae often well developed,
29.0–60.0×6.0–13.0μm,cylindrical,oftenseptate,colourless,
clamped.Pileipellis trichoderm at centre, a plagiotrichoderm
towards margin, composed of cylindrical to slightly inflated hy-
phae15–25μmwide,withbothpaleintracellularandincrusting
pigment and abundant clamp-connections.
 Habitat—Ondead woodofconiferoustreesor onsoilin
forestsandopenplaces,includinggrasslands.
 Knowndistribution—Northern,Western and EasternEu-
rope,Caucasus,WesternSiberia,RussianFarEast.
Fig. 8 Entoloma lampropus.a.Basidium;b.cheilocystidia;c.spores
(UPS: BOT:F-176490,neotype).―Scalebars=10μm.

156 Persoonia–Volume32,2014
Fig. 9 a, b: Entoloma lampropus.a.RM0855;b.LE254315.—c.E. placidum.JVG1060830-6.—d.E. sublaevisporumLIPJVG1070823T,holotype.―Scale
bars=1cm.—Photosby:a.S.Català;b.V.Kapitonov;c,d.J.Vila.
Additional specimens examined.austria, Frankenberg, Ried,Hinterzein-
ing,22 Sept1994, F. Sucti(WU13198,asE. dichroum); Rastenfeld,NW
Dobra,onwood,9Sept.2009,A. Hausknecht(WU24148,asE. dichroum);
Burgenland,Oberpullendorf,Tschnurdorf,Seltzabachtal, Pflanzen with
Salix and Alnus, 12 Oct. 1991, W. Klofoc & A. Hausknecht(WU 10092,
as E. dichroum)(WU10092,as E. dichroum).– Germany,Nettetal,3Oct.
2009, G. Wölfel (1509, L). – russia, Murmansk Region, Khibiny,Botani-
calGarden,onsoilin Picea-Betula forest, 4 Sept. 1974, L. Mikhailovsky
(LE9121,asLeptonia placida);VologdaRegion,KirillovDistrict,Kovarzino,
N59°44'28"E038°23'33",onsoilinPicea abiesforest,26Aug.2005,O. Ki-
rillova (LE235263); Bryansk Region, Syzemka District, ‘Bryansky Les’
NatureReserve, Chykhrai Village,onthe base ofFraxinus exelcior trunk
inbroad-leaved forest,24 Oct.2012,A. Fedosova (LE254339);Tatarstan
Republic,Zelenodolsk Region,on decayingwoodinPinus sylvestris-Tilia
cordataforest,27Sept.2010,K. Potapov(LE262991);UdmurtiaRepublic,
vicinitiesof Izhevsk, on rotten Picea stump, 19July 2009, V. Kapitonov
(LE254315);ibid.,onrottenPiceastumpinmixedforest,19July2009,V. Ka-
pitonov(LE254338); Karachaevo-CherkesiaRepublic,TeberdaBiosphere
Reserve,Arkhyz,onsoilinAbies nordmanniana-Fagus orientalisforest,19
Aug.2009,O. Morozova(LE254316);SverdlovskRegion,VisimskyNatura
Reserve,LipovySutukMt,N57°24'18"E059°43'34",onburntsoil,L. Marina,
3Sept.1999(LE258111);OrenburgRegion,TyulganDistrict,TashlaVillage,
onsoilinbroad-leavedforest,13July2007,O. Desyatova,(LE253584,as
E. tjallingiorum);Novosibirsk,Akademgorodok,mixedforest(Pinus sylvestris,
Acer negundo, Betula pendula)nearICGSB RASmodule, 11Aug.2011,
T. Bulyonkova(LE262992); TomskRegion, near airport, on Larix stump,
11Aug.2006,N. Agafonova(LE262985);Kamchatka Region,vicinities of
Esso,ondecaying wood,8Aug.2005,O. Morozova(LE254349).–sPain,
València,LaPuigmola,alt.350m,onCorylus avellana,amongmosses,16
Oct.2011,S. Català(RM0855).
 Notes—Entoloma lampropusisaratherconfusingspecies.
ItwasdescribedbyFriesasaspecieswith“pileosubcarnoso
convexo cinereo-griseo fibrilloso, lamellis albidis denticulo-
adnatis,stipitenitido coeruleo fistuloso” (Fries 1815). Inthe
sanctioningworkitwascharacterizedby “pileo demum um-
bilicatefibrillosogriseo,lamellisadnatisalbido-griseis,stipite
fistulosocoeruleo”(Fries1821).Thebrevityofthedescription
ledtovariousinterpretationsofthespecies.Intheconceptof
Lange (1937, pl. 76C), Bresadola (1929, pl. 570-1), Hesler
(1967)itrepresentsaspecieswithasmoothstipewithoutclamp
connections,andisconsideredamemberofsubg.Cyanula.
ThepresentworkfollowstheinterpretationofKühner&Romag-
nesi(1953),whichwasbasedonakeyphraseintheFries’s
description“primoobtutusimillimuspriori(Agaricus placidus)”,
stressing the resemblance to E. placidum.Thisinterpretation
wasadopted by Noordeloos (1982a,1992).The misapplied
interpretation of Agaricus lampropus mentioned above, was
described as a new species, Rhodophyllus sodalisKühner&
Romagn.,and lateronacceptedas E. sodaleKühner&Ro-
magn.exNoordel.Noordeloos(1982a)publisheda modern

157
O.V.Morozovaetal.:Entoloma subgenus Leptonia
description of E. lampropus, but nevertheless, due to the limited
numberofrecordsthespeciescontinuedtobeinsufficiently
knownandlotofmisidentificationswere still encountered in
severalherbariaduringourstudy.Thepresentstudyexpands
the concept of E. lampropus with more morphological, ecologi-
cal and molecular data, and its phylogenetic position is now
known,andfixedwithaneotype.
Entoloma lampropus belongs to the group of species character-
izedbythemany-anglednodulosesporesandthepresenceof
‘cheilocystidia’informofhyphalelements,oftenseptate,arising
fromthehymenophoraltrama.Thepredominantlygreybrown
pileus sometimes has a slight lilac tinge near the margin, and
thestipeisblueandlongitudinallyfibrillose,withoutsquamules
(contrarytoE. tjallingiorum). It grows bothterrestrialandon
rottenwood(mainlyconiferous).
4. Entoloma placidum(Fr.:Fr.)Noordel.,Persoonia11,2:150.
1981.—Fig.9c,10
Syn.: Agaricus placidusFr.,Observ.Mycol.2:94.1818;Agaricus placidusFr.:
Fr.,Syst.Mycol.1:202.1821;Leptonia placida(Fr.:Fr.)P.Kumm.,Führer
Pilzk.:96.1871;Rhodophyllus lampropus(Fr.:Fr.)Quél.,Enchir.Fung.:60.
1886;Entoloma placidum(Fr.)Zerova,inZerovetal.,ViznachnikUkraĭnĭ
5Basidiomycetes:104.1979[nom.inval.,Art.33.2].
Epitype(designated here): sweden, Småland,Femsjö,Hägnen, NW part,
onbeechstump,10Sept.1948,S. Lundell(5276)andG. Haglund(UPS:
BOT:F-121714,asLeptonia placida).
