A monograph of Otidea (Pyronemataceae, Pezizomycetes)

Abstract

The easily recognised genus Otidea is subjected to numerous problems in species identification. A number of old names have undergone various interpretations, materials from different continents have not been compared and misidentifications occur commonly. In this context, Otidea is monographed, based on our multiple gene phylogenies assessing species boundaries and comparative morphological characters (see Hansen & Olariaga 2015). All names combined in or synonymised with Otidea are dealt with. Thirty-three species are treated, with full descriptions and colour illustrations provided for 25 of these. Five new species are described, viz. O. borealis, O. brunneo­parva, O. oregonensis, O. pseudoleporina and O. subformicarum. Otidea cantharella var. minor and O. onotica var. brevispora are elevated to species rank. Otideopsis kaushalii is combined in the genus Otidea. A key to the species of Otidea is given. An LSU dataset containing 167 sequences (with 44 newly generated in this study) is analysed to place collections and determine whether the named Otidea sequences in GenBank were identified correctly. Fourty-nine new ITS sequences were generated in this study. The ITS region is too variable to align across Otidea, but had low intraspecific variation and it aided in species identifications. Thirty type collections were studied, and ITS and LSU sequences are provided for 12 of these. A neotype is designated for O. cantharella and epitypes for O. concinna, O. leporina and O. onotica, along with several lectotypifications. The apothecial colour and shape, and spore characters are important for species identification. We conclude that to distinguish closely related or morphologically similar species, a combination of additional features are needed, i.e. the shape of the paraphyses, ectal excipulum structure, types of ectal excipulum resinous exudates and their reactions in Melzer’s reagent and KOH, tomentum and basal mycelium colours and exudates. The KOH reaction of excipular resinous exudates and basal mycelium are introduced as novel taxonomic characters.

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Persoonia35,2015:166–229
www.ingentaconnect.com/content/nhn/pimj http://dx.doi.org/10.3767/003158515X688000
RESEARCH ARTICLE
INTRODUCTION
Species of Otidea produce typically ear-shaped apothecia that
are unique within Pyronemataceae(Pezizomycetes).Thegenus
is monophyletic based on multilocus phylogenetic analyses
from a few, but broadly sampled, Otideaspecies (Hansen
et al. 2013). Despite being distinct at the generic level, the
speciesidentificationand nomenclature ofOtidea are highly
controversial.Afewrecenttypifications havebeenproposed
(Carbone2009,2010a),butmanynamesarestillsubjectedto
differentinterpretations.Severalnewspeciesweredescribed
inthelastdecadesfromEurope(Harmaja1976,2009a)and
Asia (Cao et al. 1990, Zhuang & Yang 2008), with detailed
descriptionsandupdated identificationkeys.However,often
no illustrations were presented and colour photographs have
rarely been published when describing new species. Many
names of European species currently used in North America
andAsiaaremisapplied.Multilocusphylogeneticanalyseshave
not been previously implemented to critically address species
delimitation issues and material from different continents has
notbeencompared.AworldwidecriticalrevisionofOtidea to
clarifyspecieslimitsishighlyneeded.Theaimsofthisstudy
were:i)toundertakeanomenclaturalandtaxonomicrevision
of Otidea, to clarify misinterpretations and to propose pertinent
typificationstostabilisethe use of names;andii)toprovide
detailed species descriptions and colour photographs of both
macro-andmicroscopicstructures,andakeyforidentification.
Our multilocus phylogenies and robust hypotheses of species
limits, employing genealogical concordance phylogenetic spe-
ciesrecognition(GCPSR;Tayloretal.2000),whichisthebasis
forthepresentwork,aregiveninHansen&Olariaga(2015).
Inthepresent study wepresentanLSUrDNAphylogenyto
place a larger number of collections for which we have been
unable to obtain multiple genes, including several sequences
fromGenBank,manyofwhichweherere-identify.
Taxonomic history
Thefirst valid publicationof Otidea is by Bonorden (1851),
based on Peziza(unranked)Otidea Pers.,althoughithassome-
times been attributed to Fuckel (Kanouse 1949, Nannfeldt
1966,Liu&Zhuang2006,Smith&Healy2009).Otidea species
were treated in a broad heterogeneous genus Peziza by early
authors. Persoon (1822) defined Peziza (unranked) Otidea
as producing auriculate apothecia with a split, sometimes
elongatedononeside,andincluded10species.Fries(1822)
referred to this group as Peziza(unranked)Cochleatae, but he
includedalsotaxawithnon-splitapothecia.Bonorden(1851)
elevated Otidea to generic rank with split apothecia as the key
feature,butalsoreferredtoFries’(1822)Cochleatae.Hedid
notmakeanycombinationsorlistanyspecies.Fuckel(1870)
refinedthe genus using microscopic details, namely uni-or
biguttulatesporesandfiliformtosubclaviformparaphyses,and
included four species: O. abietina, O. cochleata O. leporina
and O. onotica. Boudier(1885)notablycontributed todisas-
semble the large genus Peziza into smaller and more natural
genera.HeplacedinthegenusOtidea species with entire or
split apothecia, biguttulate spores and, importantly non-amyloid
asciandcurved paraphyses,whichhewas the firsttointro-
duce.HedividedOtidea into two subgenera: Otidea with split
A monograph of Otidea (Pyronemataceae, Pezizomycetes)
I.Olariaga1,N.VanVooren2,M.Carbone3,K.Hansen1
1 SwedishMuseumofNaturalHistory,DepartmentofBotany,P.O.Box50007,
SE-10405Stockholm,Sweden;
 correspondingauthore-mail:karen.hansen@nrm.se.
2 36ruedelaGarde,F-69005Lyon,France.
3 ViaDonLuigiSturzo173I-16148Genova,Italy.
Key words
Flavoscypha
ITS
ITS1minisatellites
LSU
Otideopsis
resinous exudates
AbstractTheeasilyrecognisedgenusOtideaissubjectedtonumerousproblemsinspeciesidentification.Anumber
of old names have undergone various interpretations, materials from different continents have not been compared and
misidentificationsoccurcommonly.Inthiscontext,Otidea is monographed, based on our multiple gene phylogenies
assessingspeciesboundariesandcomparativemorphologicalcharacters(seeHansen&Olariaga2015).Allnames
combined in or synonymised with Otidea aredealtwith.Thirty-threespeciesaretreated,withfulldescriptionsand
colourillustrationsprovidedfor25ofthese.Fivenewspeciesaredescribed,viz.O. borealis, O. brunneo parva, O. ore-
gonensis, O. pseudoleporina and O. subformicarum.Otidea cantharella var.minor and O. onoticavar.brevispora
areelevatedtospeciesrank.Otideopsis kaushalii is combined in the genus Otidea. A key to the species of Otidea
isgiven.AnLSUdatasetcontaining167sequences(with44newlygeneratedinthisstudy)isanalysedtoplace
collections and determine whether the named OtideasequencesinGenBankwereidentifiedcorrectly.Fourty-nine
newITSsequencesweregeneratedinthisstudy.TheITSregionistoovariabletoalignacrossOtidea, but had low
intraspecificvariationanditaidedinspeciesidentifications.Thirtytypecollectionswerestudied,andITSandLSU
sequencesareprovidedfor12ofthese.AneotypeisdesignatedforO. cantharella and epitypes for O. concinna,
O. leporina and O. onotica,alongwithseverallectotypifications.Theapothecialcolourandshape,andsporechar-
actersare importantfor speciesidentification. Weconclude thattodistinguishcloselyrelatedormorphologically
similarspecies,acombinationofadditionalfeaturesareneeded,i.e.theshapeoftheparaphyses,ectalexcipulum
structure,typesofectalexcipulumresinousexudatesandtheirreactionsinMelzer’sreagentandKOH,tomentum
andbasalmyceliumcoloursandexudates.TheKOHreactionofexcipularresinousexudatesandbasalmycelium
areintroducedasnoveltaxonomiccharacters.
Article infoReceived:26November2013;Accepted:1February2015;Published:10April2015.

167
I.Olariagaetal.:AmonographofOtidea
apothecia and Pseudotis withentire apothecia. Interestingly,
Boudier erected the genus Wynnella(Helvellaceae)toaccom-
modate W. silvicola (asP. leporina / P. auricula),aspecies
with distinctly ear-shaped apothecia, but differing from Otidea
intheuniguttulatesporesandtoughconsistency.InBoudier’s
(1907)subsequenttreatment,heelevatedPseudotis to genus
rank and placed here species with entire apothecia, including
O. da liensis(asP. apophysata)andO. propinquata(asP. abi-
etina),bothwith biguttulate sporesandhookedparaphyses.
Saccardo listed O. onoticaasan‘exemplar’speciesofPeziza
subg.Otideainhissynopsisofthediscomycetegenera(1884;
he listed in general only one species per genus / subgenus that
appeartohavebeenselectedastypicalforthegenera) and
ithassincebeenacceptedasthetypespeciesbymost(Rifai
1968,Eckblad1968,Korf1972,Liu&Zhuang2006,Parslow&
Spooner2013),IndexNominumGenericorum(EckbladinFarr
etal.1979),NCU-3(Greuteretal.1993)andispreparedtobe
adoptedontheListofProtectedgenericNamesforfungi(Kirk
etal.2013).Clements&Shear(1931)listedO. cochleata as
the type, but this name was not among the original species ac-
ceptedbyPersoon(1822)andhasfurthermorebeenconsidered
anambiguousname.Kanouse(1949)proposedO. leporina as
the type species, but because of the confusions surrounding
the identity of this species until now, it has been considered an
inappropriatechoice.
ThegenusScodellinawasdescribedbyGray(1821)andin-
volved species attributed today to Otidea, and also Aleuria,
Peziza and Tarzetta. Seaver (1928) refined Scodellina to
specieswithsplittoear-shapedapotheciaonlyandtypifiedit
with Peziza leporina(Seaver1927),consideringOtidea a later
synonym.ThiswasforaperiodfollowedbyseveralAmerican
authors(e.g.Korf1963,Kimbrough1966).Thetypificationby
Seaver can however, be considered largely mechanical, taken
asthefirstspecieslistedbyGray,andbesupersededunder
the ICN (Art. 10.5; McNeill et al. 2012). Also, even though
Gray(1821)includedseveralspecieswithsplitapothecia,he
did not mention this feature in the diagnosis of Scodellina, but
emphasised “thallus … hemispherical, spreading” and coined
thevernacularname“spreadcup”.ThereforeRifai(1968)desig-
nated P. vesiculosaBull.:Fr.asthetypespeciesofScodellina,
consequently making it a later synonym of the genus Peziza.
Eckblad(1968)cametothesameconclusion.
Kanouse(1949) broadened theconcept of Otidea and de-
scribed in detail a number of NorthAmerican species. She
included species with split apothecia and straight paraphyses
withswollenapices.ShealsoincludedWynnella silvicola(as
Otidea auricula), with straight paraphyses and uniguttulate
spores.Nannfeldt (1966) delimitedOtidea to species with
non-amyloidasci;smooth, uninucleate,biguttulatespores;a
medullary excipulum of textura intricata;andanectalexcipu-
lum with isodiametric cells, covered by short chains of barrel-
shapedcells.Nannfeldt’sconceptmoreorlessconformstothe
genus Otideaaswerecogniseittoday.HeconsideredWyn-
nella so distant that it should be treated in a separate tribe of
Pezizaceae.Korf(1963)reviewedthemonotypic,sparassoid
genus Ascosparassis,andsubsequently(Korf1973a)assigned
A. shimizuensis to Otidea, based on the hooked paraphyses,
smallbiguttulatesporesandexcipulumstructure.Pfister(1979)
however, considered Ascosparassis a distinct monotypic genus,
based on “small asci and spores and peculiar growth habit”, and
combined the older name Midotis heinricheri in Ascosparas-
sis.LaterPfistercollectedA. heinricheri in South America, in
northcoastalmountainsofVenezuela(Pfister&Halling1989),
extendingitsAsiandistribution(China,IndonesiaandJapan),
still considering the species separate from Otidea.
A new genus Flavoscypha was erected for two species of
Otidea, O. concinna(asFlavoscypha cantharella)andO. phle-
bophora, with strong emphasis on the ectal excipulum of textura
prismatica(vs textura angularis in Otidea) (Harmaja1974).
Otidea was further emended to include a species with orna-
mented spores, O. unicisa,otherwise‘fittingperfectly’Otidea
(Harmaja 1986). Otideopsis was published with Otideopsis
yunnanensis as the type species, distinguished from Otidea
by having ornamented spores and paraphyses fused at the
apices(Liu&Cao1987).Flavoscypha and Otideopsis are now
considered synonyms of Otidea based on molecular phyloge-
neticanalyses(Liu&Zhuang2006,Hansen&Olariaga2015).
RecentlythecircumscriptionofOtidea was further broadened
whenthefirsthypogeousspecies,O. subterranea, was discov-
eredusingITSandLSUsequences(Smith&Healy2009).All
the characters proposed so far as diagnostic for Otidea have ex-
ceptionsacrossthegenus.Nevertheless,Otidea can be recog-
nised by the non-amyloid asci, in combination with at least
twoofthesecharacters(exceptO. subterranea):a)biguttulate
spores;b)hookedorbentparaphyses;c)medium-large,split
apothecia;andd)amedullaryexcipulumoftextura intricata, and
an ectal excipulum of textura angularis or textura prismatica.
Systematic position and relationships
Nannfeldt(1937,1938,1966)suggestedacloserelationship
between Otidea(incl.Pseudotis),Tarzetta(asPustularia)and
Helvella (Pezizaceae, tribe Acetabuleae sensu Nannfeldt),
based on similarities in asci, paraphyses and anatomical struc-
turesof the apothecia,i.e. a medullaryexcipulum of dense
textura intricata, ectal excipulum of almost isodiametric large
cells, and an outermost layer of shorter or longer chains of
cells,possibleformingdistinctclustersorwarts(andinTarzetta
prolongedtocylindrical,hyaline,wavyhairs).LeGal(1947)
similarly placed Otidea in the tribe Otideeae in her Aleuriaceae
(i.e.afamilyincludingtaxawithbothamyloidandnon-amyloid
asci),togetherwithPseudotis and Tarzetta(asPustularia),but
placed Helvella in Helvellaceae.FollowingtheideasofNann-
feldt (1966), Eckblad (1968) erected the family Otideaceae
as a small taxon of closely related genera that produce larger
apothecia, most of which typically lack bright orange to red col-
ours, including in it Tarzetta (asPustulina)andOtidea, but also
Ascosparassis, Geopyxis and Sowerbyella.Eckbladconsidered
Helvella(Helvellaceae) to be distant, but having a possible
shared origin with members of Otideaceae(andMorchellaceae
and Rhizinaceae),duetothestructureoftheexcipulum,spores
and asci, especially of Tarzetta.Atthesametime,heexpanded
the concept of Pyronemataceae(to21genera)totaxamostly
characterised by the presence of carotenoid pigments, stating
the inability to satisfactorily subdivide the family on the basis
ofcommoncharacters.Korf (1972, 1973b) placed Otidea in
the tribe Otideeae(in thesubfamilyOtideoideae)inaneven
more encompassing Pyronemataceae (49genera),together
with Ascosparassis and Psilopezia. He followed the ideas
ofArpin (1969), inexcludingtaxawith carotenoids from the
Otideoideae, instead including taxa with prominent hairs such
as Geopora, Humaria and Trichophaea.Otidea has generally
been included in a broadly circumscribed Pyronemataceae
inrecent treatments (Dissing 2000, Hansen& Pfister 2006,
Perryetal.2007,Hansenetal.2013).Multigenephylogenetic
analyses of Pyronemataceae do not support a close relation-
ship between Otidea and Geopyxis, Psilopezia, Sowerbyella
or Tarzetta(Hansenetal.2013).Surprisingly,thecleistothecial
Warcupia and the highly reduced (gymnohymenial) Mona-
scella are suggested as the closest relatives of Otidea.Otidea,
Monascella and Warcupia are strongly supported as a distinct
sister group to the rest of the Pyronemataceae in a strict sense
(Hansenetal.2013).