Pileus10–30mmbroad,conicaltoconvexwithstraightmargin
and slightly depressed or umbonate centre, not hygrophanous,
nottranslucently striate,entirelyradiallyfibrillose tominutely
squamulose, especially in the centre, with small dark grey-
brownsquamulesongreyishbackground.Lamellae moderately
distant, adnate or emarginate, with small decurrent tooth, whit-
ishtocreambecomingpink,withconcolorousedge.Stipe25 –
65×1–3mm,cylindricalorslightlybroadenedtowardsbase,
distinctlylongitudinallyfibrillose-striatewithwhitishorpaleblue
fibrilsondeepbluebackground,pruinoseinupperpart,base
withwhitetomentum.Contextwhitish,bluebeneaththestipe
surface.Smellandtastefarinaceous.Spores8.0–11.0(–11.5)
×6.0–7.0(–7.5)μm,Q=1.2–1.6,heterodiametrical,with6–8
bluntanglesinsideview.Basidia27.0–33.0× 8.8 –11.5μm,
4-spored,narrowlyclavate,clamped.Lamellae edge fertile or
heterogeneous. Scattered cystidia-like elements sometimes
present in the edge of the lamellae as vacuolised basidioles
orseptateterminalcellsof hyphaeofthe trama.Pileipellis a
trichoderm in centre, plagiotrichoderm towards margin, com-
posedofcylindricaltoslightlyinflatedhyphae10–25μmwide,
with intracellular pigment and abundant clamp-connections.
 Habitat—Ondeadwoodofdeciduoustrees,mostlyFagus,
but also Corylus, Betula,etc.
 Knowndistribution—WesternandEasternEurope,Cauca-
sus.
Specimens examined.russia,Karachaevo-CherkesiaRepublic,Teberda
BiosphereReserve, vicinities of Teberda,on beech trunk inAbies nord-
manniana-Fagus orientalisforest,7Aug.2009,O. Morozova (LE254335).
–sPain,Lleida,Bòscd’Aubàs,Bossòst,alt.1090m,onplantremnants,es-
pecially on small twigs of Corylus avellana and Betula pendula,30Aug.2006,
J. Carreras, J. Vila, F. Caballero, A. Duran & A. Mayoral(JVG1060830-6);
ibid.,2Sept.2006,J. Vila, F. Caballero & A. Mayoral(JVG1060902-2);ibid.,
26Aug.2008,F. Caballero(EFC2682008-151).–sweden,Småland(Inre),
Femsjö,Hägnen,NWpart,onbeechstump,30Aug.1949,S. Lundell(6020)
and J. Stordal(UPS:BOT:F-121715,neotype).
 Notes—Entoloma placidum is very similar to E. lampropus
duetothegrey-brownpileus,bluelongitudinallyfibrillosestipe
withoutsquamules, andmoreorlessnodulose spores.True
cheilocystidia are absent in both species, but in some speci-
mens cystidia-like elements can be observed as vacuolised
basidioles(Noordeloos1982a,b)orseptateterminalendings
ofthehyphaeofthetrama(Vila&Caballero2007).Entoloma
placidumcanberecognizedbythefarinaceoussmell,distinctly
nodulose spores, and habitat on the wood of deciduous trees,
especially Fagus.Foralongtimeitwasknownasaspecies
growingexclusivelyon beech wood. Recently somerecords
weremadeonotherdeciduoustrees(Corylus avellana, Betula
pendula)(Vila&Caballero2007).Moleculardataconfirmthat
they all belong to E. placidum.Entoloma lampropus grows on
conifersoronsoil.Entoloma tjallingiorum is also very similar but
differsbythedistinctlysquamulosestipeandwelldifferentiated
cheilocystidia.
5. Entoloma sublaevisporumVila,Noordel.&O.V.Morozova,
sp. nov.―MycobankMB803971;Fig.9d,11,12
Etymology. From latin ‘laevus’(smooth), referring to the very weakly
angled,subnodulosetoalmostsmoothspores.
Holotype.sPain,Ripollès,Girona,ValldeCarlat,Setcases,alt.1550m,
in acid soil, under Pinus uncinata, Sarothamnussp.andUrticasp.,23Aug.
2007,J. Vila & F. Caballero(LIPJVG1070823T).
Diagnosis.The species is characterized by the grey-brown pileus,bluish
finelylongitudinallystriatestipecombinedwiththemany-angled,nodulose
spores.
Pileus up to 20 mm broad, flattened or slightly convex, with a
shallowcentraldepression;greytopalegrey-brown,withdarker
centre, without violaceous tinges or only a hint in central depres-
sion;nothygrophanous,nottranslucentlystriate,withfineto
heavyfibrils,especiallyintheapex,whereitissubsquamulose;
marginstraighttorevolute,protrudingabovethelamellae.La-
mellaemoderatelydistant(L=15– 20)withabundantlamellulae
(1:3to1:5),adnatetoemarginate,thin,slightlyventricoseor
not;whitish,turningpalepinkishwhensporesmature;edgeof
thesamecolour,entireorsomewhatirregular.Stipe central, up
to 40 ×2mm,cylindrical,straightorslightlycurved;darkblueto
bluishgrey;surfacesmoothorwithweakfibrils,finelylongitudi-
nallystriate;pruinoseatapexandwithwhitebasaltomentum.
Flesh very thin, greyish on the pileus and grey-bluish grey on
thestipe;smellfungoid.Spores7.7– 9.3×4.8–5.9µm,aver-
age8.5×5.3µm,Q=1.4–1.8,Qm=1.6,heterodiametrical,
withveryweakangles,subnodulosetoalmostsmooth.Basidia
27.6–32.8×9.7–11.7µm,4-spored,rarely2-spored,narrowly
clavate, clamped. Lamellae edge fertile or heterogeneous.
Cheilocystidia absent in holotype, sometimes present, in-
termixedwith basidia, 23.2–45.0 × 5.0 –7.8, subcylindrical,
flexuose to lageniform, sometimes with long tapering neck,
frequentlyseptate.Hymenialtramacyanophilous.Pileipellis a
trichodermstructurecomposedofcylindricalhyphae,20–28µm
wide,withfusiformterminalelements;pigmentmixed,brownish
Fig. 10 Entoloma placidum.a.Basidium;b.spores(UPS:BOT:F-121714,
epitype).―Scalebars=10μm.

158 Persoonia–Volume32,2014
differs from E. chytrophilum particularly by the greyish brown
colourof the pileus and smallerspores. The many-angled,
nodulose spores distinguish it from the other species with
grey-brownpileus(E. lampropus, E. placidum, E. tjallingiorum).
CheilocystidiawerefoundonlyinthespecimenfromAustria.
The p-distance between E. chytrophilum and E. sublaevisporum
is7.4%.
6. Entoloma tjallingiorum Noordel.,Persoonia11:465.1982
Syn.:Leptonia tjallingiorum(Noordel.)P.D.Orton,Mycologist5,3:135.1991.
Misappliednames: Agaricus dichrous sensu Fr.,Summa Veg.Scand. 2:
287.1849; Agaricus dichrous sensuFr.,Ic.Sel.Hymenomyc. 1, pl.92.
1867;Entoloma dichroumsensuBres.,Iconogr.Mycol.12,pl.554.1929;
Entoloma dichroumsensuKonrad&Maubl.,Icon.Sel.Fung.2,pl.190,
f. 2. 1932; Rhodophyllus dichrous sensu J.E. Lange, Fl.Agar. Dan. 2,
pl.72A.1937;Rhodophyllus dichrous sensuRomagn.,Bull.Soc.Mycol.
France92:292.1976;Agaricus placidussensuFr.,Ic.Sel.Hymenomyc.
1,pl.97,f.1.1867.
a. var.tjallingiorum—Fig.13a‒ c,14
Pileus20–80mmbroad,hemisphericalexpandingtoconico-
convex or applanate, with low broad umbo, sometimes de-
pressed, with involute then straight margin, not hygrophanous,
nottranslucentlystriate,entirelyradiallyfibrillosetosquamulose
in centre, light to moderately dark grey-brown, often with vio-
laceousbluetingesnearpileusmargin.Lamellae adnate with
small decurrent tooth or arcuate, whitish to cream in youth
becomingpink,withirregularconcolorousedge.Stipe25–100
×5–10mm,cylindricalorbroadenedtowardsbase,fibrillose,
entirelysquamulosewithdarkbluesquamules,moreintensively
coloured in upper part, greyish blue or violaceous-blue below,
base with white tomentum. Flesh whitish, blue beneath the
stipesurface.Smell indistinct, tastemildorbitterish.Spores
8.0–11.0(–12.0)×5.8–7.2(–7.6)μm,Q=1.3–1.5,heterodia-
metrical, with 6 –9 rather blunt angles in side view. Basidia
32.6–40.4×8.6–10.3μm,4-spored,narrowlyclavate,clamped.
Lamellae edgeheterogeneousor,rarely,sterile.Cheilocystidia
29.7–71.1×5.9–8.7μm,cylindrical,narrowlylageniformorir-
regularly shaped, sometimes septate, sometimes represented
by strands of terminal elements of hyphae of the hymenial
trama,colourless.Pileipellis a plagiotrichoderm of cylindrical to
slightlyinflatedhyphae7–20μmwide,withbothbrownintracel-
lular and incrusting pigment and abundant clamp-connections.
Caulocystidiapresentaslong,upto250μm,septate,clamped
hairs,withcylindricalorlageniformterminalelements,60–120
×8–12μm,colourlessorwithincrustingandblueintracellular
pigment.
 Habitat—Onsoilorondeadwoodofmostlybroad-leaved
trees(Quercusspp.),butalsoonBetula and Alnus in deciduous
andmixedforests.
 Knowndistribution—WesternandEasternEurope,Western
Siberia.
Specimens examined.russia,LeningradRegion,Sovkhozy,Sept.1936,
around trunks in Alnus forest, R. Singer(LE9123, as Leptonia placida);
NovgorodRegion, ‘Valdajsky’NationalPark,ValdajDistrict, Poddub’e,22
Sept.2011,onsoilinbroad-leaved-coniferous(Picea abies)forest,O. Mo-
rozova(LE254318);MoscowRegion,TaldomDistrict,BolshoyeStrashevo,27
Sept.2010,E. Lukashina(LE254320);TatarstanRepublic,Kazan’,Sovetsky
District,‘SkotskiyeGory’Park,onsoilinbroad-leavedforest(Tilia cordata,
Quercus robur, Betula pendula),29Sept.2011,K. Potapov (LE254317);
UlyanovskRegion,onsoilinbroad-leavedforest,14Aug.2009,E. Ilyukhin
(LE254319).–sPain,Barcelona,CanRomegosa,Sant FostdeCampsen-
telles,alt.140m,ontrunkofQuercus pubescens,5Oct.2008,F. Caballero
(SFC081005-01);ibid.,F. Caballero(SFC081019-01);ibid.,16Nov.2008,
J. Vila & F. Caballero (JVG 1081116-5).– sweden, Uppland, Bondkyrka,
Predikstolen, on soil and rotten wood of Quercus,4Oct.1980,S. Ryman
(6124)(UPS:BOT:F-016378,holotype).
Fig. 11 Entoloma sublaevisporum.a.Basidium;b.spores(LIPJVG
1070823T,holotype).―Scalebars=10μm.
vacuolar pigment abundant, also encrusting epiparietal present,
especiallyonthinhyphae.Clamp-connections very abundant
inhymeniumstructures,scarceonthehyphaeofthepileipellis.
 Habitat—Onsoil,inpine(Pinus uncinata)forest.
 Knowndistribution—WesternEurope(Austria,Spain(holo-
type)).
Additional material examined.austria, Kärnten,Eisenkappel,Vellacher
Kotscha,7Sept.1998,A. Hausknecht(MEN9858).
 Notes—Accordingtothephylogeneticanalysisthenewspe-
cies is close to E. chytrophilum. Morphologicallythissimilarityis
confirmedbythesporemorphology.Entoloma sublaevisporum
Fig. 12 Entoloma sublaevisporum.a.Basidium;b.cheilocystidia;c.spores
(MEN9858).―Scalebars=10μm.

159
O.V.Morozovaetal.:Entoloma subgenus Leptonia
Fig. 13a– c:Entoloma tjallingiorumvar.tjallingiorum.a.LE254317;b.JVG1081116-5;c.LE227507.—d,e.E. tjallingiorumvar.alnetorumLE254321.—
f, g: E. tjallingiorumvar.laricinum.f.LE254343,holotype;g.LE254344.―Scalebars=1cm.—Photosby:a.K.Potapov;b.J.Vila;c.A.Kovalenko;d,e,
g.O.Morozova.

160 Persoonia–Volume32,2014
Additional specimens examined (grouped together with the E. alnetorum
holotype). russia,SamaraRegion,ZhigulevskyNatureReserve,vicinitiesof
BakhilovaPolyana,MaloyeKamennoyePole,Tilia cordata-Acer platanoides
forest,16Aug.2004,O. Morozova(LE227507);ibid.,17Sept.2004,E. Maly-
sheva(LE234285);ibid.,vicinitiesofBakhilovaPolyana,Tilia cordata-Acer
platanoidesforest,1 Sept. 2004, E. Malysheva (LE227584, as Entoloma
placidum).
 Notes—Entoloma tjallingiorum belongs to the group of spe-
ciescharacterizedbythemany-angledalmostnodulosespores
and septate terminal elements of the hymenophoral trama that
protrudethrough thehymenium(‘cheilocystidia’).Amongthe
species with greyish brown pileus and blue stipe it stands out
bythestouterbasidiocarpsanddistinctlysquamulosestipe.
Inanearlierpaper(Noordeloos1982a,1992)somecollections
with a bluish tinge in the lamellae and pigmented cheilocystidia
were also assigned to E. tjallingiorum.Moleculardatashowthat
these specimens are discolored forms of E. euchroum.The
current concept of E. tjallingiorum therefore excludes forms
withbluetingesinthelamellae.
As was shown by the phylogenetic analysis, the evolutionary pro-
cess within this species continues at the present time: several
lineagesintheearlystageofdivergencewererevealed.Asa
result, based on molecular and morphological differences, as
well as geographical distribution, two more varieties of E. tjallin-
giorumcanbedistinguished.Butthelimitsbetweenthesevarie-
ties are sometimes vague and intermediate forms have been
found.So,thespecimensfromZhiguli(LE227507,LE227584,
LE234285)despitethegeneticaffinitywithvar.alnetorum are
morphologicallyclosertothetypevariety.Thespecimenfrom
Novgorod(LE254318)possessesinsomelamellaeratherlarge
sitesofsterileedgewhicharecharacteristicforvar.laricinum.
b. var. alnetorum(Monthoux&Röllin)O.V.Morozova,Noordel.
&Vila,comb. nov.―MycobankMB803972;Fig.13d,e,15
Basionym.Entoloma alnetorumMonthoux&Röllin,Mycol.Helv.3,1:43.
1988.
Pileus20 – 40 mmbroad,hemisphericalexpandingto plano-
convexwithincurvedfimbriatemargin,nothygrophanous,not
translucently striate, entirely tomentose when young, becoming
radiallyfibrillosetosquamuloseatcentre,creamtopaleochra-
ceousatfirst,thensordidochre.Lamellae adnate with small
decurrent tooth or arcuate, whitish to cream in youth becoming
pink,withirregularconcolorousedge.Stipe25– 80×5–10mm,
cylindrical,broadenedtowardsbase,fibrillose-striate,entirely
squamulosewithdarksquamules,violaceousinupperpartand
whitishbelow,basewithwhitetomentum.Fleshwhitish,blue
beneaththestipesurface.Smell fruitish, taste mild or slightly
spermatic.Spores8.1–11.3×5.5–7.2μm,Q=1.4–1.8(–1.9),
heterodiametrical,with6 – 9 rather blunt anglesinsideview.
Basidia37.0 – 39.6×11.7–13.0μm,2-or4-spored,narrowly
clavate, clamped. Lamellae edge sterile or heterogeneous.