168 Persoonia–Volume35,2015
MATERIALS AND METHODS
Material and morphological methods
Thisstudyisbasedonatotalof450specimens.Onehundred
and forty two of these were collected and studied fresh dur-
ingthisproject and aredepositedinSanda few inTUR-A.
Specimenswerestudiedfromthefollowingherbaria:AH,ARAN,
BIO,C,FH,H,HKAS,K,MCVE,MICH,MIN,OSC,PC,PRM,
S,TURandUPS(Thiers2014),GMFN(gruppoAMBdiFara
Novarese,Italy),SEST(Sociedad de Ciencias Naturales de
Sestao,Spain),andfromtheprivateherbariaofG.Corriol,GC;
C.Lavorato,CL;N.VanVooren,NV;andM.Tabarés,MT.Thirty
type collections were examined, along with other original mate-
rial.ColourcodesarebasedonKornerup&Wanscher(1961).
For O. apophysata, O. borealis, O. daliensis, O. oregonensis,
O. phlebophora, O. pseudoleporina, O. smithii and O. unicisa
colourcodesforfreshmaterialweretakenfromphotographs.
Apothecialsections,i.e.thethicknessoftheapotheciaincluding
the hymenium, was measured midway between the apothecial
marginand base.Smellandtastearelistedwhen recorded.
Only discharged, mature spores were measured from living
material.Toobtainandensurematuresporesfromdriedma-
terial,asquareofc.3mm2 of an apothecium was revived in
a drop of water on a slide, with the hymenium surface fac-
ingdownandthenremovedafter1 hour.Inthisway mature
spores deposited on top of the hymenium were recovered and
measured. Spore measures exclude ornamentation. Spore
statistics are based on measurements of 20 spores from each
collection: Lm=meanlength, Wm = mean width and Qm = Lm/
Wm.Thenumberofpopulationsthatthestatisticsarebasedon
isindicatedby‘n’.Extremevaluesaregiveninparentheses.
Hymenialelementswereobservedby teasing apart a small
pieceof hymeniumwitha needle.Toobservetheexcipulum
structure, sections of apothecia were made by hand prior to
soakingthematerialinwater.Basalmyceliumwasexamined
bymountingclumpsofhyphaefromtheapothecialbase(the
tomentum)andfromamongthesubstrateparticles.Onlyasci
withmature sporesweremeasured.Allmeasurements were
madeinwater: in living statewheneverpossible;onlywhen
notpossible,measurementsweremadeonrehydrated(over
2hoursinwater)material.Melzer’sreagent(MLZ)and10%
KOHwereusedtoobservethereactionofresinousexudates
andotherpigmentation.CottonBlueinlacticacidwasusedto
observe spore ornamentation. Microanatomical terminology
followsKorf(1973b).Thenotation‘!’indicatesthattypeorother
originalmaterialwasexaminedbyus.
If not otherwise indicated in the legends, the photographs pre-
sentedinthispaperweretakenbyK.HansenandI.Olariaga.
DNA extraction, PCR amplification, sequencing and
alignment
DNAwas extracted fromdriedmaterial,orfromfreshmate-
rialstored in1%SDSDNAextractionbuffer.The extraction
methodfollowsHansenetal.(1999),exceptdriedmaterialwas
groundinaMini-BeadbeaterTM(BiospecProducts,Bartlesville,
OK,USA)andfreshmaterialusingaplasticpestle,ineppen-
dorftubes.TheprimercombinationITS5-ITS4,andinafewin-
stancesITS1–ITS4,ITS5 – 5.8SandITS3– ITS4,wereusedto
PCRamplifytheITSregion,andLR0R – LR5theLSUregion.
ForDNAextractedfromfreshmaterial(storedinSDSextrac-
tionbuffer), the ITS and LSU regions were amplified in a
single piece using the primers ITS1– LR5.The ITS2 region
for O. integra (possible original material from 1892) was
successfullyamplified in threepieces, in combination with
newly designed primers for the O. concinna clade: ITS3 –
ConcITS2midR (5´-GCCTGTAAATTTTAAAGACGAA-3´);
ConcITS2midF(5´-CCAGGGTTGCTTTGGTA-3´)–ConcITS4
intR (5´-CACTGGGTAATTGGAGGTTT-3´); ConcITS2midF
(5´-CCAGGGTTGCTTTGGTA-3´)–ITS4.PCRproductswere
cleanedusingExoSAP-IT®(USB,Cleveland,OH,USA).The
ITSwassequencedinbothdirections,usingtheprimersITS1
andITS4(and/orinafewinstancesITS5,5.8SandITS3)and
theLSUusingLR0RandLR5.ForO. integra, the same prim-
ersas used for PCR,were also used for sequencing.PCR
andsequencingconditionsfollowsHansen&Olariaga(2015).
SequenceswereeditedandassembledusingSequencherv.4.10
(GeneCodesCorporationAnnArbor,Michigan,USA)andhave
beendepositedin GenBank (Table1).Thesequenceswere
alignedmanuallyinSe-Alv.2.0a11Carbon(Rambaut2002).
Anall taxaLSUdataset wasprepared.Monascella botry osa
and Warcupia terrestris were used as outgroup, based on a
higher level phylogenetic study of Pyronemataceae(Hansen
etal.2013),whichsupportstheseastheclosestsistergroup.
Toexploreinter-andintraspecificvariationofthenewspecies
O. borealis, O. subformicarum and their closest relatives, two
smallerdatasetswerepreparedusingITSandLSUrDNAse-
quences, based on a more species-inclusive, multi-gene phy-
logeny of Otidea(Hansen&Olariaga2015).Allthreealignments
areavailablefromTreeBASEas accessionno.S15887.The
firstdataset(theO. borealisdataset)containedO. borealis and
related species in the O. concinna clade.Theseconddataset
(theO. formicarumdataset)containedspecimensoftheO. for-
micarum clade. Otidea caeruleopruinosa and O. nannfeldtii
wereusedasoutgroupforeachofthesedatasets.Nucleotide
diversity(e.g.Nei1987,equation10.6),astheaveragenumber
ofnucleotidedifferencespersitesbetweentwoITSsequences,
was calculated within O. brunneoparva and O. subformicarum.
One insertion or deletion, despite the length, was calculated as
onlyonebasepairdifference.
Phylogenetic analyses
MaximumLikelihood(ML)analysesofthealltaxaLSUdata-
setwereperformedusingthe‘RAxMLHPC2onXSEDE’tool
(Stamatakis2006)viaCIPRESScienceGateway(Milleretal.
2010),employingmixedmodelsofevolutionandstartingfroma
randomtree.ForthetwosmallerO. borealis and O. formicarum
datasets,MLanalyseswereconductedusingRAxMLv.7.3.1
(Stamatakis2006)ontheBioportal,UniversityofOslo(Kumar
etal.2009).AGTR-GAMMAmodelwithfourratecategories
was assigned and all free model parameters were estimated
bytheprogram.FortheMLbootstrapanalyses(ML-BP)1000
rapid bootstrapping replicates from random starting trees were
performed,followedbyasubsequentMLsearchsimilarlyusing
1000replicates.Asnostronglysupportedconflictwasdetected
(ML-BP≥75%,PP≥95%),theITSandLSUregionwerecon-
catenated for the O. borealis and O. formicarum datasets.Each
combineddatasetwasanalysedusingMLanalysesunderthe
samesettingsasspecifiedabove.Relationshipswerelikewise
constructed using Metropolis-coupled Markov Chain Monte
Carlo (MCMCMC) and ‘model-jumping’ as implemented in
MrBayesv.3.2.1(Ronquistetal.2012).Thesubstitutionmodel
wassampled across theGTRspaceby theMCMCanalysis
(Huelsenbecketal.2004).Fourparallelsearches,eachwith
four chains, were run for ten and three million generations,
respectively, for the all-inclusive LSU dataset and the O. borea-
lis and O. formicarum datasets, initiated with random starting
trees.Thechainsweresampledevery100generations from
theposteriordistribution.Thefirst25%ofthetreessampled
wasdiscardedasthe‘burn-in’,andtheremainingtreeswere
usedtocalculatetheposteriorprobabilities(PP)oftheclades.
For the combined ML and Bayesian analyses the ITS and
LSUregionswerespecifiedasdistinctpartitions.MLbootstrap
values≥70%andPP≥95%wereconsideredtobesignificant.

169
I.Olariagaetal.:AmonographofOtidea
Monascella botryosa CBS 233.85 Spain,1985,J.Guarro – KC012688
Otidea alutacea(1) KH.09.170(S) Sweden,2009,K.Hansen&I.Olariaga KM0100591 KC012691
O. alutacea(2) KH.10.193(S) Sweden,2010,K.Hansen,K.Gillen&I.Olariaga KM0100601 KM8231881
O. alutacea (3) KH.07.46(S) Denmark,2007,H.Knudsen KM010061 KM823457
O. alutacea (4) JS.08.81(S) Sweden,2008,J.Santos KM0100621 KM8231871
O. alutacea(5) OSC56747 USA,1996,E.T.Peterson – KM8231891
O. alutacea(6) OSC56770 USA,1997,E.T.Peterson – AF072073
O. alutacea(7) OSC56798 USA,1996,E.T.Peterson – AF086583
O. alutacea (8) OSC56777 USA,1997,E.T.Peterson – AF086582
O. alutacea (9) JS.08.43(S) Sweden,2008,J.Santos KM010063 KM823458
O. alutacea (10) KH.09.135(S) Norway,2009, V.Kučera&I.Kautmanova KM0100641 KM8231901
O. alutacea (11) KH.10.198(S) Sweden,2010,K.Hansen,K.Gillen&I.Olariaga KM010065 KM823459
O. alutacea (12) KH.09.178(S) Sweden,2009,K.Hansen&I.Olariaga KM0100661 KM8231911
O. alutacea (13) KS-94-192(C) Denmark,1994,K.Hansen&S.K.Sandal KM010067 KM823460
O. alutacea (14) C-F-48045 Sweden,1974,D.Paulsen&N.Tams KM010068 KM823461
O. alutacea (15) HMAS52742 China – DQ443438
O. alutacea (16) HMAS57844 China – DQ443439
O. alutacea (17) S-F257085 Italy,2010,M.Carbone KM0100691 KM8231921
O. alutacea (18) Moorefun19(OSC) USA,2010,J.Moore KM0100701 KM8231941
O. alutacea(19) OSC56758 USA,1996,E.T.Peterson – KM8231931
O. alutacea (20)asO. umbrina OSC56813 USA,1997,E.T.Peterson – AF086584
O. alutacea (21) as O. umbrina OSC56782 USA,2010,E.T.Peterson – AF086586
O. alutacea (22) KH.09.133(S) Norway,2009,K.Hansen&I.Olariaga KM0100711 KM8231851
O. alutacea (23) ARANA3023204 Spain,2009,J.I.LópezAmiano KM0100721 KM8231861
O. alutacea (24) GC98092002 France,1998,G.Corriol KM010073 KM823462
O. alutacea (25) HMAS83560 China,2003,W.Y.Zhuang&Y.Nong – DQ443442
O. alutacea (26) HMAS83563 China,2003,W.Y.Zhuang&Y.Nong – DQ443440
O. alutacea (27) KS-94-111(C) Denmark,1994,K.Hansen&S.K.Sandal KM010074 KM823463
O. alutacea (28) HMAS83559 China,2003,W.Y.Zhuang&Y.Nong – DQ443441
O. alutacea (29) S-F257084 Italy,2010,M.Carbone KM010075 KM823464
O. alutacea(30) KH.13.50(S) Sweden,2013,K.Hansen&X.Wang KM010076 KM823465
O. angusta H6010804 Finland,1965,H.Harmaja KF7175741 KM8231951
O. apophysata S-F257062,dupl.privateherb. Germany,1999,F.Kasparek KM0100771 KM8231961
Kaspareks.n.
O. borealis S-F242694 Finland,2010,M.Carbone KM0100231 KM8231971
O. brevispora as O. onoticavar. HKAS 43003 China,2003,Z.L.Yang – DQ443450
brevispora
O. brunneoparva (1) KH.09.82(S) Sweden,2009,K.Hansen&I.Olariaga KM0100291 KM8231981
O. brunneoparva(2) S-F249386(Ex-H6017193) Finland,1978,H.Harmaja KM010024 KM823466
O. brunneoparva(3) S-F257086,dupl.TUR–A198579 Finland,2009,M.Carbone KM0100251 KM8231991
O. brunneoparva (4) JS.08.66(S) Sweden,2008,J.Santos KM010028 KM823467
O. brunneoparva (5) KH.08.107 (S) Sweden,2008,K.Hansen KM0100261 KM8232001
O. brunneoparva(6) TUR-A198582 Finland,2011,M.Lahti KM010027 KM823468
O. bufonia (1) KH.09.172(S) Sweden,2009,K.Hansen&I.Olariaga JN942764 JN941097
O. bufonia(2) JS.08.55(S) Sweden,2008,J.Santos KM010078 KM823469
O. bufonia (3) KH.07.37(S) Denmark,2007,K.Hansen JN942767 JN941098
O. bufonia(4) KH.09.248(S) Spain,2009,J.L.Teres&P.M.Pasaban JN942766 JN941084
O. bufonia (5) KH.09.249(S) France,2009,J.L.Teres KM0100791 KM8232011
O. bufonia (6) NV2009.11.01(S) France,2009,G.Moyne JN942765 JN941085
O. bufonia(7) C-F-94240 Denmark,2011,M.Sasa KP119674 –
O. caeruleopruinosa (1) H6010805 Finland,1978,H.Harmaja KF7175751 KM8232021
O. caeruleopruinosa(2) MT10082601(SCM,dupl.S) Spain,2010,M.Tabarés&S.Santamaría KM0100301 KM8232031
O. caeruleopruinosa(3) KH.13.48(S) Sweden,2013,I.-L.Walter KM010081 KM823470
O. cantharella(1) JS.08.18(S) Sweden,2008,J.Santos KM010082 KM823471
O. cantharella(2) JS.08.47(S) Sweden,2008,J.Santos KM010083 KM823472
O. cantharella (3) KH.09.125 (S) Sweden,2009,K.Hansen&I.Olariaga KM0100841 KM8232051
O. cantharella(4) NV2008.09.16(dupl.S) France,2008,J.Cavet KM0100851 KM8232041
O. concinna(1) JS.08.59(S) Sweden,2008,B.Wasstorp KM010031 KM823473
O. concinna (2) KH.09.183 (S) Sweden,2009,K.Hansen&I.Olariaga KM0100321 JN941089
O. concinna(3) KH.09.250(S) Spain,2009,F.Prieto&A.González JN942775 JN941095
O. crassa HMAS583571 China,2003,W.Y.Zhuang&Y.Nong – DQ443444
O. daliensis(1) HMAS57688 China,1988,S.Wang&W.Y.Zhuang – DQ443445
O. daliensis(2) SEST-06081702 Spain,2003,J.L.PérezButrón KM0100861 KM8232061
O. flavidobrunneola (1) H6010806 Finland,1978,H.Harmaja KF7175761 KM8232091
O. flavidobrunneola (2) H6010830 Finland,1987,P.Askola KM0100871 KM8232081
O. flavidobrunneola (3) KH.09.153(S) Norway,2009,K.Hansen&I.Olariaga KM0100881 KM8232071
O. formicarum (1) H6003350 Finland,2005,U.Salo&P.Salo KM010036 KM823474
O. formicarum (2) JS.08.63(S) Sweden,2008,J.Santos KM0100351 KM8232121
O. formicarum(3) H6003549 Finland,1970,L.Fagerström KF7175771 KM8232111
O. formicarum (4) S-F244372(dupl.O) Norway,2009,J.Lorås KM0100341 KM8232101
O. formicarum (5) KH.11.104(S) Sweden,2011,J.C.Zamora&I.Olariaga KM010033 KM823475
O. fusconigra GMFN2293 Italy,2003,G.Jamoni KM010037 KM823476
O. integra S-F108342 Italy,1892,G.Bresadola KP006504 –
O. kauffmanii(1) AH21147(MICH) USA,1917,A.H.Smith&R.J.Porter AF072095 –
Table 1Collectionsusedinthemolecularphylogeneticanalyses,withvoucherinformationandGenBankaccessionnumbersforITSandLSUregions.Some
GenBanksequencesarere-identifiedbyusandthenamesoriginallyusedinGenBankarelistedafterthetaxonnames(‘as’).Fortypespecimens(inbold)the
originalnamesarekeptregardlessofsynonymy.Numbersinparenthesesfollowingthespeciesnamesindicatemultiplecollectionsofaspecies.TheGenBank
accessions of sequences generated in this study are in bold.
Taxon Voucher Locality / year/collector GenBankAccessionno4
  ITS LSU

170 Persoonia–Volume35,2015
O. kauffmanii (2) MICH14409 USA,1915,C.H.Kauffman KF717579 –
O. kaushalii T.Læssøe6236(C,dupl.BORH) Malaysia,1999,T.Læssøe KM0101191 AF335111
O. lactea HMAS61359 (ex-MHSU 1803) China,1987,J.Z.Cao – DQ443447
O. leporina (1) HMAS83579 China,2003,W.Y.Zhuang&Y.Nong – DQ443448
O. leporina(2) HMAS83568 China,2003,W.Y.Zhuang&Y.Nong – DQ443449
O. leporina (3)asO. smithii  − − – AF0865732
O. leporina(4) OSC56824 USA,1997,E.T.Peterson – KM8232161
O. leporina (5) OSC56784 USA,1997,E.T.Peterson – KM8232151
O. leporina (6)asO.sp. HMAS583570 China,2003,W.Y.Zhuang&Y.Nong – DQ443443
O. leporina (7) JS.08.46(S) Sweden,2008,J.Santos KM010089 KM823477
O. leporina (8) KH.09.93 (S) Sweden,2009,K.Hansen&I.Olariaga KM0100901 KM8232131
O. leporina (9) JS.08.92(S) Sweden,2008,J.Santos KM010091 KM823478
O. leporina (10) NV2008.09.28(dupl.S) France,2008,N.VanVooren KM0100921 KM8232141
O. microspora AH30502(MICH) USA,1948,A.H.Smith AF072094 –
O. minor (1) H6003841 Finland,2006,U.Salo&P.Salo KM010040 KM823479
O. minor (2) KH.10.311(S) Sweden,2010,K.Hansen,K.Gillen&I.Olariaga KM0100421 KM8232181
O. minor (3) H6008618 Finland,1992,R.Saarenoksa KM0100391 KM8232191
O. minor (4) TL-Vorsø-0754(C) Denmark,1982,T.Læssøe KM010043 KM823480
O. minor (5) CL950914-01(dupl.S) Italy,1995,C.Lavorato KM0100441 KM8232201
O. minor (6) KH.98.84(C) Denmark,1998,K.Hansen KM0100411 KM8232171
O. minor (7) C-F-83445 Denmark,2007,T.Læssøe KM010038 KM823481
O. mirabilis (1) KH.09.188(S) Sweden,2009,E.Bohus-Jensen,K.Hansen& JN942770 JN941086
   I.Olariaga
O. mirabilis (2) KH.10.285(S) Sweden,2010,K.Hansen,K.Gillen&I.Olariaga KM0100941 KM8232211
O. mirabilis (3)asO. umbrina KH.01.09(C) Denmark,2001,C.Lange JN942769 AY500540
O. mirabilis (4) S-F257083 Finland,2010,M.Carbone KM010095 KM823482
O. mirabilis (5) NV2008.09.14(dupl.S) France,2008,J.Cavet JN942768 JN941094
O. mirabilis(6) S-F256929 Italy,1999,D.Bolognini KF717580 KM823483
O. myosotis H6003548 Finland,1970,L.Fagerström KF7175781 KM8232221
O. nannfeldtii (1) CL091116-17(S) Italy,2009,C.Lavorato KM010096 KM823484
O. nannfeldtii(2) S-F257096 Italy,2009,B.DeRuvo KM010097 KM823485
O. nannfeldtii (3) CL091207-01(S) Italy,2009,C.Lavorato KM010098 KM823486
O. nannfeldtii(4)(=O. lohjaënsisnom. S-F249387(Ex-H6017194) Finland,1978,H.Harmaja KM0100931 KM8232251
prov.Harmaja)
O. nannfeldtii(5) JS.08.103(S) Sweden,2008,J.Santos KM0100451 KM8232241
O. nannfeldtii(6) NV2008.10.01(dupl.S) France,2008,N.VanVooren KM0100991 KM8232271
O. nannfeldtii (7) H6002902 Finland,1972,C.-A.Haeggström KF7175811 KM8232281
O. nannfeldtii (8) rh101310(OSC) USA,2010,R.Helliwell KM0101001 KM8232261
O. nannfeldtii(9) KH.10.302(S) Sweden,2010,K.Hansen,K.Gillen&I.Olariaga KM0101011 KM8232231
O. onotica (1) OSC56801 USA,1997,E.T.Peterson AF072067 AF086578
O. onotica(2) OSC56734 USA,1996,E.T.Peterson AF072066 AF086577
O. onotica(3) OSC56759 USA,1996,E.T.Peterson – JN941088
O. onotica (4) C-F-89691 Denmark,2008,H.Knudsen JN942773 JN941090
O. onotica (5) JS.08.48(S) Sweden,2008,J.Santos KM010102 KM823487
O. onotica (6) KH.10.284 (S) Sweden,2010,K.Hansen,K.Gillen&I.Olariaga KP0065051 KM8232291
O. onotica(7) KH.09.132(S) Norway,2009,K.Hansen&I.Olariaga KM010103 KC012692
O. onotica(8) KH.09.136(S) Norway,2009,K.Hansen&I.Olariaga JN942772 JN941096
O. onotica (9) MCVE23277 Italy,2008,M.Carbone KM010104 KM823488
O. onotica(10) KH.98.107(C) Denmark,1998,K.Hansen,T.Læssøe&C.Lange – AF335121
O. oregonensis(1) rh139(S) USA,2010,R.Helliwell KM010046 KM823489
O. oregonensis(2) Moorefun 58(OSC,S) USA,2010,J.Moore KM0100481 KM8232311
O. oregonensis(3) Moorefun31(S) USA,2010,J.Moore KM0100471 KM8232301
O. oregonensis (4)asO. rainierensis OSC56829 USA,1997,M.Castellano AF072087 AF086597
O. oregonensis(5)asO. rainierensis NSW6354(OSC) USA,1990,D.McKay AF072088 AF086598
O. oregonensis(6)asO. rainierensis  OSC56745 USA,1996,J.Trappe AF072089 KM8232321
O. oregonensis(7) EGS2179(MICH) USA,1948,E.G.Simmons AF072088 –
O. papillata(1) H6003547 Finland,1971,H.Harmaja KF7175821 KM8232341
O. papillata (2) TUR102134 Finland,1990,T.Lindholm KM0101051 KM8232331
O. papillata f.pallidefurfuracea NV 2007.09.27 (S) France,2007,N.VanVooren KF7175841 KM8232351
O. phlebophora (1) JV06-385(C) Denmark,2006,L.&J.Vesterholt KM0100491 KM8232361
O. phlebophora(2) S-F108338 Sweden,1949,G.Haglund&R.Rydberg KM010050 KM823490
O. phlebophora (3) K(M)33068 UK EU784392 –
O. platyspora (1) KH.09.163(S) Sweden,2009,K.Hansen&I.Olariaga KM0101061 KM8232381
O. platyspora (2) HK0846(S) Sweden,2008,H.Kauffman KM010107 KM823491
O. platyspora(3) JV06-656(C-F-75309) Denmark,2006,J.Vesterholt KM0101081 KM8232371
O. propinquata(1) KH.09.99(S) Sweden,2009,K.Hansen&I.Olariaga KM0101091 KM8232391
O. propinquata(2) JS.08.67(S) Sweden,2008,J.Santos KM010110 KM823492
O. propinquata (3) NV2008.09.15(dupl.S) France,2008,J.Cavet KM0101111 KM8232401
O. pseudoleporina(1)asO. concinna  NSW7574(OSC) USA,N.S.Weber AF072083 AF086593
O. pseudoleporina (2)asO. concinna  OSC56749 USA,1996,E.T.Peterson AF072082 AF086592
O. pseudoleporina (3)asO. concinna  OSC56760 USA,1996,E.T.Peterson AF072081 KM8232441
O. pseudoleporina (4) rh101910 (OSC) USA,2010,R.Helliwell KM0101121 KM8232431
O. pseudoleporina (5) Moorefun14(S) USA,2010,J.Moore KM0101131 KM8232421
O. pseudoleporina (6) OSC56809 USA,1997,J.Spatafora AF072080 KM8232411
O. rainierensis A.H. Smith 30553 (MICH) USA,1948,A.H.Smith KF7175831 KM8232451
O. sinensis HMAS61360 China – DQ443451
O. smithii(1) OSC56799 USA,1997,E.T.Peterson AF072063 JN941087
O. smithii(2) ecv3345(S) USA,2005,E.Vellinga JN942771 JN941093
Table 1(cont.)
Taxon Voucher Locality / year/collector GenBankAccessionno4
  ITS LSU