Cheilocystidia39.8–69.1 × 6.6 –15 μm,cylindrical, narrowly
lageniform or irregularly shaped, sometimes septate, colour-
less.Pileipellis a plagiotrichoderm of cylindrical to slightly in-
flatedhyphae15– 25μmwide,withbothpaleintracellularand
Fig. 14 Entoloma tjallingiorumvar.tjallingiorum.a.Basidium;b.cheilocys-
tidia;c.spores(LE254318).―Scalebars=10μm.
Fig. 15 Entoloma tjallingiorumvar.alnetorum.a.Basidium;b.cheilocystidia;
c.spores(LE254321).―Scalebars=10μm.

161
O.V.Morozovaetal.:Entoloma subgenus Leptonia
incrusting pigment and abundant clamp-connections.Caulo -
cystidiapresentaslong,upto250μm,septateclampedhairs,
with lageniform, cylindrical or subcapitate terminal elements
30–90×6–12μm,colourlessorwithblueorviolaceousintra-
cellularpigment.
 Habitat—OndeadwoodofAlnus incana in deciduous fo-
restsinMay–June,rarelyalsoinJulyorAugust.
 Knowndistribution—WesternandEasternEurope,Western
Siberia.
Specimens examined. russia,LeningradRegion,KirovskDistrict,Vasil-
kovo,leftbankofLavaRiver,ondeadtrunkofAlnus incana in Alnus incana-
Ulmus glabraforest,11June2009,E. Popov(LE254321);SamaraRegion,
ZhigulevskyNatureReserve,vicinitiesofBakhilovaPolyana,onsoilinAlnus
glutinosafloodplainforest,15June2004,E. Malysheva(LE234287);ibid.,
on soil in Acer platanoides-Tilia cordataforest,18Aug.2004,E. Malysheva
(LE227527);TumenRegion,Surgutdistrict,YuganskyNatureReserve,Ka-
menny, on soil in Pinus sibirica-Abies sibiricaforest,9July2006,E. Zvyagina
(LE254322).–switzerland, Nant-Bride, Sixt, on old trunks and branches of
Alnus incana in submontane Alnus forest,alt.±850m,28June1986,O. Röllin
(G00111402,holotype);ibid.,29May1994,O. Röllin (L).
 Notes—Entoloma alnetorum has been described as a spe-
cies very similar to E. tjallingiorum due to the thin-walled, al-
most nodulose spores, however it is distinguished by the pale
ochraceus pileus and the vernal appearance in Alnus forests
(Monthoux&Röllin1988).Phylogeneticanalysisshowsthatall
specimenspossessingthesefeaturesaregroupedtogether.At
the same time the difference between them and typical E. tjallin-
giorumisverysmall(p-distancebetweenholotypesis1.3%).
Somespecimens(LE227507,LE227584,LE234285)withthe
typical habit of E. tjallingiorum occur in the E. alnetorum-clade
withthedifferenceonlyin0.2%.Forthisreasonwedecided
to consider E. alnetorum as a variety of E. tjallingiorum.It is
noteworthy that in E. dichroum also specimens with a pale
pileuscan be encountered (JVG 1070821-4).They can be
separated from the species of the tjallingiorum-group by the
characteristicsporeswithsharpangles.
c. var. laricinumO.V.Morozova,Noordel.,Vila&E.S.Popov,
var. nov.―MycobankMB803973;Fig.13f,g,16
Etymology.Thenamereferstothesubstrateonwhichithasbeenfound
(Larix cajanderi).
Holotype.russia,KamchatkaRegion,vicinitiesofEsso,ondeadwood
of Larix cajanderi,8Aug.2005,E. Popov & O. Morozova(LE254343).
Diagnosis.Entoloma tjallingiorumvar.laricinum differs from the type variety
bythesterilelamellaeedgewithlongseptate‘cheilocystidia’(terminalele-
mentsofthehymenophoraltrama)aswellasbygrowingonLarixwood.
Pileus20–60mmbroad,hemisphericalexpandingtoconico-
convex or applanate, with low broad umbo, with involute then
straight margin, not hygrophanous, not translucently striate,
entirelyradiallyfibrillosewithlightgrey-brownfibrilsandsqua-
mules on whitish background, with violaceous blue tinge near
pileusmargin.Lamellae adnate with small decurrent tooth or
arcuate, whitish to cream in youth becoming pink, with irregu-
larconcolorousedge.Stipe25–80×5 –15mm,cylindricalor
broadenedtowardsbaseupto20mm,fibrillose,entirelysqua-
mulosewithdarkviolaceous-blueorviolaceous-greysquamules
onpaler,almostwhitebackground,basewithwhitetomentum.
Fleshwhitish,bluebeneaththestipesurface.Smell indistinct,
tastemild.Spores8.5–11.3×5.5–7.4μm,Q=1.4–1.7,hetero-
diametrical,with6–9ratherbluntanglesinsideview.Basidia
35.4–42.2× 10.3 –13.1μm,4-spored,clavate, clamped.La-
mellae edgesterile.Cheilocystidia29.7–71.1 × 5.9 – 8.7 μm,
cylindrical, flexuous, septate, represented by strands of terminal
elementsofhyphaeofthehymenialtrama,colourless.Pileipellis
aplagiotrichodermofcylindricaltoslightlyinflatedhyphae7–20
μmwide,withbothbrownintracellularandincrustingpigments
and abundant clamp-connections.Caulocystidia present as
long,upto350μm,septateclampedhairs,withlageniformor
cylindricalterminalelements50 –120 × 6 –12 μm, colourless
orwithincrustingandblueintracellularpigment.
 Habitat—Onrottenwood of Larix in mixed L. cajanderi-
Betula ermaniiforest.
 Knowndistribution—RussianFarEast.
Additional specimen examined.russia,KamchatkaRegion,vicinitiesof
Esso, on burnt wood of Larix cajanderi,8Aug.2005,E. Popov & O. Morozova
(LE254344).
 Notes—AlthoughE. tjallingiorumvar.laricinum can be dis-
tinguished from the typical variety by the sterile lamella edge
with abundant, septate terminal elements of the hymenophoral
trama, this character is not reliable enough to make a clear-cut
morphological distinction. Sometimes we find this character
alsointhetypicalvariety.Themaindistinctivefeaturesarethe
habitat(growingonLarix cajanderi)andthegeographicorigin,
theratherisolatedKamchatkaPeninsula.Thismayexplainthe
significantgeneticdivergence(p-distance3.3%).
Section Dichroi O.V.Morozova,Noordel.&Vila,sect. nov.―
MycobankMB803974
Type species.Entoloma dichroum(Pers.:Fr.)P.Kumm.
Habitcollybioidortricholomatoid;pileusconico-convexorplano-
convex;lamellaealmostfreetoemarginate;stipe flocculose to
Fig. 16 Entoloma tjallingiorum var.laricinum.a.Basidium;b.cheilocystidia;
c.spores(LE254343,holotype).―Scalebars=10μm.

162 Persoonia–Volume32,2014
Fig. 17a– c:Entoloma dichroum.a.LE227472(phototakenattheneotypelocality);b.herbariumSGC;c.JVG1070821- 4.—d.E. eugeneiLE253771,
holotype.—e,f.E. allochroumLE254324.―Scalebars=1cm.—Photosby:a,d.O.Morozova;b.S.Català;c.J.Vila;e,f.K.Potapov.
squamulose;lamellae edgesterileorheterogeneous;cheilo-
cystidia present, spores with sharp angles, pileipellis more or
lesstrichodermofinflatedelementswithintracellularpigment;
brilliantgranulesabsent;clamp-connections present and often
frequent;terrestrial,inforests.
7. Entoloma dichroum(Pers.:Fr.)P.Kumm.,FührerPilzk.:
97.1871.—Fig.17a–c,18
Syn.:Agaricus dichrousPers.,Syn.Meth.Fung.(Göttingen)2:343.1801;
Rhodophyllus dichrous(Pers.:Fr.)Quél.,Enchir.Fung.(Paris):58.1886;
Leptonia dichroa(Pers.)P.D.Orton,Mycologist5,3:132.1991.