171
I.Olariagaetal.:AmonographofOtidea
RESULTS
Alignment and ITS minisatellites
Fourty-nineITSand44LSUsequenceswerenewlygenerated
inthisstudy(Table1).Intotal146ITSsequenceswereutilised,
including34 obtainedfromGenBankand63fromHansen&
Olariaga(2015).TheITSsequencesweretoovariabletoalign
across all of Otidea,duetoahighlypolymorphicpartinITS1
andlargelengthvariability(insertionsanddeletions),andthere-
fore were not included in phylogenetic analyses of the entire
genus.TheITS sequences were alignedamong closely re-
lated species or species groups, and used as an aid to verify
identifications.TheITSregionwasespeciallyusefulincases
weretheprotein-codinggenes(RPB1,RPB2andEF1)failed
toamplify,duetopoorqualityDNAfromoldorpoorlytreated
material.ITSsequencesofthetypespecimensofO. mirabilis
and O. kauffmanii thatcouldnotbeamplifiedforthemultiple
genesareprovidedhere.ThealltaxaLSUalignmentconsisted
of167LSUsequences,including57 from GenBank and 68
fromHansen&Olariaga(2015)(Table1)and956bpincluding
insertedgaps, of which 263 bpwere parsimony informative
characters.
TheO. borealisdatasetconsistedof31taxa,representedby
24complete ITS-LSU andsixITSsequences, and1555bp
includinginsertedgaps(ITS 614 bp; LSU941bp),ofwhich
216wereparsimonyinformativecharacters.TheO. formicarum
datasetconsistedof14sequencesand2531bp(ITS1695bp;
LSU836bp).PartoftheITS1(805bp)intheO. formicarum
dataset was omitted from the analyses, due to a long insertion
and tandem repeats in O. subformicarumandthetwoMexican
specimens,andthecombineddatasetthusincluded1726bp,
ofwhich145wereparsimonyinformativecharacters.Theinser-
tionwascomposedoffourtandemrepeats(minisatellites)in
the four O. subformicarumsequences(212bpintotal),andin
oneofthem(S-F256979)therepeatwaspresentafifthtime
(275bpintotal).IntheMexicanFH301036theinsertionwas
extremelylong(715bpintotal),ofvariabletorandomrepeats.
Thetandem repeats were 43 or 63 nucleotides.Theywere
composedofthreeparts(A-B-C)of30,20and13nucleotides,
respectively, which were duplicates of the preceding part of
theITS1 sequence. Inthe first tworepeats the B part was
missing,whereasinthethird-fifthallpartswerepresent.The
Cpartwas100%identicalinallspecimensandrepeats;the
Apartwas mostlyidentical,butshowed3.3– 6.7%variation
inonerepeat;andtheBpartshowed5–15%variationinthe
repeats.Toascertainthecorrectnessofthelonginsertionin
theMexicanspecimens,theITSsequenceofFH301036was
amplifiedandsequencedtwice,usingdifferentsetsofprimers
(inonepieceusingITS1-ITS4andintwopiecesusingITS5-
5.8S/ITS3-ITS4).Thetwosequenceswerefoundtobeidenti-
cal.Unfortunately,wewerenotabletosequencethecomplete
insertionofFH301035andtheITS1wasonlysequencedinone
direction;boththepartoftheinsertionrecoveredandtheITS1
werehighlydifferentfromFH301036.
All taxa LSU phylogeny
TheMLanalysisofthealltaxaLSU dataset resulted in a single
best ML tree of -lnL=6511.68. Bayesian analyses reached
an average standard deviation of split frequencies of 0.004
after 10M generations. A majority rule consensus tree was
constructed from the 300000 trees sampled from the four
runs,eachconsistingof75000treessampledfromthestation-
O. smithii(3) OSC56753 USA,1996,E.T.Peterson AF072062 AF086574
O. smithii(4) OSC56811 USA,1997,E.T.Peterson AF072060 AF086572
O. subformicarum (1) S-F242696 Spain,2012,J.Herranz&J.C.Campos KM010054 KM823495
O. subformicarum(2) S-F256979 Spain,2008,J.FernándezVicenteetal. KM010051 KM823494
O. subformicarum(3) CL050928-30,dupl.S-F256978 Italy,2005,C.Lavorato KM0100521 KM8232471
O. subformicarum(4) Privateherb.CMP1179,RM1095, Spain,2009,C.M.PérezdelAmo&R.Gil KM0100531 KM8232461
dupl.S-F256980
O.aff.subformicarum (1) FH301035 Mexico,2007,M.Hernández KM0100551 KM8232491
O.aff.subformicarum(2) FH301036 Mexico,2007,M.E.Smith KM0100561 KM8232481
O. subterranea(1) RH97(FH) USA,1997,R.Healy FJ404766 FJ404766
O. subterranea(2) RH69 (FH) USA,1997,R.Healy FJ404767 FJ404767
O. tuomikoskii(1) JS.08.68(S) Sweden,2008,J.Santos KM010114 KM823496
O. tuomikoskii(2) MK200065(S) Sweden,2000,M.Karström KM010115 KM823497
O. tuomikoskii (3) H6002901 Finland,1972,R.Tuomikoski KF7175851 KM8232501
O. tuomikoskii(4) JS.08.100(S) Sweden,2008,J.Santos KM010116 KM823498
O. tuomikoskii(5) NV2008.09.08(S) France,2008,N.VanVooren JN942777 JN941091
O. tuomikoskii(6) KH.09.130(S) Norway,2009,K.Hansen&I.Olariaga JN942776 JN941092
O. tuomikoskii(7) KH.11.77(S) Sweden,2011,M.Prieto&I.Olariaga KM010117 KM823499
O. tuomikoskii(8)asO. leporina − − – AF0865883
O. tuomikoskii(9) OSC56756 USA,1996,E.T.Peterson AF072084 AF086594
O. tuomikoskii(10) OSC56826 USA,1996,M.Madsen&R.Davis AF072086 AF086596
O. tuomikoskii(11) OSC56761 USA,1996,E.T.Peterson AF072085 KM8232511
O. unicisa(1) KH.06.06(FH) USA,2006,L.Millman – KC012693
O. unicisa (2)asO. grandis HMAS51684 USA,Burdsall – DQ443446
O. unicisa(3)asO. grandis  ZWGeo65-Clark(S) USA,2003,Z.Wang KM010118 AY789369
O. yunnanensis HMAS82166 China,2003,Z.L.Yang – DQ443452
O.sp.‘a’(1) MK0942(S) Sweden,2009,M.Karström KM010057 KM823500
O.sp.‘a’(2) MK1081(S) Sweden,2010,M.Karström KM010058 KM823501
O.sp.‘b’ KH.09.79(S) Sweden,2009,K.Hansen&I.Olariaga KM0101201 KM8232521
Warcupia terrestris CBS891.69 Canada,1966,J.W.Paden – DQ220467
1 SequencesfromHansen&Olariaga(2015).
2 ThevoucherspecimenforAF086573ismistakenlygivenasOSC 56823inGenBank.Thisvoucher (OSC56823)isO. smithiibasedonmorphologicalre-examinationand theITSsequence
(AF072061)depositedbythesameauthors.TheLSUsequenceAF086573isO. leporina(Fig.1).
3 ThevoucherspecimenforAF086588ismistakenlygivenasOSC56825inGenBank.Thisvoucher(OSC56825)isO. leporinabasedonmorphologicalre-examinationandtheITSsequence
(AF072078)depositedbythesameauthors.TheLSUsequenceAF086588isO. tuomikoskii(Fig.1).
4 ITS:Internaltranscribedspacers(ITS1andITS2)andthe5.8SgeneofthenrDNA;LSU:28SlargesubunitofthenrRNAgene.
Table 1(cont.)
Taxon Voucher Locality / year/collector GenBankAccessionno4
  ITS LSU

172 Persoonia–Volume35,2015
Fig. 1Bayesianinference 50% majorityruleconsensus phylogram ofOtideafrom LSU sequencedata.Maximum Likelihood bootstrapvalues(ML-BP)
≥70%andBayesian posteriorprobabilities(PP)≥95%are shownaboveandbelowthe branches,respectively.Thickenedbranchesreceivedsupportby
bothML-BP≥70%andPP≥95%.Typecollectionsarehighlightedinbold.CountryoforiginforeachcollectionisgivenusingISOcountrycodes.Namesof
speciesrecognisedareindicatedbytheverticalbars.
O. cantharella (1) SE
O. cantharella (2) SE
O. cantharella (3) SE
O. cantharella (4) FR
O. brunneoparva (1) SE
O. brunneoparva (2) FI
O. brunneoparva (4) SE
O. brunneoparva (5) SE
O. propinquata (1) SE
O. propinquata (2) SE
O. propinquata (3) FR
O. tuomikoskii (1) SE
O. tuomikoskii (2) SE
O. tuomikoskii (3) FI
O. papillata f. pallidefurfuracea FR
O. tuomikoskii (4) SE
O. tuomikoskii (5) FR
O. tuomikoskii (6) NO
O. tuomikoskii (7) SE
O. tuomikoskii (8) US
O. tuomikoskii (9) US
O. tuomikoskii (10) US
O. tuomikoskii (11) US
O. pseudoleporina (1) US
O. pseudoleporina (2) US
O. pseudoleporina (3) US
O. pseudoleporina (4) US
O. pseudoleporina (5) US
O. pseudoleporina (6) US
O. leporina (3) US
O. leporina (4) US
O. leporina (5) US
O. crassa CHN
O. leporina (6) CHN
O. leporina (7) SE
O. myosotis FI
O. leporina (8) SE
O. leporina (9) SE
O. leporina (10) FR
O. papillata (1) FI
O. papillata (2) FI
O. alutacea (1) SE
O. alutacea (2) SE
O. alutacea (3) DK
O. alutacea (4) SE
O. alutacea (5) US
O. alutacea (6) US
O. alutacea (7) US
O. alutacea (8) US
O. alutacea (9) SE
O. alutacea (10) NO
O. alutacea (11, 30) SE
O. alutacea (12) SE
O. alutacea (13) DK
O. alutacea (14) SE
O. alutacea (15) CHN
O. alutacea (16) CHN
O. alutacea (17) IT
O. alutacea (18) US
O. alutacea (19) US
O. alutacea (20) US
O. alutacea (21) US
O. alutacea (22) NO
O. alutacea (23) ES
O. alutacea (24) FR
O. alutacea (25) CHN
O. alutacea (26) CHN
O. alutacea (27) DK
O. alutacea (28) CHN
O. alutacea (29) IT
O. apophysata GE
O. daliensis (1) CHN
O. daliensis (2) ES
O. subterranea (1) US
O. subterranea (2) US
O. platyspora (1) SE
O. platyspora (2) SE
O. platyspora (3) DK
Monascella botryosa ES
Warcupia terrestris CA
93
100
91
96
89
100
100
100
100
99
99
-
74
96
-
98
100
99
-
93
100/100
100
clade 1
clade 2
clade 3a
clade 4
O. alutacea s.str.
clade 3b
100
100
100
97
100
100
98
84
78
98
100
100
100
100
100
100
100
100 100
98
100
100
75
95
97
94
100
71
100
100
100
100
98
100
100
O. cantharella
O. brunneoparva
O. propinquata
O. tuomikoskii
O. leporina
O. pseudoleporina
O. papillata
O. alutacea complex
O. daliensis
O. subterranea
O. apophysata
O. platyspora
O. brunneoparva (3) FI
75
-
99
100
75
76
0.4

173
I.Olariagaetal.:AmonographofOtidea
Fig. 1(cont.)
100
O. bufonia (1) SE
O. bufonia (2) SE
O. bufonia (3) DK
O. bufonia (4) ES
O. bufonia (5) FR
O. bufonia (6) FR
99
O. mirabilis (1) SE
O. mirabilis (2) SE
O. mirabilis (3) DK
O. mirabilis (4) FI
O. mirabilis (5) FR
O. mirabilis (6) IT
O. leporina (1) CHN
O. leporina (2) CHN
O. smithii (1) US
O. smithii (2) US
O. smithii (4) US
O. smithii (3) US
O. flavidobrunneola (1) FI
O. flavidobrunneola (2) FI
O. flavidobrunneola (3) NO
O. brevispora CHN
O. onotica (1) US
O. onotica (3) US
O. onotica (2) US
O. onotica (4) DK
O. onotica (5) SE
O. onotica (6, 7, 8) SE, NO
O. onotica (9) IT
O. onotica (10) DK
O. unicisa (1) US
O. unicisa (2) US
O. unicisa (3) US
O. kaushalii MY
O. yunnanensis CHN
O. minor (1) FI
O. minor (2) SE
O. minor (3) FIO. minor (3) FI
O. minor (4) DK
O. minor (5) IT
O. minor (6) DK
O. minor (7) DK
O. lactea CHN
O. oregonensis (1) US
O. oregonensis (2) US
O. oregonensis (3) US
O. oregonensis (4) US
O. oregonensis (5) US
O. oregonensis (6) US
O. phlebophora (1) DK
O. sp. ‘a’ (1) SE
O. sp. ‘a’ (2) SE
O. phlebophora (2) SE
O. borealis FI
O. rainierensis US
O. concinna (1) SE
O. concinna (2) SE
O. concinna (3) ES
O. caeruleopruinosa (1, 3) FI, SE
O. caeruleopruinosa (2) ES
O. sinensis CHN
O. subformicarum (1) ES
O. subformicarum (2) ES
O. aff. subformicarum (1) ME
O. aff. subformicarum (2) ME
O. formicarum (1) FI
O. formicarum (2) SE
O. formicarum (3) FI
O. formicarum (4) NO
O. formicarum (5) SE
O. sp. ‘b’ SE
O. nannfeldtii (1) IT
O. nannfeldtii (2) IT
O. nannfeldtii (3) IT
O. angusta FI
O. nannfeldtii (4) FI
O. nannfeldtii (5) SE
O. nannfeldtii (6) FR
O. nannfeldtii (7) FI
O. nannfeldtii (8) US
O. nannfeldtii (9) SE
94
97
100
99
91
100
100
100
98
71
100
100
-
100
100
99
89
98
100
99
91
-
99
98
100
73
-
100
100
99
100
-
98
100
100
100
71
91
82
100
100
100
100
100
100
95
99
100
100
100
95
100
100
100
97
98
100
97
100
99
91
-
100
100-
O. bufonia
O. mirabilis
O. smithii
O. flavidobrunneola
O. brevispora
O. onotica
O. unicisa
O. yunnanensis
O. minor
O. lactea
O. oregonensis
O. sp. ‘a’
O. sp. ‘b’
O. phlebophora
O. borealis
O. rainierensis
O. concinna
O. caeruleopruinosa
O. sinensis
O. subformicarum
O. formicarum
O. nannfeldtii
O. kaushalii

174 Persoonia–Volume35,2015
arytreedistribution (the first25%discardedas the burn-in)
(Fig.1).The MLandBayesiantreetopologies were congru-
ent and recovered the same moderate to strongly supported
clades(Fig.1).Theterminalcladesthatconstitutespecieswe
recognise(Hansen&Olariaga2015)havemoderatetostrong
support in both analyses, except for O. leporina and O. mirabilis.
A few synonymies inferred from sequences of type collections
from which we were not able to obtain the protein-coding
genes, and clear misidentifications of sequences deposited
inGenBankareevidencedbytheLSUphylogeny(seeTable
1).BasedonLSUsequencesfromGenBank:theholotypeof
O. crassa is nested within the O. leporinaclade(=O. leporina
and O. pseudoleporina);O. lactea is suggested to be a sister
lineage to O. minor; O. sinensis is resolved as a sister species
to O. caeruleopruinosa,butwithoutsupport;O. yunnanensis
forms a monophyletic group with O. kaushalii and O. unicisa;
and the type of O. onotica var.brevispora(=O. brevispora)is
supported as a sister lineage to O. onotica. Theholotypeof
O. mirabilis is nested among other collections of O. mirabilis
and the LSU sequence differs only in 1 bp from sequences of
collectionsfromScandinaviaandFrance.
The O. formicarum and O. concinna clades in separate
ITS-LSU phylogenies
TheMLanalysisoftheO. formicarum dataset recovered a single
treeof -lnL=3579.93 (Fig. 2). Bayesian analyses reached
an average standard deviation of split frequencies of 0.003
after 3M generations. A majority rule consensus tree was
constructedfromthe90004treessampledfromthefourruns,
each consisting of 22 501 trees sampled from the stationary
treedistribution(thefirst25%discardedastheburn-in).The
four collections of O. subformicarum from Spain and Italy form
adistinct,stronglysupportedmonophyleticgroup(ML-BPand
PP100%).Likewise,thefivecollectionsofO. formicarum from
Fennoscandia, and the two collections of O. aff. subformicarum
fromMexico, eachformseparate, stronglysupportedclades
(ML-BPandPP100%;ML-BP84%,PP100%,respectively).
Phylogeneticanalyses ofthecombined ITS-LSUdatasetfail
however, to resolve relationships among these three clades
withanycertainty.
ThecombinedMLanalysisoftheO. borealis dataset resulted in
asinglebestMLtreeof-lnL=4230.82(Fig.3).Bayesiananaly-
ses reached an average standard deviation of split frequencies
of0.005after3Mgenerations.Amajorityruleconsensustree
was constructed, as for the O. formicarumdataset (above).
The supported topology (PP≥95%) did not differ from the
supportedtopologyrecoveredbyMLanalysis.Theplacement
of the single O. borealiscollectionisunresolved,buttheITS-
LSUphylogenyconfirmsitisgeneticallydivergentfromitssister
species(Fig.3).Otherspecieswithayellowoutersurfaceofthe
apothecia, O. concinna, O. minor, O. oregonensis and O. phle-
bophora,are each stronglysupported as monophyletic (ML
97–99%,PP100%).AnexceptionisO. minor(5)fromItaly
that is resolved as a sister lineage to the rest of the O. minor
collections and O. integrawithoutsupport.TheITSandLSU
sequences of O. minor(5)differby16and5bp,respectively,
from the rest of the sequences of O. minor,whichare100%
identical, except for the ITS sequence of O. minor (7) that
differsin1bp.Otidea integraisrepresentedonlybytheITS2
region(281 bp).TheholotypeofO. rainierensis is forming a
0.0080
O. aff. subformicarum (1) ME
O. formicarum (5) SE
O. formicarum (4) NO
O. formicarum (3) FI
O. formicarum (2) SE
O. formicarum (1) FI
O. aff. subformicarum (2) ME
O. subformicarum (2) ES
O. subformicarum (3) IT
O. subformicarum (4) ES
O. subformicarum (1) ES
O. nannfeldtii (7) FI
O. angusta FI
O. nannfeldtii (5) SE
100
84
100
100
100
100
100
100
100
100
100
100
Fig. 2ThesinglebesttreeresultingfromtheMaximumLikelihoodanalysis
oftheITS-LSU regions of the O. formicarum clade.MLbootstrap values
(ML-BP)areshownabovenodesandBayesianposteriorprobabilities(PP)
belownodes.Thickenedbranchesarenodeswithhighsupport(ML-BP≥75;
PP≥95).Typecollectionsareinbold.
0.02
O. microspora US
O. rainierensis US
O. kauffmanii (1) US
O. kauffmanii (2) US
O. integra IT
O. minor (7) DK
O. minor (3) FI
O. minor (1) FI
O. minor (6) DK
O. minor (2) SE
O. minor (4) DKO. minor (4) DK
O. minor (5) IT
O. oregonensis (1) US
O. oregonensis (3) US
O. oregonensis (2) US
O. oregonensis (4) US
O. oregonensis (5) US
O. oregonensis (6) US
O. oregonensis (7) US
O. sp. ‘a’ (1) SE
O. sp. ‘a’ (2) SE
O. phlebophora (3) UK
O. phlebophora (2) SE
O. phlebophora (1) DK
O. borealis FI
O. concinna (3) ES
O. concinna (1) SE
O. concinna (2) SE
O. caeruleopruinosa (2) ES
O. caeruleopruinosa (1) FI
100
100
100
100
97
99
100
100
100
100
98
99
99
100
97
100 98
100
100
100
Fig. 3ThesinglebesttreeresultingfromtheMaximumLikelihoodanalysis
oftheITS-LSUregionsofO. borealisandcloselyalliedspecies.MLbootstrap
values(ML-BP)areshownabovenodesandBayesianposteriorprobabilities
(PP)belownodes.Thickenedbranchesarenodeswithhighsupport(ML-BP
≥75%;PP≥95%).Typecollectionsareinbold.

175
I.Olariagaetal.:AmonographofOtidea
strongly supported clade with the holotype of O. kauffmanii
and a paratype of O. microspora(ML97%,PP100%).The
O. kauffmanii and O. microspora types are only represented
byITSandarethereforenotincluded inanyofourprevious
analyses.WeconcludeO. kauffmanii is a synonym of O. rainie-
rensis and O. microsporaadoubtfulname(seefurtherunder
Taxonomy).Otidea concinna is strongly supported as a sister
group to the rest of the ingroup, but otherwise the relationships
amongthespeciesarewithoutsupport.
Thephylogeneticresults,andtheITSsequencesimilarityand
divergence(forspeciesidentification),willbefurtherdiscussed
whereapplicableinthedescriptivenotesbelow.
Morphological characters for species delimitation
All Otidea species recognised by concordance of our four genes
phylogenies(Hansen&Olariaga2015)canberecognisedby
acombinationofmorphologicalcharacters.Weevaluatedthe
characters in the context of the phylogeny and discovered
severalnewcharacters.The apothecial shape, colours, and
sporecharacters(size, shape,ornamentation)areimportant
forspeciesidentification,buttodistinguishcloselyrelatedspe-
cies(orotherwisemorphologicallysimilarspecies)additional
charactersareneeded.Thesearetheshapeoftheparaphyses,
ectal excipulum structure, type of exudates on the medullary
excipulum hyphae, resinous exudates on the outer surface of
the ectal excipulum and on the mycelium at the base of the
apothecia,andtheirpossiblereactionsinMLZandKOH(see
furtherinHansen&Olariaga2015).Belowweprovidedetails
ontheresinousexudates,andtheirreactionsinKOHandMLZ,
becausetheylargelyhavebeenoverlooked.
Excipular resinous exudates and reactions in MLZ and KOH
A resinous exudate is here used for a substance that is released
from the cells and in many species is vulnerable to common
mountants,butnotwater(followingHuhtinen1990).InOtidea
the exudates are concentrated in the excipulum cells, and on
the tomentum and mycelium at the base of the apothecia.
The resinous exudates are deposited on the outside of the
cellorhyphalwalls.Harmaja(2009a)introducedthereaction
of coloured resinous exudates on the outermost cells of the
ectalexcipuluminMLZasataxonomiccharacter.Ourstudy
revealed in addition, different reaction patterns of exudates
on the medullary excipulum cells and on the mycelium at the
baseoftheapothecia(includingthetomentum),extendingout
among the soil particles, which turned out to be diagnostic for
somespecies(seeunderMyceliumatthebaseoftheapothe-
cia). In the medullary excipulum, most frequently scattered,
goldenbrown,resinousexudatesarepresentatsepta,e.g.in
O. alutacea, O. leporina, O. nannfeldtii(Fig.4a).Otidea bufonia
has unique exudates, wrapping some hyphae and appearing
striate(referredtoas‘fingerprint-like’byKorf&Zhuang(1991)),
sometimesformingbigcrystal-likeaggregates(Fig.4b,c).In
contrast, the sibling species, O. mirabilis, has only sometimes
biflabellate crystal-like exudates in the medullary excipulum
(Fig.4d).Otidea papillata also possesses unique brown exu-
dates, embedding some hyphae of the medullary excipulum
andsometimesappearingrod-like.
Small, resinous drops or amorphous matter are present in vari-
able amounts on the outer surface of the apothecia of nearly
all Otidea species. In most species they are abundant and
Fig. 4Medullaryexcipulum resinous exudatesinOtidea. a. Hyphaewithgolden brown resinousexudatesat septa inO. leporina (KH.11.02),inwater*;
b,c.browncrystal-likeexudatesinO. bufonia (KH.07.37)inwater†:b.overview;c.close-upofhyphaewrappedinstriateexudates;d.close-upofbiflabellate
crystal-like exudates in O. mirabilis(GMFN1951,holotype),inwater†—Scalebars=10µm;*=freshmaterial;†=driedmaterial.

176 Persoonia–Volume35,2015
easytoobserveinwater,butscarceanddifficulttodetectina
few(O. alutacea, O. formicarum).Onespecies,O. kaushalii,
hasauniquetypeofexudate,i.e.crystal-like,oblatespheroid,
striatebodies,withaconstrictedcentre.Thepossiblereaction
oftheexudatesinMLZandKOHisusefultoseparatecertain
speciesorgroups.Usingthesecharactersrequiresexperience.
Thereactionappearstovarydependingontheamountofexu-
dateandtheconcentrationofMLZ.AsHarmaja(2009a),we
observedtwotypesofreactionsinMLZ:i)Resinousexudates
dissolveandcoalesceintospheroiddrops,‘amberdrops’,that
containhyalinebubbles(Fig.5a).The‘amberdrops’areover-
lookediftheectalexcipulumisdirectlymountedinMLZ,since
the exudates coalesce instantly and the drops can be washed
away.ThereactionisbestobservedifMLZisaddedtoawater
mount.Thisreactionispresentinmanyspecies.ii)Resinous
exudatespartlyconvertintosmallreddishparticles(Fig.5b).
Thereactionisoftensubtleandvisibleonlyincertainpartsof
amount.ThisreactionistypicalinO. bufonia, O. mirabilis and
O. smithii.InsomespeciestheexudatesdonotreactinMLZ
ortheysimplydissolve.
Fig. 5ReactionsofresinousexudatesontheoutermostectalexcipulumcellsinOtidea.a.O. nannfeldtii† (H6010804,holotypeofO. angusta),inwater(left)
andconvertingintoamberdropsinMelzer’sreagent(right);b.O. bufonia*(KH.09.171),inwater(left)andconvertingintoreddishparticlesinMelzer’sreagent
(right);c.O. borealis† (S-F242694,holotype),inwater(left)andturningbrightyellowinKOH(right);d.O. nannfeldtii † (H6010804),inwater(left)andturning
reddishbrowninKOH(right);e.O. pseudoleporina†(rh101910,holotype),inwater(left)andconvertingintoreddishgreydropsinKOH(right);f.ectalexcipulum
cells showing a gelatinous sheath in O. formicarum* (KH.11.104).—Scalebars=10µm;*=freshmaterial;†=driedmaterial.

177
I.Olariagaetal.:AmonographofOtidea
In this study we detected three discriminative reactions of re-
sinousexudatesin10%KOH,whichweproposeasanovel
taxonomiccharacter:i)Inwatertheresinousexudatesrange
fromyellow todarkreddish yellow(Fig.5c, left)andin KOH
these dissolve completely ± exuding a yellow pigment, or partly
withtherestturningdistinctlybrighteryellow(Fig.5c,right).This
reaction occurs in O. concinnaandcloselyrelatedspecies,viz.
O. borealis, O. caeruleopruinosa, O. flavidobrunneola, O. kaus-
halii, O. minor and O. oregonensis, and slightly less strikingly in
O. unicisa.ii)Theresinousexudatesareyellow-browninwater
(Fig.5d,left)and turnreddishbrowninKOH(Fig.5d,right).
ThisreactionhasbeenobservedinO. nannfeldtii and O. pro-
pinquata.iii) The resinous exudates,yellow brown inwater
(Fig.5e,left),partlydissolveinKOH,andpartlycoalesceinto
heterogeneous, pale reddish grey drops, with bubbles inside
(Fig.5e, right).Thesedropsare similartothe‘amberdrops’
observedinMLZinmanyspecies,exceptforthepaleredcolour.
ThisreactionhasbeenobservedinO. leporina and O. pseudo-
leporina.Anumberofspecieshavetheoutermostcellsofthe
ectal excipulum sometimes covered with a thin gelatinous
sheath(Fig.5f).
Mycelium at the base of the apothecia
All Otidea species studied showed a conspicuous tomentum
covering the base of the apothecia and spreading out in the
substrate.Inthemicroscopicdescriptionswerefertobothas
thebasalmycelium.Thehyphaeareseptate,straightandfre-
quentlybranchandanastomose.Norhizomorphshavebeen
observed, although slender hyphal threads are sometimes
formed.Resinousexudatesareoftenpresentonthesurface,
especially on the mycelia extending out in the substrate, and
canappearlikehyphalornamentation.
Harmaja(2009a)proposedthecolourofthebasaltomentum
as a taxonomic character, but microscopic features of the basal
mycelium have been largely overlooked and have not been
includedinspeciesdescriptions.Twotypesofcharactersare
usefulforspeciesidentification:i)Resinousexudatesoccurring
onthehyphalwalls(Fig.6a).Thesecanbeinconspicuousor
nearly absent in some species, such as in O. alutaceas.l.,in
which only a few refractive drops or minute hyaline exudates
aresometimespresent.Resinousexudatesarealsoscarcein
some species of the O. concinnaclade. Manyotherspecies
show hyphae densely covered with resinous exudates, such as
O. formicarum, O. propinquata and O. tuomikoskii.Theshapeof
theexudatesisvariable,i.e.rod-shaped,hemispherical,conical
orbipyramidal.TheydissolveandcompletelydisappearinMLZ
(Fig.6b).Sometimesdoubtscanariseaboutthenatureofthe
differently shaped exudates, but the facts that they detach from
the hyphal wall when the mount is squashed and dissolve in
MLZ,showthesearenottrueornamentation,i.e.notpartofthe
hyphalwall.ii)ThehyphalwallturnsyellowinKOH(Fig.6d;
seeFig.6cinwater).Wehaveobservedthisreactionclearly
in O. borealis and O. onotica. Itcanbeobserved inisolated
hyphae, but is more conspicuous when a mass of hyphae is
observedtogether.Itcanalsobeobservedmacroscopically.
Fig. 6MyceliumatthebaseoftheapotheciaandextendingoutinthesubstrateinOtidea†.a,b.O. flavidobrunneola(H6010806,holotype):a.resinousexu-
datesonthehyphalwallsinwater;b.dissolvedinMelzer’sreagent;c,d.O. borealis(S-F242694,holotype):c.paleyellowinwater;d.turningbrightyellowin
KOH.—Scalebars=10µm;†=alldriedmaterial.

178 Persoonia–Volume35,2015
TAXONOMY
Otidea (Pers.)Bonord.,Handb.Mykol.:205.1851
 ≡Peziza(unranked)Otidea Pers.,Mycol.Eur.1:220.1822.
 ≡Peziza (unranked)Cochleatae Fr., Syst.Mycol.2:46.1822:Fr.loc.cit.
Type species.Otidea onotica (Pers.:Fr.)Fuckel,indicated by Saccardo,
Bot.Centralbl.18:215.1884(‘P. onoticaPers.’).
= FlavoscyphaHarmaja,Karstenia14:107.1974.
Type species.Peziza phlebophoraBerk.&Broome.
= OtideopsisB.Liu&J.Z.Cao,ShanxiUniv.J.,Nat.Sci.Ed.4:70.1987.
Type species.Otideopsis yunnanensis B.Liu&J.Z.Cao.
Apotheciasmalltolarge,3–75mmhigh,4 – 80mmwide,often
in fascicles or caespitose, epigeous, cup- to ear-shaped and
splittothebaseononeside,lessoftenentire,stipitateornot;
orhypogeousandenclosed.Hymenium white, yellow, ochre,
brown,almostblack,oftenwithpinkstains.Receptacle surface
concolorous or with similar colours as hymenium, sometimes
with purplish, greenish or bluish tones, with conical to broadly
conical warts or pustules, less often smooth or furfuraceous,
concolorousordarkerthanthebackground.Baseoftheapo-
thecium tomentose, mycelium white, ochre, yellow, orange or
brown, extending out in the substrate, base ribbed-veined in
afew species.Spores uniseriate, ellipsoid, oblong or fusoid,
typically with 2 guttules, sometimes with a few smaller gran-
ules,smooth (orverruculoseinSEM),rarelyspinoseorwith
lowridges,withdeBarybubblesinMLZandCottonBluewhen
dried, thin-walled to slightly thick-walled, hyaline to very pale
brown.Paraphyses typically curved to hooked, rarely straight,
sometimes with notches or swollen at the apices, septate, typi-
cally containing refractive small guttules at the apices, fading in
colourandcollapsedwhendried.Asci cylindrical, operculate,
8-spored,116–275×8–19µm,withpleurorhynchousbase.
Subhymeniumc.100 –150µmthick,ofdensetextura intricata,
hyphae sometimes swollen, often with scattered pigmented
exudatesatsepta.Medullary excipulum400–1500µmthick,
of textura intricata, hyphae cylindrical to slightly swollen, thin-
walled to thick-walled, hyaline to pale brown, often with pig-
mentedresinousexudatesatsepta.Ectal excipulum 70 –150
µm thick, of textura angularis, less often of textura prismatica.
Surface with warts up to 180 µm high, formed by fascicu-
late, short hyphoid hairs of globose to elongated cells, or of
textura globulosa-angulariswithsinglehyphoidhairs.Resinous
exudates often present on the surface, yellow to dark brown,
sometimesdissolvinginMLZ,turningreddishorintobrownish
yellowamberdrops,sometimeschangingcolourinKOH.Basal
mycelium of septate, straight hyphae, that frequently branch
andanastomose,turningyellowornotinKOH,oftencovered
withpigmented,small,resinousexudates.
 Ecology&Distribution—seeHansen&Olariaga(2015).
Key to species of Otidea
WewerenotabletostudyandinterpretthefollowingChinese
species and these are therefore not treated nor included in the
key: Otidea bicolor W.Y.Zhuang&ZhuL.Yang,O. kunmingen-
sisW.Y.Zhuang,O. olivaceobrunneaHarmaja, O. sinensisJ.Z.
Cao&L.Fan,O. subpurpureaW.Y.ZhuangandO. tianshuien-
sis J.Z.Cao,L.Fan&B.Liu.ForO. integra(Bres.)Harmaja
see notes under O. phlebophora.
1. Ascomata hypogeous, globose to subglobose, truffle-like
................................4.O. subterranea
1. Ascomata epigeous, cup-shaped to ear-shaped, split or
entire........................................ 2
2. Sporesornamented ............................. 3
2. Sporessmooth................................5
3. Sporeswithfinewarts±ridges.......... 17.O. unicisa
3. Sporesspiny..................................4
4. Spores14–17×7–9µm.............16.O. kaushalii
4. Spores16.5–20×7.6–10µm ......18.O. yunnanensis
5. SporesLm>17µm ............................6
5. SporesLm<17µm ...........................10
6. Apotheciabroadlyear-shaped,split;typicallyochraceous
yellowtoochreorange;oftenassociatedwithCudonia in
mossy Picea forests ..............11.O. cantharella
6. Apotheciacup-shaped,splitorentire;brown;underPicea
or other trees .................................7
7. Apotheciaentire;reddishbrown;basalmyceliumnormally
withabundantminuteresinousexudates;ectalexcipulum
hyphoidhairsoftenwithagelatinoussheath(moreeasily
seeninMLZ);withPicea ......... 12.O. propinquata
7. Apothecia split or entire; purple or ochre-brown; basal
myceliumwithoutorwithsparseresinousexudates;ectal
excipulumhyphoid hairs without aconspicuous sheath;
with angiosperms.............................. 8
8. Apotheciasplit,upto75mmwide;sporesbroadlyellipsoid
to oblong, Qm=1.7–1.8;associatedwithFagaceae ....
................................3.O. platyspora
8. Apotheciaentireorsplit,upto16mmdiam;sporesnarrowly
ellipsoid to fusoid, Qm=1.9–2.1;mostlikely associated
with Betulaceae and Salicaceae ..................9
9. Apotheciadeeplycup-shaped,split;ectalexcipulumwithout
resinous exudates................1.O. apophysata
9. Apotheciashallowlycup-shaped,usuallyentire;ectalex-
cipulum surface with abundant resinous exudates.....
................................. 2.O. daliensis
10. Receptaclesurfacewithbrightcitrineyellowtonesinyoung
apothecia ................................... 11
10. Receptaclesurfacewithoutcitrineyellowtones..... 15
11. Spores Qm=1.7– 2; apothecial base at most wrinkled,
without high ribs or veins ....................... 12
11. SporesQm=2– 2.3;apothecialbasewithribsorstrongly
veined at least in some apothecia ................14
12. Receptaclesurface ochraceousyellow;basal mycelium
yellowinKOH;sporesWm=6.5µm,Qm=1.7........
.................................25.O. borealis
12. Receptaclesurfacecitrineyellow;basalmyceliumunchang-
edinKOH;sporesWm=5.6–6µm,Qm=1.8– 2 ..... 13
13. Apotheciasometimesentire,sometimeswithbluntribsat
thebase;NorthAmerica..........31.O. oregonensis
13. Apotheciasplit,withoutribsatthebase;Europe.......
................................27.O. concinna
14. Apotheciamostlyentire,basestronglyribbed-veinedand
anastomosing in all the apothecia ...32.O. phlebophora
14. Apotheciamostlysplit,basewithafewribs-veins,smooth
in some apothecia.................... 30.O. minor
15. At least some paraphyses straight or curved, claviform
toalmostcapitateatapices;and/orresinousexudatesof
the ectal excipulum yellow or reddish yellow, turning bright
yellowinKOH;apotheciasplitornot............. 16
15. Paraphysesnotas such;resinousexudatesof theectal
excipulum,whenpresent,notturningbrightyellowinKOH;
apothecia always split.........................20
16. Apotheciashallowlycup-shapedandirregular,entire ...
...................................29.O. lactea
16. Apotheciaear-shapedordeeplycup-shaped,split...17
17. Paraphysesoftenclaviformorcapitateatapices,3–9µm
broad;sporesQm=1.6–1.8....................18
17. Paraphysesat mostslightly swollenatapices,2–5µm
broad;sporesQm=1.9–2 ......................19
18. BasalmyceliumturningyellowinKOH;Europe.......
................... (seeunderO. borealis)O. sp.‘a’