Excl.:Agaricus dichroussensuFries1849,1867;Entoloma dichroum sensu
Bresadola1929;sensuKonrad&Maublanc1932;Rhodophyllus dichrous
sensuLange1937;sensuKühner&Romagnesi1953(=E. tjallingiorum).
Neotype (designated here).russia,SamaraRegion,ZhigulevskyNature
Reserve,vicinitiesofBakhilovaPolyana,MalayaBakhilovaHill,broad-leaved
forest,26Aug.2003,E. Malysheva(LE227472).
Pileus5 – 35 mm broad,conicalorhemispherical,hardly ex-
panding with age, with involute then straight margin, not hy-
grophanous, not translucently striate, dark violaceous-blue,
purplish brown, or very pale, greyish with lilac tinge, entirely
granular-fibrillose,becomingsquamulosewithviolaceous-blue
squamulesonpalebackground.Lamellae adnate-emarginate
with decurrent tooth, white then pinkish, with entire concolor-
ousedge.Stipe30–80×2 – 5mm,clavateorcylindricalwith
slightly swollen base, deep blue, different from colour of pileus,
longitudinallyfibrillose,withdarkbluesquamulesonthepaler
background,basewithwhitetomentum.Fleshwhite,darkblue
underthesurface.Smellindistinct, tasteunpleasant.Spores
9.2–11.5×6.4–7.7μm, Q=1.3 –1.7,heterodiametrical,with
5 –7 pronounced angles in side view.Basidia 44.2 – 51.4 ×
8.5–10.5μm,clavate,clamped.Lamellae edge heterogene-
ous.Cheilocystidia 20 – 45 × 5 –11μm, cylindrical, narrowly
lageniform or irregularly shaped, sometimes septate, colourless,
scatteredamongbasidia.Pileipellis a trichoderm of cylindrical
hyphaewithterminalelements35–120×15–35μmwithblue
intracellular pigment and abundant clamp-connections.Caulo-
cystidiapresentaslong,upto300μm,septateclampedhairs,
withtaperingterminalelements,50–110×8–12μm,withdark
blueintracellularpigment.
 Habitat—Onsoilindeciduousforests.
 Knowndistribution—WesternandEasternEurope.
Additional specimens examined.russia, Samara Region, Zhigulevsky
NatureReserve,vicinitiesofBakhilovaPolyana,MaloyeKamennoyePole,
Tilia cordataforest,3July2005,E. Malysheva(LE234260).–sPain,Girona,

163
O.V.Morozovaetal.:Entoloma subgenus Leptonia
TorrentBurgil,Pardines,alt.1425m,inacidsoil,underBuxus sempervirens,
21Aug.2007,J. Vila & F. Caballero(JVG1070821-4);València,LaPuigmola,
alt.350m,inbasicsoil,underPinus halepensis, amongmosses, 16Oct.
2011, S. Català(herbariumSGC).
 Notes—Entoloma dichroum together with E. eugenei forms
aseparatecladegeneticallycharacterizedbythelargeinser-
tionin the ITS1-region.Morphologically,E. dichroum can be
recognizedby the bright blue squamulosestipe and spores
with5–7 sharp angles.The pileus colour,however,varies
considerably among the studied collections, from bright blue
to violaceous-blue, violaceous-brown and pale brown. The
ITS-sequencesslightlyvary,howeverthisvariability(p-distance
1.4–2%base-pairdifference)mightwellbeacceptablewithin
aspecies.Morematerialwouldpossiblyallowforthedistinc-
tionofvarieties.Entoloma allochroum, another species with
sharply-angled spores possesses a lilaceous or violaceous,
lesssquamulose,morelongitudinallyfibrillosestipe.Entoloma
dichroum differs from the closely related E. eugenei mainly by
the slender collybioid habit, the heterogeneous lamellae edge,
andslightlysmallerandlesspronouncedlyangledspores.
8. Entoloma eugeneiNoordel.&О.V.Morozova,Mycotaxon
112:234.2010.—Fig.17d,19
Pileus13 – 60 mmbroad,hemisphericalexpandingto plano-
convex with incurved margin, fleshy, not hygrophanous, not
translucently striate, entirely velvety when young, becoming
glabrousatthemargin,uniformlydeepblue(Indianblue)atfirst,
thenwithviolettingeatmargin.Lamellae adnate-emarginate
with decurrent tooth, pure white then pinkish, with irregular
concolorousedge.Stipe30–80×4–8mm,clavateorcylindrical
withswollenbase(to15mm),concolourouswiththepileusor
slightlypaler,entirelysquamulosewithconcoloroussquamules,
basewhite tomentouse. Flesh white, darkblue beneath the
surface.Smellslightly spicy,tastemild. Spores 10.0 –12.5 ×
6.0–8.0μm,Q=1.3 –1.7,heterodiametrical,with5–7angles
insideview.Basidia34–44×9–12μm,clavate,clamped.La-
mellae edgesterile.Cheilocystidia28.0–39.0×6.5–15.5μm,
cylindrical, narrowly lageniform or irregularly shaped, colour-
less.Pileipellis a trichoderm of cylindrical hyphae with terminal
elements90–200×12–20μmwithblueintracellularpigment
and abundant clamp-connections.Caulocystidia present as
long,upto250μm,septateclampedhairs, with tapering or
cylindricalterminal elements 50–100 × 8 –12 μm, with dark
blueintracellularpigment.
 Habitat—Onsoilinthefloodplainforest.
 Knowndistribution—Russian FarEast,Japan(GenBank
AB509605,asEntolomaaff.kujuense).
Specimens examined.russia, Primorsky Territory, Kedrovaya Pad Nature
Reserve,therightbankoftheKedrovayaRiver,N43°05'51"E131°33'34",24
Aug.2005,E. Popov(LE253771,holotype);ibid., 8Sept.1994,E. Bulach
(VLAM-3556);ibid.,LeopardovySanctuary,watershedoftheriversGryaz-
nayaandAnanjevka,onthebaseofdeadtrunk,1Sept.2011,A. Kovalenko
(LE254340);ibid.,1Sept.2011,T. Svetasheva(LE254347).
 Notes—Entoloma eugenei is morphologically very close
to E. dichroum.Themainmorphologicaldifferenceisinitstri-
Fig. 18 Entoloma dichroum.a.Basidium;b.cheilocystidia;c.spores
(LE227472,neotype).―Scalebars=10μm.
Fig. 19 Entoloma eugenei.a.Basidium;b.cheilocystidia;c.spores
(LE253771,holotype).―Scalebars=10μm.

164 Persoonia–Volume32,2014
cholomatoid habit, the sterile lamellae edge, and slightly larger
andmorepronouncedlyangledspores. Genetically it differs
from E. dichroumamongotherthingsinoneratherlarge(about
40base-pair)insertionintheITS1-region.Thesignificantdiver-
gencebetweenthesetwospecies(p-distance9.8%base-pair
difference)couldbeexplainedbygeographicalreasons–the
natural isolated habitat of E. eugenei in the Southern Far East
andJapanwithuniqueclimaticconditions(Noordeloos&Mo-
rozova2010),whileE. dichroumisknownfromEurope.
INCERTAE SEDIS
9. Entoloma allochroum Noordel., Persoonia11,4:463.
1982.—Fig.17e,f,20
Pileus20–40 mm diam, conical, thenconvex with umbo,
expanded to plano-convex, not or hardly hygrophanous, not
translucently striate or at margin only, with straight margin,
firstwithgreyishbrownishlilaceustingedvelvetycoveringlater
breakingupintobrownishlilacsquamulesonpalebackground.
Lamellae moderately crowded, adnate-emarginate, adnexed to
almost free, ventricose, whitish, becoming pinkish, with entire to
irregularconcolorousedge.Stipe40–70×3– 6mm,cylindrical,
broadened towards the base, pale violaceous, entirely covered
withdarkerfibrillosesquamules,withwhitetomentumatbase.