179
I.Olariagaetal.:AmonographofOtidea
18. BasalmyceliumnotyellowinKOH;NorthAmerica .....
..............................33.O. rainierensis
19. SporesLm=11.2–11.8µm;sometimesreceptaclewitha
bluishhue,ochraceousgreywhendried;basaltomentum
light ochre in dried specimens..26.O. caeruleopruinosa
19. SporesLm=10–10.6µm;receptacle without bluish hue
whenfresh,reddish brown whendried;basaltomentum
orange-ochre in dried specimens 28.O. flavidobrunneola
20. SporesLm < 12 µm ...........................21
20. SporesLm > 12 µm ...........................28
21. Apotheciadarkbrownwithlilaceoustones;Asia.......
................................23.O. purpurea
21. Apothecianot darkbrown,without lilaceoustones;Asia,
Europe or North America.......................22
22. Medullaryexcipulumwithreddishbrownresinousexudates
scatteredamongandcoveringsomehyphae;ectalexcipu-
lum of textura prismatica to textura intricata;receptaclesur-
face with contrasting brown warts .......6.O. papillata
22. Medullaryexcipulumsometimeswithyellowishbrownre-
sinousexudatesatsepta,notcoveringhyphae;ectalexci-
pulum of textura angularis;receptaclesurfacewithcontrast-
ing warts or not.............................. 23
23. Atleastsomewartshigherthan85µm;basaltomentum
orange-ochrewhendried;apothecialsectionsoftenyellow
inKOH;withoutresinousexudatesatseptainthemedullary
excipulum ....................... 9.O. tuomikoskii
23. Wartsupto85µm;basaltomentumpaleochreoryellow
whendried;apothecialsectionsnotyellowinKOH;some-
times with resinous exudates at septa in the medullary
excipulum ..................................24
24. Hymenium with distinct yellow or orange tones, ochre-
yellowtopinkishorange;resinousexudatesontheectal
excipulum partly dissolving into reddish grey, heterogene-
ousdropsinKOHorbasalmyceliumyellowinKOH. 25
24. Hymeniumwithoutorwith weakorange tones;resinous
exudates sometimes turning reddish brown but not dissol-
vingintodropsinKOH;basalmyceliumnotturningyellow
inKOH .....................................26
25. Resinousexudates ontheectalexcipulum partlydissol-
vingintoreddishgreyheterogeneousdropsinKOH;basal
myceliumnotturningyellowinKOH;NorthAmerica...
............................8.O. pseudoleporina
25. Resinousexudatesontheectalexcipulumnotdissolving
intoreddishgreyheterogeneousdropsinKOH;basalmyce-
liumturningyellowinKOH;Asia..... 19.O. brevispora
26. Hymeniumsometimeswithpinktones;apotheciayellowish
ochretobrown;narrowlyear-shapedinthebeginning. .
................................14.O. nannfeldtii
26. Hymeniumwithoutpinktones;apotheciareddishbrownto
orange-brown;broadlyear-shaped...............27
27. Spores Lm=10 –10.7 µm; Qm=1.6–1.7............
...............................13.O. formicarum
27. SporesLm=11.1–11.7µm;Qm=1.7–1.9............
............................15. O. subformicarum
28. SporesQm <1.8;ear-shaped................... 29
28. SporesQm >1.8;ear-orcup-shaped.............30
29. Apotheciadarkbrown,sometimeswitholivaceoustint;para-
physes mostly with distinct notches.10.O. brunneoparva
29. Apotheciacinnamonbrown,withoutolivaceoustint;para-
physes not or slightly notched.......... 7.O. leporina
30. Apotheciaochraceous yellow, hymeniumoftenwith pink
tones;basalmyceliumturningyellowinKOH.........
..................................22.O. onotica
30. Apotheciapaleor darkbrown,withoutpink tones;basal
myceliumnotturningyellowinKOH.............. 31
31. Receptaclemediumbrown;ectalexcipulumresinousexu-
datesabsentorscarce,lightyellowishbrown;basalmy-
celium without dark brown resinous exudates.........
...............................5.O. alutacea s.l.
31. Receptacledarkpurplebrown;ectalexcipulumresinous
exudatesabundant,darkbrown;basalmyceliumwithdark
brown resinous exudates ......................32
32. SporesQm <2, ellipsoid; apothecia mostly ear-shaped;
North America...................... 24.O. smithii
32. Spores Qm > 2, ellipsoid-fusoid; apothecia mostly cup-
shaped,split;EurasiaandNorthAmerica..........33
33. Receptaclestrikinglypurple-violaceous(fresh);medullary
excipulum without, or rarely with flabellate crystal-like exu-
dates,formingcross-likeaggregates;underconifersoncal-
careous ground....................21.O. mirabilis
33. Receptaclemostlywithoutpurpletones;medullaryexcipu-
lum with striate exudates covering some hyphae, some-
timesformingcrystal-likeaggregates;oftenunderdecidu-
ous trees or on acidic ground..........20.O. bufonia
Based on genealogical concordance phylogenetic species
recognition(GCPSR:Tayloret al. 2000),usingthefourloci,
RPB1,RPB2,EF1 and LSU rDNA, wedelimited25species
within Otidea (seeHansen&Olariaga2015). In addition eight
species were recognised by genetic divergence from their sis-
ters.Twenty-eightofthesearetreatedanddiscussedbelow,
along with O. brevispora, O. lactea, O. subterranea and O. yun-
nanensis included in our LSU phylogeny, and O. purpurea that
hasonlybeenstudied morphologically.Thespeciesarepre-
sented following their phylogenetic relationships, inferred from
ourcombined three-andfour-geneanalyses(f.3inHansen
&Olariaga2015).
Otidea platyspora clade
Apothecia disc-shaped, cup-shaped and split, or globose and
hypogeous, brown. Spores large, exceeding 20 µm, except
14–16.5µmifhypogeous.Basalmyceliumsmoothorwithvery
sparseresinousexudates.
Species — Otidea apophysata, O. daliensis, O. platyspora,
O. subterranea.
1. Otidea apophysata(Cooke&W.Phillips)Sacc.,Syll.Fung.
8:96.1889
Basionym.Peziza apophysataCooke&W.PhillipsinCooke,Grevillea 5:
60.1876.
 ≡Pseudotis apophysata (Cooke&W.Phillips)Boud.,Hist.Classific.Disco-
myc.Europe:52.1907.
Holotype.England,Shrewsbury,inadampditch,1876,W. Phillips(K(M)
30410exHerb.Phillips).Isotype(K(M)167215exHerb.Cooke)!
Misapplied names
– Otidea felinasensuBoudier,Icon.Mycol. livr.29:n°.512,pl. 331.1910
(preliminarytextwith‘circulaires’).
Apotheciasolitarytocaespitose,8–30mmhigh,upto15mm
wide, initially ear-shaped, then soon expanding and becom-
ingcup-shaped, split, sessileor stipitate. Hymenium purple
brown(6D3,6D4),whendrieddarkorangebrown(6E7,6F7).
Receptacle surface palegreyishorange(6C5),purplebrown
(6D4),whendrieddarkorangebrown(6E7,6F7),furfuraceous.
Wartsscarcetoabsent,low.Stipeabsent.Basal tomentum and
mycelium whitishtopaleorangegrey(5B2). Spores narrowly el-
lipsoid to fusoid, narrowing toward the poles, sometimes inequi-
lateral, with two large guttules, and often with several smaller
guttules,smooth,hyaline,20– 24.5×9–11µm(Lm=21.6µm,
Wm = 10 µm, Qm=2.1;n=1).Paraphyses curved to hooked,
seldomstraight,slightlyenlargedatapices,3–4(–5)µmwide,

180 Persoonia–Volume35,2015
sometimeswithasinuousundersideorwith1–2notches,fre-
quently branching, entangled and interconnected, when dried
containingsmall,refractive,hyalineguttules.Asci172 –197×
12–13µm. Apothecial section700–850µmthick. Medullary
excipulum of textura intricata,400 – 500 µm thick, hyphae
slightlythick-walled,5 – 9µmwide,palebrown.Ectal excipulum
of textura angularis,70–110µm,cellsthin-walled,yellowish
brown,18–33×13– 22µm.Surfacewithlowwarts,upto40
µmhigh,cellsovoidtoglobose,constrictedatsepta,8–13µm
wide.Resinousexudatesabsent.Basal myceliumof4–5µm
broad, pale brown hyphae, sometimes with oily refractive drops
onthesurface.
Specimens examined. gErmany,Nordrhein,Herten,1Sept.1999,F. Kas-
parek,privateherb.Kaspareks.n.(dupl.S-F257062).
Notes — Otidea apophysata is characterised by deeply cup-
shaped, split, brown apothecia, and large, ellipsoid to fusoid
spores.Otidea daliensis is a closely related species, distin-
guished by darker brown, shallowly cup-shaped, usually entire
apothecia, and abundant dark brown resinous exudates on the
outermostcellsoftheectalexcipulum.Otidea platyspora has
also brown apothecia and large spores, but it differs from O. apo-
physata in the larger apothecia, partly buried in the substrate,
broadly ellipsoid to oblong spores, and non-entangled para-
physeswithoutnotches.
ThenameO. apophysata has been misapplied twice for O. da-
liensis (Boudier 1909b, Pérez-Butrón & Fernández-Vicente
2008).ThetypematerialofO. apophysata lacks resinous exu-
dates on the ectal excipulum, which clearly distinguishes it
from O. daliensis.An originalpaintingbyW.Phillips,based
on the type material of O. apophysataandpreservedatRBG
Kew(reproducedinParslow&Spooner2013),showstypical
brown,split,deeplycup-shapedapothecia.Theillustrationby
Cooke (1878, f. 350), based on drawings and (likely dried)
specimenscommunicatedbyW.Phillips,showsslightlydarker
apotheciathantypical.
Otidea apophysata is only known from very few reports from
France(Boudier 1910 as O. felina), Germany (Häffner &
Winterhoff1989, Kasparek 2000), Belgiumand Spain (Van
Vooren2011a).InMidtoSouthBritainitiswidelydistributed,
with collections from fourteen different localities (Parslow &
Spooner2013).UnlikemostOtidea species, O. apophysata
shows preference for damp habitats, and might be associated
with Alnus and Populus (Häffner&Winterhoff1989,Parslow
&Spooner2013).
2. Otidea daliensis W.Y.Zhuang&Korf,Mycotaxon35:300.
1989
Holotype.China,Yunnan,Dali,HudiequanPark,alt.2100m,onbaresoil
under seedlings of Plantago major,5Nov.1988,R.P. Korf, L.S. Wang & W.Y.
Zhuang(HMAS57688).Isotype(CUP-CH2532).
Misapplied names
– Pseudotis apophysatasensuBoudier,Icon.Mycol. livr.24:n°.471,pl.332.
1909(preliminarytextwith‘circulaires’).
– Otidea apophysatasensuPérez-Butrón&Fernández-Vicente,Errotari5:
37.2008.
Apotheciagregarious,upto9mmhigh,3–16mmwide,initially
cup-shaped, sometimes split, then becoming shallowly cup-
shaped,sessileorstipitate.Hymenium when dried dark purple
brown (7F5, 7F6) to dark brown (6F5). Receptacle surface
darkpurplebrown(7F5,7F6),whendrieddark brown(6F5),
furfuraceous.Wartsabsentor verylow.Stipe if present very
short.Basal tomentum and mycelium whitish to pale orange
grey(5B2).Spores broadly ellipsoid to ellipsoid and narrowing
toward the poles, sometimes inequilateral, with two large gut-
tules, and often with several smaller granules, smooth, hyaline,
(19.5–)20.5 – 23 × 10.5–12(–13) µm (Lm=21.2–21.5µm,
Wm=10.7–11µm,Qm=1.9– 2.1;n=2).Paraphyses curved to
hooked,sometimesslightlyenlargedatapices,2.5 – 3.5(–5)µm
wide, sometimes with slightly swollen areas, apices sometimes
embedded in a brown matter, when dried containing small,
refractive,brownguttules.Asci199–212×15 –17µm.Apothe-
cial section600 – 850µmthick.Subhymeniumc.90–110µm
thick, of dense textura intricata,visibleasadarkerzone,cells
cylindrical to swollen, with scattered brown resinous exudates
atsepta.Medullary excipulum of textura intricata,300–400µm
thick,hyphae4 – 6.5µmwide,slightlythick-walled,palebrown,
with brown resinous exudates at septa. Ectal excipulum of
textura angularis,90–120µmthick,cellsthin-walled,yellowish
brown,18–28×11–28µm.Surfacewithbroadlyconicalwarts.
Non-wartedpartswith2– 5-celledhyphoidhairs,withclaviform
uppermost cell, more rarely cylindrical, constricted at septa,
6–9µmwide.Resinousexudatesabundant,darkbrown,partly
dissolvinginMLZ.Basal mycelium of3.5–4µmwide,hyaline
hyphae,withyellowishbrown,small,resinousexudates.
Specimens examined. FranCE,1869,L. Quélet(UPSF-629790).–Spain,
BasqueCountry,Bizkaia, Galdames, Presa deAguas Juntas, sandy soil
under Populus nigra,11Aug.2003,J.L. Pérez Butrón,SEST-03071103;17
Aug.2006,SEST-06081702.
Notes — Otidea daliensis is recognised by small, usually
entire, dark purple brown, shallowly cup-shaped apothecia,
large ellipsoid spores often narrowing toward the poles, and
abundantbrownresinousexudatesontheectalexcipulum.This
species has been confused with O. apophysata(seeO. apo-
physata).
ThefirstknownreportofO. daliensis wasbyBoudier(1909b),
as Pseudotis apophysata.Theplate332(n°.471)showsthe
typical dark purple, shallowly cup-shaped apothecia, in contrast
to O. apophysata,depictedinplate331(n°.512)asO. felina
(Boudier1910).Mornand&Courtecuisse(2005)proposed a
provisional name, O. boudieri, for the Boudier P. apophysata
plate(=O. daliensis,n°.332).Zhuang&Korf(1989)described
O. daliensis without comparing it to O. apophysata.Material with
small, shallowly cup-shaped apothecia and darker colour was
reported from the Iberian Peninsula as O. apophysata (Pérez-
Butrón&Fernández-Vicente2008),andthushadsimilarities
with O. daliensis andBoudier’splate332.VanVooren(2011a)
considered the Iberian O. apophysata material to represent
O. daliensis.After restudying the Iberian materialand com-
paring it to O. apophysata,weagreewiththatstatement.LSU
sequences obtained from the Iberian material and from the
Chinese holotype of O. daliensis areidentical.
3. Otidea platysporaNannf.,Ann.Bot.Fenn.3:317.1966.
—Fig.7
Holotype.SwEdEn,Uppland,Djursholm,Oct.1951,A. Zander, FungiExs.
Suec.3284(UPSF-005428).Isotype(S-F88395)!
Misapplied names
– Otidea cochleatasensuBoudier,Icon.Mycol. livr.21:n°.461,pl.329.1908
(preliminarytextwith‘circulaires’).
Apotheciacaespitose,60–70mmhigh,40 –75mmwide,ini-
tially ear-shaped, then soon expanding and becoming deeply
cup-shaped,split,sessileorshortlystipitate.Hymenium initially
yellowishbrown(5C6,5C7),palegreyishbrown(5C3,5C4)to
darkbrown(6F3–6F6),whenbruisedmarginblackish,when
driedbrownishochre(5B4,5B5).Receptacle surface dark ochre
brown(5D7,5D8),slightlyhygrophanous,indryingpaleochre
brown(5B6),whendriedyellowishbrown(5C6,5C7),some-
timeswrinkledatthebase,finelyfurfuraceousinthemargin.
Wartsabsent.Stipenotwelldeveloped.Basal tomentum and
myceliumwhitishtopalebrown(5A3). Spores broadly ellipsoid