Context greyish, darker under the surface, yellowish at the stem
base.Smell agreeable or indistinct, taste indistinct. Spores
8.3–12.5×6.2 – 9.5μm,Q=1.3–1.6,heterodiametrical,with
5–8ratherpronouncedangles.Basidia32.6– 44.2×11.9–15.7
μm,4-spored,clavate,clamped.Lamellae edgeheterogeneous.
Cheilocystidia32.4–57.4×5.5–15.7μm,cylindricaltoflexuose,
septate,rareor absent. Pileipellis a trichoderm of cylindrical
toinflatedhyphaewithterminalelements15– 40μmwidewith
brownish violaceous intracellular pigment. Some hyphae of
pileitramaincrusted.Clamp-connectionspresent.
 Habitat—Onsoilandplantremainsinbroad-leavedforests
andparks.
 Knowndistribution—WesternEurope,Caucasus.
Specimens examined.austria,VellacherKotscha,Eisenkappel,Karinthia,
7Sept.1998,K.F. Reinwald & A. Hausknecht(L9860).–Germany,Mühltal
bei Willisau, im feuchten Eschen-haselnusswald zwischen Moosen und
Schachtelhalmenauffeuchter,lehmigerErdeinderNähedesBachufers,21
Sept.1984,G. Wölfel(L:E4884,asE. dichroum(Noordeloos2004:1276,
upperfig.).–netherlands,Aerdenhout,29July1973,Kits van Waveren(holo-
typeL);Apeldoorn,Vellertsdijk,7Aug.1993,L. Bos(L,asE. tjallingiorum);
Paterswolde,Vennebroek,14Aug.2000,R. Chrispijn(L).–russia, Caucasus
BiosphereReserve,valleyofthePslukhRiver,onsoilinbroad-leavedforest
with Fagus orientalis, Alnus glutinosa, Abies nordmanniana,29Aug.2006,
A. Kiyashko (LE262984); Karachaevo-Cherkesia Republik, Teberda Bio-
sphereReserve, Dombaj, onrottenstump in Abies nordmanniana-Fagus
orientalisforest,14Aug.2012,T. Svetasheva(LE254342);ibid.,Teberda,on
rotten stump in Fagus orientalisforest,19Aug.2012,K. Potapov(LE254324).
–sPain,Vald’Aran,Lleida, alt. 1090m,on plantdebrisin forestofAlnus
glutinosa, Populussp., Fraxinus excelsior,2Sept.2006,J. Vila, F. Cabal-
lero, A. Mayoral(JVG1060902-1);ibid.,24Aug.2008,F. Caballero(EFC
2482008-148);Espinavell,Girona,alt.1350 m,inacidsoil,underCorylus
avellana,12July2008,F. Caballero(EFC1272008-137).– switzerland,
Schonau,Tunau,20Sept.2009,G. Wölfel (L,E0809).
 Notes—Entoloma allochroumisaneasilyrecognizablespe-
cies due to the presence of the lilaceous or violaceous colours
both in the pileus and, especially, in the stipe, white lamellae,
aswellasratherthick-walledandpronouncedlyangledspores.
Due to the sharply-angled spores E. allochroum is similar to
E. dichroum and E. eugenei, however the molecular evidence
doesnotallowtheplacingofthisspeciesinsect.Dichroi (Fig.2).
10. Entoloma callichroum E.Horak&Noordel.,inNoordeloos,
Cryptog.Mycol.4,1:33.1983
a. var.callichroum—Fig.21
Pileus to 22 mm broad, convex with small papilla, not hygropha-
nous,nottranslucentlystriate,radiallyfibrillose,lilaceous-pink.
Lamellae distant, emarginate, ventricose, whitish with lilac tint
towardsentireconcolorousedge.Stipe 40 × 2 mm, cylindrical,
steel-blue,basewithwhitetomentum,fistulose.Smell and taste
notreported. Spores 9.5–13.2 × 7.2 – 9.4 μm, Q=1.4–1.6,
heterodiametrical,with6–9bluntanglesinsideview,almost
nodulose. Basidia 39.0 –48.9 × 14.4 – 20.1 μm, 4-spored,
broadlyclavate,clamped.Lamellae edgefertile.Cheilocystidia
absent.Pileipellis a plagiotrichoderm of cylindrical to slightly
inflatedhyphae 10 –18 μm wide, with intracellular pigment.
Clamp-connectionspresent.
 Habitat—OnsoilinAlnus incanaforest.
 Knowndistribution—WesternEurope.
Specimen examined.switzerland,Graubunden,Forna,3Aug.1971,E.
Horak(ZT71/ 58,holotype).
 Notes—Thisrarespeciesischaracterizedbypinkishvio-
laceoustingeinthepileusincombinationwithsteelbluestipe.
The type variety is distinguished by the broad, almost nodulose
spores,andtheabsenceofcheilocystidia.
Fig. 20 Entoloma allochroum.a.Basidium;b.cheilocystidia;c.spores
(LE262984).―Scalebars=10μm.

165
O.V.Morozovaetal.:Entoloma subgenus Leptonia
b.var.venustum(Wölfel&F.Hampe)O.V.Morozova,Noordel.
&Vila,comb. nov.―MycobankMB804535;Fig.22,23a,b
Basionym.Entoloma venustumWölfel&F.Hampe,Z.Mykol.77/2:185.
2011.
Pileus0.5–30mmbroad,convextoplano-convex,nothygro-
phanous, not translucently striate, radially silky fibrillose to
squamuloseincentre,pinkishlilaceoustoviolaceous,becoming
brownishpinkwithage.Lamellae moderately distant, adnate-
emarginate with small decurrent tooth, ventricose, brightly pink-
ish lilaceous, pinkish violaceus or whitish with blue tinge towards
entireconcolorousedge,becomingpink.Stipe20–50×0.3– 2.5
mm,cylindrical,longitudinallyfibrillose-striate,darkblue,steel
blueorviolaceous-blue,basewithwhitetomentum,fistulose.
Smelloffruitsorflowers(viola),tasteunknown.Spores11.5–
13.0(–16.0)×5.7–8.6μm,Q=1.3–1.8(–2.5),heterodiametri-
cal,with6– 8moderatelypronouncedanglesinsideview.Basi-
dia35.0–45.0×12.0–18.0μm,4-spored,broadlyclavate,clamp-
ed.Lamellae edge heterogeneous. Cheilocystidia 30 – 60 ×
15–28μm,broadlyclavateorsphaeropedunculate,intermixed
with the basidia, sometimes hardly separated from the ba-
sidioles.Pileipellis a plagiotrichoderm of cylindrical to slightly
inflatedhyphae 10 – 20 μm wide, withintracellular pigment.
Clamp-connectionspresent.
 Habitat—Onsoilingrasslandsandinwetdeciduousforest.
 Knowndistribution—Western Europe, WesternSiberia,
RussianFarEast.
Specimens examined.Belarus,VitebskRegion,VerkhnedvinskDistrict,
vicinitiesof Rositsa Village,onsoilin Alnus incana forest,19Aug. 2003,
P. Kolmakov(LE226909,asE. lepidissimum).–Germany,Hannover-Nord,
Kuglfanger,13Nov.2010,G. Wölfel & F. Hampe(WöE17/10,L,asE. ve-
nustum,holotype).–russia,SamaraRegion,ZhigulevskyNatureReserve,
Bakhilova Polyana, on soil in Betula pendulaforest,23Aug.2003,E. Maly-
sheva(LE227532, as E. lepidissimum);Altaj Republic,Altajsky Nature
Reserve,Komga,onsoilinfloodplain forest,18Aug.2008,V. Malysheva
(LE254312);Novosibirsk,Akademgorodok,onrottenbirchstump,15June
2008,T. Bulyonkova(LE254313);ibid.,plantedforestbetweentheSobolev
InstituteofMathematicsandtheComputingCenter,16Aug.2011,T. Bulyon-
kova(LE254314);PrimorskyTerritory,UssuriyskyNatureReserve,17Sept.