181
I.Olariagaetal.:AmonographofOtidea
Fig. 7 Otidea platyspora*.a.Apothecia;b.spores;c.paraphyses;d.ectalexcipulum(a:KH.10.183;b – d:KH.09.163).—Scalebars=10µm;*=allfresh
material.
to oblong, rarely slightly inequilateral, with two large guttules,
oftenwithseveralsmallerguttules,smooth,hyaline,18–22×
(9.5–)10.5–12µm(Lm=19.8–20.7 µm,Wm=10.9–11.6µm,
Qm=1.7–1.8; n=6). Paraphyses curved to hooked, of the
same width or slightly enlarged at apices, 2.5 – 3.5(– 5) µm
wide, without notches, rarely with a slightly swollen area on
the underside, when fresh containing small, refractive, light
brownishyellowguttules;whendriedtiny,lightyellowgranules.
Asci168–213×14–19µm.Apothecial section700 –1700µm
thick.Subhymenium80 –100µmthick,visibleasadarkerzone,
cells cylindrical, densely arranged, with scarce yellowish brown
resinousexudates at septa.Medullary excipulum of textura
intricata,550–1100µmthick,hyphae3.5–10µmwide,some-
timesslightlyswollen,thin-walled,lightyellowishbrown.Ectal
excipulum of textura angularis,70–90µm, cells thin-walled,
palebrown,13 – 37×12–22µm.Surfacewithhyphoidhairs,
33–70µmlong,of4–7ovoidtosubglobosecells,constricted
atsepta,6 – 9µmwide,sometimeswithalightbrownmatter.
Resinousexudates absent toscarce,palebrown, dissolving
inMLZ.Basal mycelium of3.5 – 4.5µmwide,hyalinetovery
pale brown hyphae, sometimes with oily refractive drops on the
surface,sometimeswithminutebipyramidalresinousexudates.
Specimens examined. azErbaijan,MontesTalysh,insilvamixta,14Oct.
1962,E. Parmasto(UPSF-629452).–dEnmark,EastJylland,Kammerher-
rensEge,MoesgårdSkov,SofÅrhus,under Fagus and Quercus,11Sept.
2006,J. Vesterholt,JV06-656(C).–FranCE, Ain, commune de Saint-Benoit,
forêtd´Évieu,underQuercus robur, Carpinus betulus, Corylus avellana and
Alnus glutinosa,15July2011,F. Armada, NV2011.07.04(dupl.S).–SwE-
dEn,Uppland, Stockholm,Drottningsholm,underQuercus robur,13 Sept.
2009,K. Hansen & I. Olariaga, KH.09.163(S);Uppland,Stockholm,Solna,
Karlbergsparken,gardenwithbroadleaftrees,29July2006,H.-G. Toresson
(S-F248339);Uppland, Stockholm, Sånga,Svartsjöslott, underQuercus,
Fagus and Corylus,31Aug. 2008,H. Kauffman, HK08046(S); Uppland,
Uppsala, mixed forest with Pinus sylvestris, Picea abies, Betula pendula
and Quercus robur,9Oct.2011,J.C. Zamora (BIO-Fungi16391);Uppland,
Uppsala,Hågadalen-NåstenNatureReserve,Predikstolen,underCorylus,
Populus, Quercus and Picea, onrich ground,6Sept. 2010,K. Hansen &
I. Olariaga, KH.10.183(S).
Notes — Otidea platyspora is characterised by large, caespi-
tose, brown apothecia, blackening in bruised margins and large
spores.MacroscopicallyitresemblesmembersoftheO. alu-
taceacomplex,butthesedifferinthesmallerspores.Otidea apo-
physata and O. daliensis have likewise brown apothecia and
large spores, but can be distinguished by having even larger
sporesandsmaller(upto30mm)apothecia(seefurtherunder
thosespecies).
Otidea platyspora is a striking species with scarce records.
It was described from Sweden and has been reported from
France(Boudier1908,Nannfeldt1966,VanVooren&Armada
2011),theNetherlands(MaasGeesteranus1967)andrecently
fromBritainandDenmark(Parslow&Spooner2013).Were-
portitasnewfromAzerbaijanandgiveadditionalrecordsfrom
DenmarkandSweden.Otidea platyspora appears to have been
overlookedandismorewidespreadthanthoughtpreviously.
4. Otidea subterranea Healy&M.E. Sm.inSmith&Healy,
Mycol.Res.113:860.2009
Holotype.USA,Iowa,LedgesStatePark,shallowlyhypogeous,erumpent
onsoil,30Aug.1997,R. Healy RH69(FH).
Notes — Otidea subterranea is the only known hypogeous
species of Otidea.Thepustules(c.50 –100µmhigh)on the

182 Persoonia–Volume35,2015
outer surface of the ptychothecia and the incrusted tomentum
are typical for Otidea.Theasciarecylindrical,with8uniseri-
atespores,placedinadefinedhymenium.UnlikemostOtidea
speciesthesporesareuniguttulate.Asanadaptationtoahypo-
geoushabitactivesporedischargehasbeenlost.Probablyfor
the same reason, the paraphyses are aggregated and fused
subapically to form an epithecium of brown thick-walled cells,
and are reminiscent of the frequently branching, entangled and
interconnected paraphyses in O. apophysata. There are no
other morphological features that support the exact placement
of O. subterranea in the O. platysporaclade.Thereceptacle
surface of O. subterranea ascomata is whitish to cream in
young and peach-cream to buff with tan-brown areas in older
specimens.Theglebaisofdarkbrownfertileveinslinedwith
a thin, light yellowish brown hypothecium, but the sterile veins
areofwhitishhyphae(Smith&Healy2009).
Otidea alutacea clade
Apotheciacup-shapedandsplit,brown.Sporestypicallyellip-
soid,withalmostparallelsides.Outermostectalexcipulumand
basalmyceliumsmoothorwithverysparseresinousexudates.
Species — Otidea alutacea s.l.
5. Otidea alutacea(Pers.)Massee,Brit.Fungus-Fl.4: 446.
1895.—Fig.8
Basionym.Peziza alutaceaPers.,Observ.Mycol.2:78.1799;non Peziza
alutacea Schumach.,Enum.Pl. 2:431.1803(homonym).
 ≡Scodellina alutacea (Pers.)Gray,Nat.Arr.Brit.Pl.1:668.1821.
 ≡Peziza cochleata var.alutacea(Pers.)Fr.,Syst.Mycol. 2:50. 1822:
Fr.loc.cit.(‘ßalutacea’).
 ≡Plicaria alutacea (Pers.)Fuckel,Jahrb.NassauischenVereinsNaturk.
23– 24:327.1870.
 ≡Aleuria alutacea (Pers.)Gillet,Champ.FranceDiscom.1:42.1879.
Lectotype.Bull.,Hist.Champ.France1:t.154,f.b.,designatedbyCar-
bone(2010a).Epitype.(L0111551,Herb.Persoon),designatedbyCarbone
(2010a).
1Apothecia gregarious, rarely caespitose, 15 –75 mm high,
8–48mmwide,initiallyear-shaped,soonexpanding,becom-
ingshallowlytodeeplycup-shaped,split,sessileorstipitate.
Hymeniuminitiallybrown(5D6), then yellowish brown (5D4,
5D5) to dark reddish brown (7D7, 7E7), when dried purple
brown(6D6,6D7).Receptacle surface slightly hygrophanous, in
dryingyellowishbrown(5D5,5D6),orsometimeswithpurplish
brown(7D5)tones,whendriedlightochre(5A4,5B4)brownish
ochre(5B5,5B6),finelyfurfuraceoustoslightlywartyinthe
margin.Wartsflattened,gregarious,concolorous,darkerwhen
theoutsideindrying.Stipe3–6×3–4mm.Basal tomentum
and mycelium whitetoverylightochre(5A2).Spores ellipsoid
to broadly ellipsoid, oblong ellipsoid, slightly inequilateral,
with two large guttules, often with several smaller granules,
smooth,hyaline,(13.5–)14.5–16.5(–17.5)×6.5–7.5(–8)µm
(Lm=14.6–16.1µm,Wm=6.6–7.5µm,Qm=2.1–2.2;n=5).
Paraphyses curved to hooked, only few straight, of the same
widthorslightlyenlargedatapices,2.5– 4.5µmwide,without
notches, sometimes embedded in a brown matter at apices,
when fresh containing small, refractive, light brownish yellow
guttules;whendriedbrownishyellow.Asci 140–187×11–13
µm. Apothecial section 750 –1050 µm thick. Subhymenium
c.80–100µmthick,visibleasadarkerzone,cellscylindrical
to swollen, densely arranged, with scattered yellowish brown
resinousexudatesatthesepta.Medullary excipulum of textura
intricata,400–650µmthick,hyphaethin-walledtoslightlythick-
walled,4 – 9µmwide,hyalinetolightbrown,sometimeswith
yellowishbrownresinousexudatesatsepta.Ectal excipulum of
textura angularisof80–100µm,cellsthin-walled,palebrown,
9–21×9–17µm.Surfacewith broad conical warts, 35 – 57
µmhigh,formed byshort,fasciculate,hyphoidhairs, of 6–7
globose to subglobose cells, constricted at septa, 7–10 µm
wide.Resinousexudatesabsenttoscarce,yellowish brown,
dissolvinginMLZ.Basal myceliumof3– 4.5µmwide,hyaline
hyphae, with oily refractive drops on the surface, sometimes
withminuteresinousexudates.
Specimens examined. O. alutacea s.str. — dEnmark,SSjælland,Møn,
StoreKlinteskov,bySvantestenen,oncalcareoussoil(pH7.0)alongforest
road, under deciduous trees, together with Humaria hemisphaerica and
Trichophaea woolhopeia,11 Sept. 1994, K. Hansen & S.K. Sandal, KS-
94-111 (C).– FranCE,Puy-de-Dôme,Auvergne,Nadayat,sousfeuillusen
terrain neutrocline, 20Sept.1998,G. Corriol,GC98092002(dupl.S).–italy,
Piemonte,Vignole Borbera(AL),Fraz. Varianosuperiore, underQuercus
pubescens and Castanea sativa,19Oct.2010,M. Carbone(S-F257084).–
norway,Nord-Trøndelag,Leksvik,Gjøråsvika,onrich,bareground,under
Corylus and Picea,onasteepslope,3Sept.2009,K. Hansen & I. Olariaga,
KH.09.133(S).–Spain,Gipuzkoa,Tolosa,Elosegimarkesarenlorategiak,
underbroadleaf treesina garden, 29May 2009, J.I. López-Amiano,JLA
2009052902(ARAN-FungiA3023204). –SwEdEn, Gotland,OllajvsNature
Reserve,close toLjugarn,under Picea and Pinus on calcareous ground,
27Sept.2010,K. Hansen, K. Gillen & I. Olariaga,KH.10.278(S).Clade 1
— dEnmark,Eastern Falster,Korselitze-forests,5Oct.2007,H. Knudsen,
KH.07.46 (S). – SwEdEn, Uppland, Stockholm, Norra Järvafältet, Hansta
NatureReserve,onrichgroundunderCorylus and Quercus,8Sept.2010,
K. Hansen, K. Gillen & I. Olariaga,KH.10.193(S);Uppland,Stockholm,
NDjurgården,Storaskuggan,onsoilin grazedopenoakforest,12Sept.
2008,J. Santos, JS.08.81(S);Uppland,Uppsala,Hågadalen-NåstenNature
Reserve,Predikstolen,underQuercus robur, Picea abies, Corylus and Salix,
onrichbareground,17Sept.2009,K. Hansen & I. Olariaga,KH.09.170(S).
Clade 2 —USA,Oregon,LincolnCo., Devil’s Punchbowl State Park, 13
Mar.1997,E.T. Peterson (OSC56770);Washington,PierceCo.,MtRainier
NationalPark,LowerTahomaCreek,underPseudotsuga, Tsuga, Picea and
Calocedrus, 29Oct.1996,E.T. Peterson(OSC56747);ibid.,30Oct.1996
(OSC56754); ibid., 18Oct.1997 (OSC 56798);Washington,Snohomish
Co.,SloanCreektrail,24Sept.1997,E.T. Peterson(OSC56777).Clade 3a
— norway,Nord-Trøndelag,Leksvik,Gjøråsvika,onslopeunderCorylus and
Picea, onrichground,3Sept.2009,V. Kučera & I. Kautmanová,KH.09.135
(S).–SwEdEn,Södermanland,Nynäshamn,Herrhamra,onsoilunderFagus,
innarrowforestareaalongtheroad,19Sept.2013,K. Hansen & X.H. Wang,
KH.13.50(S); Uppland, Norrtälje, Länna, under Corylus, 26Aug.2008,
J. Santos, JS.08.43 (S); Uppland, Uppsala, Hågadalen-Nåsten Nature
Reserve,Predikstolen, under Quercus robur, Picea abies, Corylus and
Salix,onrich,bareground,6Sept.2010,K. Hansen, K. Gillen & I. Olariaga,
KH.10.198(S). Clade 3b — dEnmark,SSjælland,Møn, StoreKlinteskov,
VestreUlvemose, on calcareoussoilin deciduous forest, 26 Sept.1994,
K. Hansen & S.K. Sandal,KS-94-192(C).– SwEdEn, Uppland, Uppsala,
Hågadalen-NåstenNatureReserve,Predikstolen,underCorylus, Populus
and Picea,onrichground,19Sept.2009,K. Hansen & I. Olariaga, KH.09.178
(S).Clade 4 — USA,Oregon, BentonCo.,Corvallis,McDonald-DunnRe-
searchForest,10Oct.1997,E.T. Peterson(OSC56782);ibid.,18Nov.1996
(OSC56758);Oregon,BentonCo.,Corvallis,westsideofNWBeechwood
Place, scattered to clustered on rotting bark mulch and in thin grass under
Pseudo tsuga menziesii,17 Nov. 2010, N.S. Weber,NSW10200 (OSC
150345);Oregon,Corvallis,WithamHill,25Nov.1997,E.T. Peterson (OSC
56813);Oregon,DouglasCo.,20Oct.2010,J. Moore,Moorefun19(OSC);
Oregon,DouglasCo.,BearGulch,underPseudotsuga menziesii,13Jan.
1999,R. Davidson (OSC67524);Oregon,DouglasCo.,Slimewater,under
Pseudotsuga menziesii, Quercus garryana, Abies grandis, etc., 12 Sept.
1999,Frymire(OSC72978);Oregon,DouglasCo.,Umpqua,underPseudo-
tsuga menziesii, Abies concolor, Calocedrus decurrens, Corylus cornuta,
etc.,23Mar.2000,E. Stewart(OSC72979);underPseudotsuga menziesii,
Pinus ponderosa, Calocedrus decurrens, Pinus lambertiana, Abies concolor,
etc.,15Dec.1999,C. Rusch(OSC72176);Oregon,LaneCo.,Willamette
NationalForest,MiddleForkRangerDistrict,underTsuga heterophylla, Thuja
plicata and Abies grandis,18Nov.2002,Smith(OSC119567).
Additional material of O. alutacea s.l.dEnmark,Sjælland,Hareskoven,Nof
Copenhagen,2Aug.1961,H. Dissing(C-F-48301).–FranCE, Orliénas,sous
Quercus et Cedrus atlantica,13Nov.2008,B. Rivoire & N. Van Vooren,NV
2008.11.01(dupl.S-F256976);Rhône,Bron,ParcdeParilly,24Sept.2008,
J. Cavet,NV2008.09.32(dupl.S-F256974); Rhône,Courzieu,hameaudes
Verchères,sousPseudotsuga menziesii,26Oct.2008,D. Carbonnel,NV
2008.10.02(dupl.S-F256975).–italy,Puglia,Mesagne(BR),BoscoLucci,
in soil, mainly under Quercus ilex,12Oct.2010,M. Carbone (S-F257085).
–norway,Nord-Trøndelag,Leksvik,Gjøråsvika,mixedforestonrichground,
1 DescriptionbasedonlyonspecimensofO. alutaceas.str.

183
I.Olariagaetal.:AmonographofOtidea
Fig. 8 Otidea alutacea s.str.(KH.09.133).a.Apothecia; b.spores inwater †;c.paraphysesinwater†;d.ectalexcipulum inKOH†.—Scale bars=10µm;
†=driedmaterial.
3Sept.2009,K. Hansen & I. Olariaga,KH.09.137(S);ibid.,R. Braathen,
KH.09.139(S);Nordland,Rana,Rausandaksla,insalico-betuletum,onlime-
stone,21Sept.1974,S. Sivertsen(C-F-60697). –Spain,Madrid,Arboreto
deETSIMontes,underQuercus suber, Pinus pinea and Nerium oleander,
21Nov.2006,L. Rubio Casas(AH42204).–SwEdEn,Gotland,nearVisby,
Värnhem,onrichgroundunderFagus and Quercus, with Hepatica nobilis,
22Sept.2009,E. Bohus-Jensen, K. Hansen & I. Olariaga,KH.09.187(S);
Gästrikland,Hofors,Sibbersbovägen,onrichgroundunderCorylus,1Sept.
2010, K. Hansen, K. Gillen & I. Olariaga,KH.10.154(S);Jämtland,Ändsjön
NatureReserve,inrichPicea forest,26Aug.2009,H. Lindström, KH.09.97
(S);Närke, Ekeby,Kvarntorp, underabig Quercusbythe road,10Sept.
2008,J. Santos & K. Hansen,JS.08.57(S);Närke,Tysslinge,Latorpsbruk,
Grytsätterskogen,grasslandwith Quercus, 13 Sept. 2008, JS.08.76 (S);
Närke,Örebro,Hästhagen,bySvartån,mixedforest,13Sept.2008,H. Kauff-
man,JS.08.74(S);Skåne,Helsingborg,FredriksdalsFriluftmuseum,16Sept.
2010, G. Hamilton,KH.10.206(S);Skåne,KjugekullNatureReserve,onbare
ground under Quercus rubra, Corylus and Fagus,24Sept.2010,K. Hansen,
K. Gillen & I. Olariaga,KH.10.262(S);Skåne,Maltesholm,forestclosetothe
castle, on the ground under Fagus, close to Alnus,25Sept.2010,K. Hansen,
K. Gillen & I. Olariaga, KH.10.265(S);TorneLappmark,Abisko,12Aug.
1974,M.D. Paulsen & N. Tams(C-F-48045);Uppland,Stockholm,Eneby-
berg,Rinkebyskogen,on bare soil in a ditch, under Picea, Betula and
Populus,28Aug.2008,J. Santos,JS.08.50(S);ibid.,indeciduousforest
under Corylus, but also Tilia, Quercus and Betula,1Sept.2008,J. Santos,
JS.08.56(S);Uppland,Stockholm,NorraJärvafältet,HanstaNatureReserve,
on naked soil among leaves, under large Corylus, also Quercus,8Sept.
2010, K. Hansen, K. Gillen & I. Olariaga,KH.10.189(S);Uppland,Uppsala,
Hågadalen-NåstenNature Reserve, Predikstolen, on rich ground under
Quercus, Corylus, Populus tremula and Picea,17Sept.2009,K. Hansen &
I. Olariaga,KH.09.173(S).
 Notes—WeconsiderO. alutaceas.l.tocompriseaspecies
complex.Itisrecognisedbythemediumbrown,cup-shaped,
split apothecia, an ectal excipulum with only sparse resinous
exudatesifany,andpredominantly oblong spores.Although
sometimestreated as awell-delimited species (Harmaja
2009a),sporesizesofO. alutacea provided by different authors
varyconsiderably,e.g.14–16×7–9µm(Dissing2000)or12.5–
14.5×6.2–7.3µm(Harmaja2009a).Infact,O. cochleata has
been separated from O. alutacea on account of larger spores,
16–18×7– 8µm(Dissing2000),ordarkerapothecia(Mornand
& Courtecuisse 2005, Liu & Zhuang 2006, Zhuang 2006).
Meanwhile,twotaxaoftheO. alutacea species complex have
been separated in North America, based on apothecia colour,
andsporesizeandshape(Peterson1998).OurLSUphylogeny
resolved several clades within O. alutacea s.l.(Fig.1),which
arestronglysupported in our multigene phylogeny(Hansen
&Olariaga2015).Itappearsthatthesporesizeswithineach
clade have a fairly narrow range, but overlap exists between
theclades.Patternsofcontinentalspeciationaresuggestedas
well;twocladeshaveNorthAmericanspecimens(clade2,4),
andtherestcontainsamplesfromEuropeandAsia.
Carbone(2010a)selectedalectotypeandanepitypeforO. alu-
tacea.Asporerangeof15.5–17×7µmwasgivenfortheepi-
typespecimen(Carbone2010a),andbasedonthisweassign
O. alutaceas.str.tothecladeinferred fromthephylogenetic
analysesencompassingthissporesize.Thedescriptionabove
is based solely on the specimens of that clade, which are
characterised by the initially shallowly cup-shaped apothecia,
with rather light ochraceous brown hymenium, which later
becomes purple brown and more deeply cup-shaped. The
lectotypificationofO. alutacea proposed by Parslow & Spooner
(2013)issuperfluous.
Among the Eurasian clades, clade 1 contains North European
specimenscharacterisedbysmallspores(12–13.5×5.5–7
µm, Lm=12.2–12.9µm, Wm=5.8 –6.6 µm, Qm=1.9–2.2),
non-overlapping with O. alutaceas.str.andclade3.Clade 1

184 Persoonia–Volume35,2015
should be compared to O. kunmingensis, a taxon belonging
to the O. alutacea complex characterised by short spores
(Zhuang&Yang2008).Clade 3aencompassesthree speci-
menswithsporessizes13.5–15×6.5 –8µm(Lm=14.5–14.6
µm,Wm=6.7–7.3 µm, Qm=2 –2.2) overlapping with those
of O. alutaceas.str.asdescribedhere.Theapothecia differ
slightly macroscopically from O. alutaceas.str.inbeingdeeply
cup-shaped in the beginning, reddish brown when young, later
paleochre-brown. Clade 3b is composedof two collections
with larger spores, 15.5 –17.5 × 7.5– 8 µm (Lm=16.1–17.4
µm,Wm=7.7–8µm,Qm=2 – 2.3),butslightlyoverlappingwith
O. alutaceas.str.andclade3a.Clade3bmaycorrespondto
O. cochleata sensuDissing(2000).The twoNorthAmerican
clades(2and 4) comprise specimenswithclearlynon-over-
lapping spore sizes, 15 –18 × 7–8 µm (Lm=15.5 –16.9 µm,
Wm=7.3 –7.9 µm, Qm=2– 2.2) and 12 –14.5 × 6.5 –8.5 µm
(Lm=12.5–14µm,Wm=6.6–7.5µm,Qm=1.8–2),respectively.
Thesetwocladescorrespondtothetwospeciesdistinguished
byPeterson(1998)inWesternNorthAmerica,asO. alutacea
(clade2) and O. umbrina (clade4).Thesespeciesweresaidto
differincolouroffreshapothecia.Clades2and4havespores
that overlap with European clades, and it is so far problematic
to distinguish them using only morphology and disregarding
thegeographicalorigin.
TobeabletofullyclarifyspeciesboundarieswithintheO. aluta-
cea complex, sampling of additional collections for molecular
studyisneeded.Distinguishingmorphologicalandecological
characters should be sought, especially through studying fresh
Fig. 9 Otidea papillata†.a.Apothecia;b.sporesinwater;c.paraphyses;d.resinousexudatesinthemedullaryexcipuluminwater;e.ectalexcipuluminKOH;
f.basalmyceliuminKOH(a– d,f:H6003547,holotype;e:TUR102134).—Scalebars=10µm;†=alldriedmaterial.—Photos:a.J.Kearey.

185
I.Olariagaetal.:AmonographofOtidea
material.Severalnamesthatbelongtothiscomplex,suchas
O. alba, O. cinerascens, O. cochleata, O. felina and O. kun-
mingensisshouldbeconsideredasthisstudyisundertaken.
Otidea papillata clade
Apotheciacup-shaped,split.Receptaclesurfacewithcontrast-
ingwarts.Sporessmall,9.5 –11µm.Medullaryexcipulumwith
brownresinousexudatesembeddingsomehyphae.Ectalexci-
pulum poorly differentiated, of textura prismatica to textura
intricata.Resinousexudates on the ectalexcipulumnotdis-
solvinginMLZ.
Species — Otidea papillata.
6. Otidea papillataHarmaja,Karstenia15:31.1976—Fig.9
Holotype.Finland,Kainuu,Paltamo,Melalahti,Myllymäki,predominantly
coniferous grass-herb forest on distinctly calcareous soil, in litter mainly
composedofspruceneedles,23Sept.1971,H. Harmaja (H6003547)!
Apothecia23–30mmhigh,7–33mmwide,initiallybroadlyear-
shaped, with upper margin rounded, then becoming cup-shaped,
split,stipitate orsessile.Hymeniumochre(5B5,5B6)toyel-
lowishochre(4A5)whendried.Receptacle surface yellowish
brown(5C6,5C7)whendried,warty.Wartsconical,angularor
rounded, gregarious, distinctly darker than the background, dark
ochrebrowntobrown.Stipe7–10×3–4mm.Basal to mentum
and myceliumabundant,palebrownishochre(5A3)toorange-
ochre(6A3). Spores broadly ellipsoid, seldom very slightly
inequilateral,with two largeguttules, smooth, hyaline,9.5–
11(–11.5)×5.5–6.5µm(Lm=10–10.7µm,Wm=6.1– 6.3µm,
Qm=1.6–1.7;n=2).Paraphyses curved to tightly hooked, usu-
allyenlarged at apices, 3–4µm wide, sometimes with1–2
shallow notches, sometimes truncate or forked at apices,
whendriedcontainingsmall,refractive,hyalinegranules.Asci
116–165×9 –10.5µm.Apothecial section900 –1200µmthick.
Subhymenium 70 –90 µm thick, of dense textura intricata,
visibleasapalebrownzone.Medullary excipulum of textura
intricata,600–800µmthick,hyphae5.5 –10.5µmwide,thin-
walled to slightly thick-walled, hyaline to very pale yellow, with
brown resinous exudates scattered among and covering some
hyphae, sometimes rod-shaped, paler and partially dissolv-
inginKOH.Ectal excipulum of textura prismatica to textura
intricata,70–100µm,cellsthin-walled,hyaline,13–35×9–18
µm.Surfacewithconicaltoroundedwarts,65 –100µmhigh,
formed by short, fasciculate, hyphoid hairs, sometimes with
agelatinoussheath.Resinousexudatesabundant,yellowish
browntoreddishbrown,palerinKOH,notdissolvinginMLZ.
Basal mycelium of 2.5 –4.5 (– 6) µm wide, septate, hyaline
to very pale yellow hyphae, unchanged in KOH, smooth or
with regularly arranged, spheroid, yellow, resinous exudates,
sometimesembeddedinayellowishmatter,dissolvinginMLZ,
partiallyandmoreslowlyinKOH.
Specimen examined. Finland, Varsinais-Suomi,Parainen,Petteby,Stor-
näset(Paltbacken),inconiferousforestamongmosses,30Sept.1990,T. Lind-
holm (TUR102134).
Notes — Otidea papillata is only known from two Finnish
collections and its apothecial colours in fresh state are still
unknown.Itisadistinctspecies,characterisedbycup-shaped
apothecia with conspicuous warts on the outside, and small
spores(Harmaja1976).Twodiagnosticcharactershavebeen
observedin the twocollections examined: 1) a very poorly
differentiated ectal excipulum of textura prismatica to textura
intricata, which is unique within the Otidea species studied by
us;and2)scatteredbrownresinousexudatesonthehyphae
of the medullary excipulum, that are somewhat reminiscent of
those in O. bufonia.
Harmaja(1976)emphasisedthehighwartsontheoutsideof
the apothecia as a unique character for O. papillata, but two ad-
ditional Otidea species, O. tuomikoskii and O. nannfeldtii, have
ashighorhigherwarts.Thesetwospecieshaveinfactbeen
confused with O. papillata(Lundelletal.1985,VanVoorenet
al.2008). Otidea tuomikoskii is distinguished from O. papillata
by narrowly ear-shaped apothecia, a yellow reaction of the
excipuluminKOH,anectalexcipulumoftextura angularis, and
by lacking pigmented exudates on the hyphae of the medullary
excipulum.HighlywartedapotheciaofO. nannfeldtii probably
resemble O. papillata more.However,O. nannfeldtii possesses
pigmented resinous exudates at the septa in the medullary ex-
cipulum, resinous exudates of the ectal excipulum that convert
into amber drops, and most importantly, an ectal excipulum of
textura angularis.
Otidea leporina clade
Apotheciaear-shaped,yellowishochretobrown.Resinousexu-
dates on the ectal excipulum converting into reddish grey drops
inKOH.Associatedwithconifers.
Species — Otidea leporina, O. pseudoleporina.
7. Otidea leporina (Batsch)Fuckel,Jahrb.NassauischenVer-
einsNaturk.23–24:330.1870‘1869–1870’—Fig.4a,10
Basionym.Peziza leporina Batsch, Elench.Fung. 1:117. 1783:Fr.,Syst.
Mycol.2:47. 1822.
 ≡Scodellina leporina(Batsch)Gray,Nat.Arr.Brit.Pl.1:668.1821.
 ≡Helvella leporina (Batsch) Franchi, L. Lami & M. Marchetti, Rivista
Micol.1:63.1999.
 ≡Helvella auricula Schaeff., Fung. Bavar. Palat. Nasc. 4: 103. 1774
(‘Elvela’).
 ≡Wynnella auricula (Schaeff.)Boud.,Icon.Mycol.listprél.600sp.:(2).
1904.
 ≡Otidea auricula (Schaeff.)Sacc.,Syll.Fung.8:95.1889.
Lectotypedesignatedhere:Schaeffer,Fung.Bavar.Palat.Nasc.2:t.156.
1763(‘Elvela decima tertia’).Epitype designated here: SwEdEn, Jämtland,
Östersund,AndersönNatureReserve,underPicea abiesonrichground,28
Aug.2009,K. Hansen & I. Olariaga,KH.09.93(S);MycoBankMBT178082.
= Otidea leporina f. minor Rehm,Ber.Naturhist.VereinsAugsburg 26:
63:1881.
 ≡Otidea leporinavar.minor(Rehm)Sacc.,Syll.Fung.8:94.1889.
Lectotype designated here: gErmany,Leipzig,inderHarth,inspruce
forest,Aug. 1873, G. Winter, Rehm Ascomyceten no. 251 (S-F88382) !
Isolectotype(UPSF-641412)!;MycoBankMBT178088.
= Otidea leporina f. major Rehm,Hedwigia 3–4:2.1883.
= Otidea leporinavar.rubescens Velen.,Monograph.Discom.Bohemiae
1:354.1934.
Lectotype designated here: CzECh rEpubliC, Kosořnear Prague,Sept.
1920,F. Fechtner (PRM614790)!;MycoBankMBT200087.
= Otidea myosotis Harmaja,Karstenia 15:32.1976.
Holotype. Finland, Etelä-Karjala, Hamina, Vehkalahti,Pyhältö, mixed
forest,3Oct.1970,L. Fagerström(H6003548)!
= Otidea crassa W.Y.Zhuang,Mycotaxon 94:366.2006(‘2005’).
= Otidea fuckelii M.Carbone&VanVooren,RivistaMicol. 52:322.2010
(‘2009’).
Holotype.auStria, Nassau, in pinetis umbrosis,FungiRhen.Exs. no.
1233(G00110768).Isotype(S-F114092)!
Misapplied names
–Non Wynnella auriculasensuBoudier,Icon.Mycol. livr.26:n°.535,pl.250.
1909(preliminarytextwith‘circulaires’)(=Wynnella silvicola(Beck)Nannf.).
Apotheciagregarious or caespitose, 17– 52 mmhigh,4–25
mm wide, narrowly to broadly ear-shaped, split, stipitate or
sessile.Hymeniumyellowishbrown(5C6), cinnamon brown
(5D6),paleochre brown (5A5, 5B6) to orange brown(6C7,
6D6),sometimesdark brown (5D8, 5F8, 6E8) whenyoung,
seldom with pale pink stains (6A2), bruised margin orange
brown(6D8),whendriedcinnamonbrown(5A5,5B5)torusty
brown (6D7, 6D8). Receptacle surface ochre brown (5B6),
hygrophanous,indryingpalerochrebrown(4A4,4A5, 4B6),

186 Persoonia–Volume35,2015
whendriedyellowishbrown(5C7,5C8),furfuraceoustofinely
warty,seldomwrinkledatthebase.Wartsconicaltoflattened,
gregarious, concolorous, sometimes darker than the back-
ground,goldenbrown. Stipe4–15×2 – 8 mm.Taste slightly
bitter.Basal tomentum and mycelium abundant, white to cream
white(5A3),sometimesverypalebrownwhendried. Spores
broadly ellipsoid, sometimes inequilateral, with two large
guttules, smooth, hyaline, (12 –)12.5–14 (–15) × 7–8.5 µm
(Lm=12.8–13.8µm,Wm=7.5 – 8.2µm,Qm=1.6–1.8;n=16).
Paraphyses curved to hooked, of the same width or slightly
enlargedatapices,2.5– 4µmwide,withoutnotchesorwith1– 3
low notches, seldom forked at apices, when fresh containing
small,refractive,lightyellowguttules;whendriedsmall,refrac-
tive,hyalinegranules.Asci170 –215×9–10.5µm.Apothecial
section700– 900µmthick.Subhymeniumc.80–100µmthick,
of dense textura intricata, visible as an orange-brown darker
zone,cellscylindricaltoswollen.Medullary excipulum of textura
intricata, 300 –550 µm thick, sometimes differentiated into
twoparts:a)textura angularis underneath the subhymenium,
40–50µmthick,cells6–12µmbroad;b)textura intricata, hy-
phae5 –13 µmwide,sometimesslightly swollen,thin-walled
to slightly thick-walled, hyaline to very pale brown, sometimes
withyellow-brownresinousexudatesatsepta.Ectal excipulum
of textura angularis, 85 –110µmthick,cells thin-walled,pale
brown,15–48×12 – 31µm.Surfacewithbroadlyconicalwarts,
50–85µm high,formedbyshort,fasciculate, hyphoidhairs,
Fig. 10 Otidea leporina*.a,b.Apothecia;c.spores;d.paraphyses;e.ectalexcipulumwithresinousexudates,insertshowsclose-upofamberdropsonthe
ectalexcipuluminMelzer’sreagent;f.basalmyceliumpalebrown,withverysmall,regularlyarranged,resinousexudates(a:KH.09.93,epitype;b:KH.09.102;
c– g:KH.11.02).—Scalebars=10µm;*=allfreshmaterial.

187
I.Olariagaetal.:AmonographofOtidea
of 2 –4 subglobose to elongated cells, constricted at septa,
10–16µmwide,sometimeswithagelatinoussheath.Resinous
exudates abundant, yellow brown, sometimes dissolving in part
intoamberdropsorconvertingintoreddishparticlesinMLZ,
dissolving into yellowish reddish grey heterogeneous drops in
KOH.Basal myceliumof3–4.5(– 6)µmwide,hyalinetopale
brownhyphae,unchangedinKOH,smoothorwithverysmall,
regularly arranged, spheroid, pale brown, resinous exudates,
dissolvinginMLZ,andpartiallyinKOH.
Specimens examined. Canada,Québec,LeVerendryePark,onground
underspruce,16Sept.1965,M.E. Elliot 65-123(UPSF-629640).–CzECh
rEpubliC, CentralBohemianregion,Zdice,Aug. 1924, F. Fechtner(PRM
614787,as O. felina); in piceto ad aciculos, Sept. 1925 (PRM 148836);
Prague-Westdistrict,KosořnearPrague,Sept.1920,J. Velenovský(PRM
614792,asO. umbrina).–dEnmark,Bornholm,RøPlantage,coniferousfor-
est,29Sept.1985,W. Rummel(C-F-47633);NJylland,LangdalPlantage
(nearTranum),under Juniperus, near Picea, 13 Aug. 2009, T. Læssøe,
TL-13769(C);NJylland,RønhøjPlantage,7Oct.1962,A. Hauerbach(C-F-
86691);Jylland,Virklund,SilkeborgSønderskov,inmoss,coniferousforest,
26Sept.1964,H. Dissing(C-F-48298); Sjælland,BrommePlantage, Nof
Sorø,underPicea, 9Oct.1965,H. Dissing(C-F-48299).–Finland, Etelä-
Häme,Mustiala,in pineto, 29Aug. 1866, P.A. Karsten(UPS F-146429);
Perä-Pohjanmaa,Rovaniemi,Kaittiainen,acidicPicea forest,11Sept.2011,
T. Kekki,TK407(TUR);Perä-Pohjanmaa,Rovaniemi,Pisajärvi,old Picea
forest,2Sept.2011,T. Kekki,TK304(TUR);Perä-Pohjanmaa,Rovaniemi,
Välljoki,calciferous Picea forest, 25Aug. 2011,T. Kekki,TK231(TUR).–
FranCE,Loire,LaChamba,ausoldanslalitièred’aiguillesd’épicéa,27Sept.
2008,N. Van Vooren,NV 2008.09.28(dupl.S).–gErmany,Thüringen,in
silvis abiegnis, Kl. & Op.(UPSF-629404,Klotzsch,Herb.Viv.Mycol.143).
–norway, Nord-Trøndelag,Kvam, NoemNortheast,under Picea, among
Rhytidiadelphus squarrosus,2Sept.2009,H. Lindström, KH.09.131 (S);
Nord-Trøndelag,Namdalseid,FlåbekkåsenNatureReserve,Picea and Pinus
old-growthforest,onacidicsoil,amongmosses,4Sept.2009,K. Hansen &
I. Olariaga,KH.09.141(S); Nord-Trøndelag, Steinkjer,Skrattåsen, in rich
Picea abies forest,5Sept.2009,K. Hansen & I. Olariaga, KH.09.145(S);
ibid.,KH.09.147 (S). – SwEdEn, Härjedalen, Torkilstöten, Ljungdalen, on
an active anthill in Piceaforest, 19Aug.2011,J.C. Zamora & I. Olariaga,
KH.11.02(S);Jämtland,in the surroundingsofSällsjö, inyoungstand of
Picea abies, with Betula and Salix,29Aug.2009,K. Hansen & I. Olariaga,
KH.09.100(S);ibid.,KH.09.102(S);Jämtland,SWofMörsil,Sandtjärndalen
NatureReserve,underPicea abies onrichground,7Sept.2009,K. Hansen &
I. Olariaga, KH.09.156(S);Lappland,EkoparkVuollerim,streamfromBränn-
myran, Picea forest, 28Aug. 2008, M. Karström, MK0828 (S);Lappland,
Jokkmokk,‘Nornaskogen’byÅllojaur,mossyPicea forest, on rich ground,
29Aug.2011,K. Hansen & I. Olariaga, KH.11.12(S);ibid., KH.11.14(S);
Lappland,Jokkmokk,UltevisFjällurskogNatureReserve,Sitoätno,neara
Picea, 31Aug.2011,K. Hansen & I. Olariaga,KH.11.33(S);ibid.,KH.11.36
(S);Lappland,Kuouka,15kmSEMessaure,herb-richPicea forest on rich
ground,amongmosses,3Sept. 2011,K. Hansen & I. Olariaga,KH.11.67
(S);Lappland,S of Kvikkjobb-KablaFURNature Reserve, by Kassavare
Mt,Köpenhamn,underPicea, mossyplaceonacidicground,1Sept.2011,
K. Hansen & I. Olariaga,KH.11.76(S);Närke,Knista,Lekhyttan,Kungshall,
under Piceaoncalcareousground,amongmossesandlitter,12Sept.2008,
J. Santos,JS.08.065(S);ibid.,coniferousforest,witholdPicea and Pinus,
K. Hansen,KH.08.108(S);Skåne,Loshult,LillaLoshult,Picea forest,5Sept.
1998,S.-Å. Hanson,SÅH 105838 (C); Uppland, Stockholm, Enebyberg,
Rinkebyskogen,underPicea and Betulaonacidic ground, 2Sept.2009,
K. Hansen & I. Olariaga,KH.09.169(S);Uppland,Täby,Rönninge,closeto
parking place by Arninge, under Piceaonthicklitterlayer,21 Sept.2008,
J. Santos, JS.08.92(S);Uppland,Uppsala,Ersta NatureReserve,onsoil
under Piceain young plantation,23Sept.2008, J. Santos & K. Hansen,
JS.08.99(S);Uppland,Uppsala,Sävja,NorraLunsenNatureReserve,under
Picea, 28Aug. 2008, J. Santos, JS.08.46 (S); Värmland, Gustav Adolf,
Hagfors,Malmbackarna,onmoss,underPicea and Betula,10Aug.2009,
F. Turander s.n.(S).–USA, California, Del Norte Co.,LakeEarlWildlife
Area,14Dec.1997,E.T. Peterson (OSC56824);ibid.,15Dec.1997(OSC
56825);ibid.,underPicea sitchensis, Pinus contorta, Abies concolor, 26
Nov.2001,M. Castellano & E. Cazares(OSC108820);California,Humboldt
Co.,BigLagoonPark,14Dec.1956,A.H. Smith56668(UPSF-629302);
ibid.,underPicea,16Dec.1956,A.H. Smith56799(UPSF-629304);ibid.,
23Dec.1956, A.H. Smith 56954(UPSF-629305);Colorado, Tolland,on
groundin coniferouswoods,28Aug.1920, F.B. Cotner(UPSF-629390);
Oregon,LincolnCo.,FogartyCreekStatePark,15Oct.1997,E.T. Peterson
(OSC56784);Washington,OkanoganNationalForest,PasaytenWilderness,
under Picea engelmannii, Pseudotsuga menziesii, Abies lasiocarpa, Pinus
contorta,16Sept.1999,R. Davis (OSC108856).
Notes — Otidea leporina is probably the most common Oti-
deaspeciesinborealconiferousforestsofEurope.IntheAlps
it often occurs together with Cudonia circinans, an association
notseeninFennoscandia.Itischaracterisedbyear-shaped,
brown apothecia, together with relatively broad spores that
are almost unique within Otidea.Otidea brunneoparva shares
similar spores, but differs from O. leporina in the darker brown
apotheciaandstronglynotchedparaphyses.Otherspeciesof
Otidea that macroscopically resemble O. leporina are distin-
guishedbydifferentsporesizeandshape.
Our morphological and molecular study of the holotype of O. my-
osotis shows it is a synonym of O. leporina (Hansen&Olariaga
2015).TheoriginaldescriptionofO. myosotis(Harmaja1976)
pointed out the apothecial shape and colours, and paraphyses
as diagnostic characters, all of which agree with our concept
of O. leporina.Recently,Harmaja(2009a)statedthattheex-
cipular resinous exudates in O. leporinaconvertin MLZinto
reddish particles and show no reaction in O. myosotis.Inthe
material of O. leporina examined by us, the resinous exudates
dissolve in part, can appear unchanged or can convert into
reddish particles. Sometimes, small amber-drops have also
beenobserved,though not asstrikinglyasin other species.
Therefore,itseemsthatthereactionoftheexudatesinMLZ
is variable within O. leporina, and cannot be used to separate
O. myosotis from O. leporina. Otidea crassa is a synonym
basedprimarilyontheGenBankLSU sequence of the type
collection(DQ443444).
 Nomenclaturalnotes—WhenBatsch(1783)describedPe-
ziza leporina,hereferredtoSchaeffer’splate(1763),beingun-
awareordisregardingSchaeffer’slaterdescriptionof‘Elvela’
auriculaSchaeff.(1774)basedonthesameplate.SinceFries
(1822)sanctionedBatsch’sname,referringtoH. auriculaSchaeff.
(asP. auriculaSchaeff.)asasynonym,P. leporinahaspriority.
Whilemost authors have interpretedthe plate by Schaeffer
(1763,t.156)asaspeciesthatbelongstoOtidea(e.g.Fuckel
1870,Rehm1883,Bresadola1898,Seaver1904),othershave
considered it to represent the monotypic Wynnella(Gonner-
mann&Rabenhorst1869,Quélet1886).Recently,Franchiet
al.(1999)statedthattheSchaefferplaterepresentsW. silvicola
and as they consider Wynnella to be part of the genus Helvella,
they made the combination Helvella leporina.Carbone&Van
Vooren(2010)expresseddoubtsabouthowtointerpretthepro-
tologuebyBatsch(1783)andSchaeffer’splate,andconcluded
thenameisambiguousandrecommendeditnotbeused.In-
stead they introduced the new name O. fuckelii for the Otidea
speciestreatedhere.Basedonourphylogeneticandmorpho-
logicalstudies(seealsoHansen&Olariaga2015)thisnew
name is, however, superfluous, since O. myosotis and O. crassa
are shown to be synonyms of O. leporina.
TheoriginalSchaefferplateshowsseveral,moreorlessevenly
coloured, light ochraceus brown apothecia, conforming to O. le-
porina,andnotbi-colouredapothecia(darkreddishbrownwith
awhitebase)asinW. silvicola.Inouropinion,itleaveslittle
doubt it shows a species of Otidea.Tosettletheuseofthename
O. leporina, and at the same time preserve the use of the well-
established name, W. silvicola(Beck)Nannf.(Nannfeldt1966),
we propose a modern epitype for Peziza leporinaBatsch:Fr.
(Fig.10a),whichrepresentstheOtidea species for which the
namehasmostoftenbeenused.TheepitypeisfromSweden
where Fries saw and studied living material, as indicated by
the abbreviation ‘v. v. ’(vidi vivam,seenliving).