1963,M. Nazarova(VLAM-20528,asRhodophyllus lampropus).
 Notes—ThephylogeneticanalysisshowsthatE. venustum
is very close to E. callichroum(p-distance1.8%base-pairdif-
ference)and,therefore,couldbeconsidereditsvariety.Both
species are morphologically distinct by the pinkish violaceous
pileus;moreorlesslilaceous-bluetingesinthelamellae,the
steelblueorviolaceous-bluestipe,andthesizeofthespores.
The description of E. venustum as a new species was based on
the bright colour of the basidiomata and on the presence of well
developed cheilocystidia, which, however, do not form a sterile
gill edge and are often hardly distinguishable from basidioles
(Wölfel& Hampe 2011).These characterscan significantly
varywithintherangeofgenetically(nrITS)identicalspecimens.
A more reliable feature for delimitation of these two taxa is spore
form.SporesofE. venustum are narrower and possess more
pronouncedangles.Alsothepresenceofanumberofextremely
long(upto16μm)germinating(?)sporeshasbeenreported
fromtheholotypeandotherspecimens(Table3).
Fig. 21 Entoloma callichroum var.callichroum.a.Spores;b.basidia(ZT
71/58,holotype).―Scalebars=10μm.
Fig. 22 Entoloma callichroum var. venustum.a.Basidium;b.cheilocystidia;
c.spores(WöE17/10, holotype).―Scalebars=10μm.

166 Persoonia–Volume32,2014
Fig. 23 a, b: Entoloma callichroum var. venustum.a. WöE17/10, holotype;b.LE254312. — c. E. coelestinum LE258103. — d, e: E. percoelestinum.
d.LE254390(JVG1130925-24),holotype;e.LE254341.—f,g:E. lepidissimumf.LE254871;g.LE234751.―Scalebars=1cm.—Photosby:a.F.Hampe
(fromWölfel&Hampe2011);b,f,g.O.Morozova;c.L.Marina;d.J.Vila;e.T.Bulyonkova

167
O.V.Morozovaetal.:Entoloma subgenus Leptonia
11. Entoloma coelestinum(Fr.)Hesler,Beih.NovaHedwigia
23:111.1967.—Fig.23c,24
Syn.:Agaricus coelestinus Fr.,Epicr.Syst. Mycol. (Upsaliae): 158. 1838
‘1836–1838’;Nolanea coelestina(Fr.)Gillet,Mém.Soc.Émul.Montbéliard,
sér.25: 536.1875;Rhodophyllus coelestinus (Fr.)Quél., Enchir.Fung.
(Paris):65.1886;Leptonia coelestina(Fr.)P.D.Orton,Trans.Brit.Mycol.
Soc.43,2:177.1960.
Pileus7–10mmbroad,conicaltohemisphericalwithumbo,not
hygrophanous, not translucently striate, with straight margin,
radiallysilkyfibrillose, squamulose at centre, uniformlydark
blue.Lamellae moderately distant, adnate-emarginate, ventri-
cose,white,becomingpinkish,withentireconcolorousedge.
Stipe20– 50×1–2mm,cylindrical,smoothoralmostsmooth,
glabrous,concolourouswithpileus,whitetomentoseatbase.
Smelland taste indistinct. Spores 6.9–8.3(– 8.9) × 5.2 –6.2
μm,Q=1.3–1.5,heterodiametrical,5–7-angledinside-view.
Basidia29.5– 37.0×8.1–9.6μm,4-spored,clavate,clamped.
Lamellae edgefertile.Cheilocystidiaabsent.Pileipellis a pla-
giotrichodermof cylindricaltoslightly inflatedhyphae10–20
μmwide with blueintracellular pigment. Clamp-connections
present.
 Habitat—Onsoilinbroad-leavedforest.
 Knowndistribution—Ural.
Specimens examined.russia,SverdlovskRegion,VisimskyNatureRe-
serve, on soil in Acer platanoides-Fraxinus excelsiorforest,21Aug.2004,
L. Marina(LE258103).
 Notes—Entoloma coelestinum is distinguished by the tiny,
very dark blue to black basidiocarps with conical hardly ex-
pandedpileuscombinedwiththesmallspores.Inthecourse
ofthephylogenetic analysis specimens previously identified
as E. coelestinum ended up in a well-supported clade, which,
however, consists itself of two sister clades that can be distin-
guishedmorphologically.Thelargercladeischaracterizedby
almostnodulosespores and a longitudinally fibrillose-striate
stipe.Itincludesblue-colouredbasidiomesandentirelyblack
ones(Fig.22e).Theothercladeconsistofonecollectionchar-
acterizedbymorepronouncedlyangled,notnodulosespores
andapolishedstipe.Thiscollectionfitswellwiththeprotologue,
and the current concept of E. coelestinum(Noordeloos2004).
Consideringthesemorphologicaldifferences,andthesignificant
p-distancebetweentheseclades(5.3%base-pairdifference)
itwasdecidedtodescribethefirstcladeasthenewspecies,
E. percoelestinum below.Unfortunately we were unable to
design a neotype for E. coelestinum since the limited material
studiedisnotfromtheoriginalgeographicarea.Moremate-
rialfromEurope,especiallyfromSwedenisneededtodoso.
12. Entoloma percoelestinumO.V.Morozova,Noordel.,Vila&
Bulyonkova, sp. nov.―MycobankMB803975;Fig.23d,e,
25
Etymology.NamedafteritssimilaritytoE. coelestinum.
Diagnosis.ThenewspeciesisclosetoE. coelestinum from which it differs
byalmostnodulosesporesandalongitudinallyfibrillose-striatestipe.
Holotype.sPain,Osona,Barcelona,LaDevesa,Rupit,alt.1050m,among
grasses and mosses, near Quercus pubescens and Fagus sylvatica, on
basicsoil,25Sept.2013,J. Vila & X. Llimona(LE254390),isotypeinJVG
1130925-24.
Pileus5–12mmbroad,conicalorhemisphericalwithumbo,not
hygrophanous, not translucently striate, with straight margin,
radiallyfibrillose,squamulose atcentre,uniformlydarkblue,
blackishblue orblack.Lamellae moderately distant, adnate-
emarginate, ventricose, white, becoming pinkish, with entire con-
colorousedge.Stipe20–40×1–2mm,cylindrical,longitudinally
fibrillose-striateoralmost smooth, concolourous with pileus,
whitelytomentoseatbase.Contextthin,concolorouswiththe
surface.Smellindistinctorfungoid,tastenotreported.Spores
6.5 –8.5 (–9.0) × 5.0 – 6.5 μm, Q=1.3 –1.5(–1.7), heterodia-
metrical,with7–9bluntanglesinside-view,almostnodulose.
Basidia 27.9 –37.0 (–45.4) × 8.1– 9.6(–13.7) μm, 4-spored,
narrowly clavate to subcylindrical, clamped. Lamellae edge
fertile.Cheilocystidiaabsent.Pileipellis a plagiotrichoderm of
cylindricaltoslightlyinflatedhyphae10–20μmwidewithblue
intracellularpigment.Clamp-connectionspresent.
 Habitat—Onsoilinbroad-leaved,mixedandpine(Pinus
sylvestris)forests.
 Knowndistribution—WesternandEasternEurope,Western
Siberia.
Additional specimens examined. russia, Penza Region, vicinities of
Poperechnoye, on soil in Fraxinus excelsiorforest,7Aug.1990,A. Ivanov
(LE18913,as E. lepidissimum); Novosibirsk, Akademgorodok, on soil in
planted Pinus sylvestrisforestSWofLavrentieva6/1,12Oct.2011,T. Buly-
onkova(LE254327);ibid.,onsoilinmixedforest,20Oct.2010,N. Filippova
(LE254341).–sPain,Barcelona,MasJoan,Espinelves,alt.730m,onplant
debris’s(Rhododendronsp.,Sequoiadendron giganteum, Piceasp.andAbies
alba),11Nov.2006,J. Vila & F. Caballero(JVG1061111-7, as E. coelestinum
(Vila&Caballero2007)).
Specimen Sporessize(μm) Q Sporesform Cheilocystidia Lamellaecolour
E. callichroumvar.callichroum 10.0–13.2×7.0 –9.4 1.4–1.6 5– 9angled, absent,somecystidia whitishwithlilactingetowards
(holotype)   almostnodulose likeclavatecellspresent edge
E. callichroumvar.venustum 8.4–12.7(16.0)×6.0 –8.6 1.3–1.6(2.5) with6 –8moderately broadlyclavateorsphaero- brightlypinkishlilaceous,pinkish
(holotype)   pronouncedangles pedunculate,intermixedwith violaceus
     basidia,45– 60×15 –28μm
E. callichroumvar.venustum 9.3–12.7×6.4 –8.2 1.4–1.8 with6–8moderately rare,broadlyclavateorsphaero- whitishwithlilactingetowards
(LE254313)   pronouncedangles pedunculate,hardlydistinguish- edge
able from the basidioles,
     30.9– 42.8×12.0 –19.0μm
E. callichroumvar.venustum 9.5–13.0(14.0)×5.7–7.2 1.4–1.8(2.0) with6–8moderately rare,broadlyclavateorsphaero- whitishwithbluishtingetowards
 (LE254312)   pronouncedangles pedunculatehardlydistinguish- edge
able from the basidioles,
     29.8– 42.7×12.9 –21.0μm
Table 3 Comparison between Entoloma callichroumvar.callichroum and E. callichroumvar.venustum.
Fig. 24 Entoloma coelestinum.a.Basidium;b.spores(LE258103).―Scale
bars=10μm.

168 Persoonia–Volume32,2014
 Notes—Intheboreal-temperate Eurasia several species
withsmallblueorblackishbluebasidiomataarerecognized.
Entoloma percoelestinum differs from E. coelestinum by almost
nodulosespores and a longitudinally fibrillose-striatestipe,
from E. chytrophilum by the smaller spores and conical, hardly
expanding pileus, and from E. lepidissimum by the smaller
sporesandlackofthebluetingeinyounglamellae.Entoloma
klofacianumischaracterizedbytheisodiametricalspores.North
American Leptonia subcoelestina is also close but it differs by
the larger spores and by the pileipellis which lacks clamps and
iscomposedofsubmoniliformcells.
13. Entoloma lepidissimum(Svrček)Noordel.,Persoonia11:
460.1982.—Fig.23f,g,26
Syn.:Leptonia lepidissimaSvrček,CzechMycol.18:205.1964;Rhodophyl-
lus lepidissimus(Svrček) M.M. Moser,Rohrlinge-Blatterpilze, 4Aufl., 2 ,
b/ 2:203.1978.
Pileus5–25mmbroad,conical,broadlyconical,hemispherical
to convex with small umbo, not hygrophanous, not translu-
centlystriate,withstraightmargin,radiallyfibrillosetoslightly
squamuloseatcentre,deepbluetoblackishblue,sometimes
discolouring to greyish violet. Lamellae moderately distant,
adnate-emarginate or almost free, ventricose, bluish, greyish
blue or bluish violaceous, becoming greyish pink, with entire
concolorousorpaleredge.Stipe20– 60×1–3mm,cylindrical,
longitudinallyfibrillose-striateoralmostsmooth,concolourous
withpileus,whitetomentoseatbase.Contextconcolorouswith
thesurface.Smellindistinct,tastenotreported.Spores7.5–11.0
×6.0– 8.0μm,Q=1.3–1.6(–1.7),heterodiametrical,with6–8
anglesinside-view.Basidia27.9– 37.0(–45.4)×8.1–9.6(–13.7)
μm,4-spored, narrowly clavate to subcylindrical,clamped.
Lamellae edgefertileorheterogeneous.Cheilocysti dia cylin-
drical, lageniform or narrowly clavate, intermixed with basidia,
sometimes rare or absent. Pileipellis a plagiotrichoderm of
cylindricaltoslightlyinflatedhyphae10– 20μmwide,withswol-
lenterminalelementsandblueintracellularpigment.Clamp-
connectionspresent.
 Habitat—Onsoilinconiferousanddeciduousforests.
 Knowndistribution—Western,CentralandEasternEurope,
RussianFarEast.
Specimens examined.CzeCh rePuBliC,Bohemiamerid., Vrabskénear
Cimelice, on fallen twigs of Alnus glutinosa in swamp Alnus forest,20Oct.
1963,M. Svrček(PRM755801,holotype,asLeptonia lepidissima).–russia,
NovgorodRegion,ValdajskyNationalPark,costofUzhynLake, onsoilin
Picea abiesforest,20Aug.2003,E. Popov (LE234755);ibid.,vicinitiesof
Sokolovo,KrasnayaGorka,onsoilinQuercus roburforest,22Aug.2003,E.
Popov(LE234751);ibid.,valleyofPoneretkaRiver,onsoilinPinus sylvestris
forest,23Sept.2011,E. Popov(LE254871);PrimorskyTerritory,Sikhote-Alin
NatureReserve,Kabanuj,25Aug.2011,E. Malysheva(LE254311).
 Notes—Entoloma lepidissimumisrecognizedbythedark
blue basidiomata with bluish lamellae. Microscopically the
scatteredcheilocystidia also can be distinctive.Despite the
fact that the blue tinge of the lamellae was not mentioned in
the protologue, all studied specimens are characterized by
bluishlamellae.Moleculardatasupporttheiridentitywiththe
holotype.ThesimilarspeciesE. coelestinum is distinguished
bythewhitelamellae,smallersporesandmoreconicalpileus.
Entoloma chytrophilum possesses white lamellae, nodulose
sporesandamoreapplanatepileus.
Acknowledgements We are very grateful to the following curators of
herbariafortheloanofthespecimens:DrsPh.Clerc(G),M.Scholler(KR),
D.Triebel(M),J.Holec(PRM), S.Ryman (UPS),E.Bulakh(VLA),W.Till
(WU)andR.Berndt(ZT).Thisworkwouldnotbepossiblewithoutthepar-
ticipation of our friends and colleagues, who contributed with material and
photosforthestudy:E. Malysheva,E.Popov,T.Bulyonkova,M.Á.Ribes,
S.Català,F.Caballero, G. Wölfel, F.Hampe,A. Hausknecht, Y.Rebriev,
K.Potapov,N.Agafonova,S.Gashkov,B.Gierzyk,V.Kapitonov,A.Kova-
lenko,V.Malysheva,A.Kiyashko,T.Svetasheva,L.Marina,O.Desyatova,
O.Shiryaeva(Kirillova),A.Fedosova,E.Ilyukhin,E.Lukashina,S.Lukashin,
E.Zvyagina,I.Ukhanova,andS.Arslanov.Weexpressoursincerethanksto
allofthem.Wearealsogratefultotheanonymousreviewersofthemanuscript
fortheirvaluableandconstructivecomments.Thisworkwas supportedin
partbytheRussianFoundationforBasicResearch(projectN12-04-33018
mol-a-vedandN13-04-00838a).
Fig. 26 Entoloma lepidissimum.a.Basidium;b.cheilocystidia;c.spores
(LE254871).―Scalebars=10μm.
Fig. 25 Entoloma percoelestinum.a.Basidium;b.spores(LE254390,
holotype).―Scalebars=10μm.

169
O.V.Morozovaetal.:Entoloma subgenus Leptonia
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