188 Persoonia–Volume35,2015
8. Otidea pseudoleporinaOlariaga&K.Hansen,sp. nov.—
MycoBankMB808972;ITSbarcodeGenBank:KM010112;
Fig.5e,11
Etymology. FromancientGreekψευδο-,whichmeans‘false,fake’,refer-
ring to a close relationship with O. leporina.
Holotype.USA,Oregon,DouglasCo.,EofMillCreek,under Pseudotsuga
menziesii, Abies concolor, Pinus lambertiana, 19 Oct.2010,R. Helliwell,
rh101910(OSC).
Misapplied names
– Otidea cantharella var. minorsensuKanouse,Mycologia41:667.1949.
Apotheciagregarious,10–30(– 50)mmhigh,8–22(– 31)mm
wide, initially narrowly to broadly ear-shaped, margin rounded,
then expanding and sometimes becoming irregularly cup-shaped,
split,stipitateorsessile.Hymeniumochre-orange(4A6,5A7)to
pinkishorange(5A6,5A7),sometimeswithpinkspotsorstains
(6A4),whendriedorange-ochre(5A5,5B5)toreddishbrown
(6D7).Receptacle surface ochre-brown(5B5),hygrophanous,
indryingyellowishochre(4A4,4A5),whendriedyellowishochre
(5A5)to brownish ochre (5B5),furfuraceous to finely warty,
sometimes wrinkled at the base. Warts conical to rounded,
gregarious, concolorous, sometimes distinctly darker than
thebackground,reddishbrown.Stipe4–11×3–5mm.Basal
tomentum and myceliumabundant,whitetopaleyellow(4A2)
orochre(5A2). Spores ellipsoid, sometimes slightly inequilat-
eral,withtwolargeguttules,sometimeswithupto4 smaller
guttules,smooth,hyaline,(9.5 –)10 –12(–12.5)×5.5–6.5µm
(Lm=10.2–11.6µm,Wm=5.7–6.4µm, Qm=1.7–1.9; n=6).
Fig. 11 Otidea pseudoleporina.a,b.Apothecia;c.apothecia†;d.sporesinwater†;e.paraphysesinwater†;f.ectalexcipuluminwater†(a,d–f:rh101910,
holotype;b,c:Moorefun14).—Scalebars=10µm;†=driedmaterial.—Photos:a.R.Helliwell;b.c.J.Moore.

189
I.Olariagaetal.:AmonographofOtidea
Paraphyses curved to hooked, of the same width or slightly
enlargedat apices, 2.5–4.5 µm wide, sometimes with 1–3
notches, apices seldom forked and rarely covered with a hyaline
coating, when dried containing small, refractive, yellow gran-
ules.Asci155– 231×9–10µm.Apothecial section900–1200
µmthick.Subhymeniumc.80–100µmthick,ofdensetextura
intricata,visibleasanorange-browndarkerzone,ofcylindrical
toswollencells.Medullary excipulum500–750µmthick,dif-
ferentiatedintotwoparts:a)textura angularis underneath the
subhymenium,150–200 µm thick, hyphae 8–18 µm broad;
b)textura intricata,hyphae 5–10(–18)µmwide,sometimes
slightly swollen, thin-walled to slightly thick-walled, very pale
yellow, sometimes with yellow-brown resinous exudates at
septa.Ectal excipulum of textura angularis, sometimes of a
textura prismatica,80–110(–150)µmthick,cellsthin-walled
toslightlythick-walled,paleyellow-brown,13–37×8–27µm.
Surfacewithbroadlyconicalwarts,77–115µmhigh,formedby
short,fasciculate,hyphoidhairs,of2–3subglobosetoelong-
atedcells,constrictedatsepta,6–12µmwide,sometimeswith
a gelatinous sheath. Resinous exudates abundant, yellow-
brown,dissolvingintoamberdropsinMLZ,partiallytoentirely
dissolving into reddish grey heterogeneous drops in KOH.
Basal mycelium of 3 –5 µm wide, very pale yellow hyphae,
unchangedinKOH,withverysmall,yellowresinousexudates,
regularlyarranged,spheroid,dissolvinginMLZ,partiallyand
moreslowlydissolvinginKOH.
Specimens examined. USA, California,Trinidad,underspruce,30Nov.
1956,A.H. Smith56168(UPSF-629690);Idaho,IdahoCo.,RickliffCreek
Public Camp, on the ground in Thuja-Tsugawoods,10Oct.1947,W.B. Cooke
21227(UPSF-629388); Idaho,PapooseCreek,SevenDevilsMts,onthe
groundinDouglasfirassociation,3Sept.1954,A.H. Smith & H.E. Bigelow,
47346(UPSF-629331);Oregon,BearSprings,MtHoodNationalForest,18
Oct.1947,A.H. Smith27946(UPSF-629430);Oregon,BentonCo.,Corvallis,
westsideofNWBeechwoodPlace,scattered toclustered onrottingbark
mulch and in thin grass under Pseudotsuga menziesii,14Nov.2010,N.S.
Weber,NSW10202(OSC150347);ibid.,17Nov.2010,N.S. Weber,NSW
10200 (OSC 150345); ibid., 27 Nov.2010, NSW 10201 (OSC 150346);
Oregon,DouglasCo.,EofMillCreek,under Pseudotsuga menziesii, Abies
concolor, Pinus lambertiana,22Oct.2010,J. Moore, Moorefun24(OSC);
Oregon,DouglasCo.,MillCreek,underconifers, 19Oct. 2010,J. Moore,
Moorefun14(S);Oregon,DouglasCo.,RoseburgDistrictBureauofLand
Management,under Pseudotsuga menziesii, Arbutus menziesii, Castanopsis
chrysophylla,11May1997, J. Klein (OSC66261);Oregon, DouglasCo.,
southofLemoloLake,underconifers,5Nov.2010,R. Heliwell, rh179(S);
Oregon,Douglas Co., ThornUnitI, under Pseudotsuga menziesii, Tsuga
heterophylla,21Oct.2010,C. Durbecq,Durbecq16(OSC);Oregon,Jackson
Co.,MedfordBureauofLandManagement,AshlandResourceArea,Beaver
Creek, under Pseudotsuga menziesii, Arbutus menziesii, Toxicodendron
diversilobum, Berberis piperiana,18Dec. 2000, R. Brock (OSC119311);
Oregon,JacksonCo.,MedfordDistrictBureauofLandManagement,Butte
FallsResourceArea,under Pseudotsuga menziesii, Calocedrus decurrens,
Pinus lambertiana, Pinus ponderosa, Quercus kellogii, Rhus diversiloba,
Berberis piperiana, Fragaria vesca, Moehringia macrophylla, grasses,8Mar.
2000, M. Wineteer (OSC72956); Oregon, LaneCo.,Willamette National
Forest,BlueRiver RangerDistrict,under Pseudotsuga menziesii, 18Nov.
1999(OSC72296);Oregon,Marion Co., Breitenbuch Hot Springs Com-
munity,nearDetroitReservoir,inwoods,8Nov.1997,J.W. Spatafora (OSC
56809);Oregon,MtHood,amongmossunderconifers,15Oct.1922,L.E.
Wehmeyer(UPSF-629375);Oregon,WarmSpringsR.,MtHoodNational
Forest,SkylineTrail,29Sept.1947,A.H. Smith & W.B. Gruber,27064(UPS
F-629689);Washington,ClallamCo.,OlympicNationalPark,WhiskeyBend
trailhead,26Nov.1996,E.T. Peterson (OSC56760);Washington,FishCreek
Region,MtRainierNationalPark,25Aug.1948,E.G. Simmons2067(UPS
F-629332);Washington,lowerslopesofRampartRidge,W.ofLongmire,Mt
RainierNationalPark,4Sept.1948,E.G. Simmons2172(UPSF-629386);
Washington,ParkCreek,MtBakerNationalForest,9Sept.1941,A.H. Smith
16755(UPSF-629820);Washington,PierceCo.,MtRainierNationalPark,
LowerTahomaCreek,29Oct.1996,E.T. Peterson (OSC56749).
Notes — Otidea pseudoleporina is recognised by the broadly
ear-shaped apothecia, ochre-orange to pinkish orange hyme-
niumandsmallspores.Ourmulti-genephylogeneticanalyses
(Hansen& Olariaga2015)suggest O. pseudoleporina is the
sister species of O. leporina.Theyshareear-shapedapothe-
cia and resinous exudates on the outer excipulum that partly
convert into heterogeneous reddish drops in KOH. Otidea
leporina differs in the brown apothecia and larger, broadly el-
lipsoidspores.Otidea pseudoleporina resembles O. nannfeldtii
and O. formicarum in the general apothecial shape and small
spores.Otidea nannfeldtii is distinguished by most often lacking
orange tones, having narrowly ear-shaped young apothecia,
and ectal excipular resinous exudates turning reddish brown in
KOH.Otidea formicarum is distinguished by having apothecia
devoid of orange tones, and spores with a lower Qm(1.6–1.7)
than O. pseudoleporina(1.7–1.9).
ThematerialcitedbyKanouse(1949)underO. cantharellavar.
minor most likely represents O. pseudoleporina.Otidea can-
tharella var.minorasdescribedbyBoudier(1909a)hasapale
ochre or grey hymenium, citrine yellow outside, and veins at the
apothecialbase,andrepresentsadifferentspecies(seeunder
O. minor).Peterson(1998)treatedunderO. concinna material
that we refer to O. pseudoleporinaandstatedthatHarmaja
(1974)usedthenameO. cantharellafor the same species.
Otidea concinna is a well-known species in Europe, clearly
distinct from O. pseudoleporina(seeO. concinna).Asforthe
name O. cantharella, the protologue describes a fungus with
the colour of Cantharellus cibarius(Fries1822),andwetypify
it with material of the large-spored species sometimes called
O. caligata (seeO. cantharella).Sofar O. pseudoleporina is
onlyknownfromWesternNorthAmerica.
Otidea tuomikoskii clade
Apothecia ear-shaped. Receptacle surface with warts often
morethan100µmhigh.Basaltomentumlightochretoorange
ochre.Sporessmall,10–11µm long. Sections of apothecia
turningyellowinKOH,especiallythesubhymeniumandectal
excipulum.Associatedwithconifers.
Species — Otidea tuomikoskii.
9. Otidea tuomikoskiiHarmaja,Karstenia15:30.1976—Fig.
12
Holotype.Finland,Etelä-Häme,Lammi,Pappilankylä,Koiransuolenoja,in
needles of Picea abies onananthill,9Sept.1972,R. Tuomikoski (H6002901)!
= Otidea papillataf.pallidefurfuracea VanVooren&Hairaud,Bull.Mycol.
Bot.Dauphiné-Savoie188:56.2008.
Holotype.FranCE,Jura,LesRousses,tourbièreprèsdulacdesRousses,
ausoldanslalitièred’aiguilles,sousépicéas(Picea abies),19Sept.2007,
N. Van Vooren, NV2007.09.27(PC).Isotype(S)!
Misapplied names
– Otidea papillatasensuLundell, Nannfeldt&Holm,FungiExs.Suec.66:
3282.1985.
Apotheciagregarioustocaespitose,17–60mmhigh,7–30mm
wide,longandnarrowlyear-shaped,split,stipitateorsessile.
Hymeniumpalewhitishochre(4A2,4A3)toochreyellow(4A4 –
4A6)rarelywithpinkstains,whendriedlightochreyellow(4A4)
toochre(5B7,5B8).Receptacle surfacebrownishochre(4B7,
4C7)toyellowbrown(5B6–5D6),hygrophanous,ochreyellow
(4A5,4A6,4B6)indrying,whendriedyellowishbrown(5C8,
5D8),warty,rarelywrinkledatthebase.Wartsconical,gregari-
ous, brown, distinctly darker than the background or rarely light-
er.Stipe3–6×2– 3mm.Smellfaintlyaromatic.Basal tomentum
and myceliumabundant, light ochre(5A2) to orange-ochre
(5A4). Spores ellipsoid, slightly inequilateral, with two large
guttules,andsometimeswith1–4smallergranules,smooth,
hyaline,(9.5–)10–11(–12)×5.5– 6.5(–7)µm(Lm=10.3–11.4
µm,Wm=(5–)5.5–6.5(–7.5)µm,Qm=1.7–1.9;n=15).Para-
physescurvedtohooked,oftenbroaderatapices,2.5–5 (–6)
µm wide, sometimes with up to two shallow notches or forked at

190 Persoonia–Volume35,2015
apices, when fresh containing small to large, refractive, hyaline
topaleyellowguttules;whendriedpaleyellow.Asci113 –199
×9–11.5µm.Apothecial section600–700(–1200) µm thick,
paletobrightyellowinKOH.Subhymeniumc.50–90µmthick,
of dense textura intricata,visibleasayellowishbrownzone.
Medullary excipulum of textura intricata,300 – 500(–850)µm
thick,hyphae3 –13µmwide,thin-walledtoslightlythick-walled,
hyalinetoverypaleyellow,withoutresinousexudatesatsepta.
Ectal excipulum of textura angularis,80–120µm,cellsthin-
walled,hyalinetolightyellow,13–40×8 – 25µm.Surfacewith
conicalwarts,55–177µmhigh,formedbyfasciculate,short,
hyphoidhairs,of3–8globosetoelongatedcells,constrictedat
septa,7–15µmwide,sometimeswithagelatinoussheath.Re-
sinous exudates abundant, yellow-orange to yellowish brown,
dissolvingintoamberdropsinMLZ,unchangedinKOH.Basal
myceliumof3.5–6(–7.5)µmwide,oftenthick-walled,septate,
hyalinetoverypaleyellowhyphae,unchangedinKOH,with
regularly arranged, spheroid, yellow to orange resinous exu-
dates,dissolvinginMLZ,partiallyandmoreslowlyinKOH.
Specimens examined. dEnmark, NWJylland,KlimBjerg,soilalongfor-
est road, S.A. Elborne & K. Hansen,16Sept.1998,KH.98.92(C-F-53155).
–EStonia,Põlvamaa,adterraminpicetohumida,11Aug.1960,A. Elango
(UPSF-629392).–Finland, Perä-Pohjanmaa,Rovaniemi,Pisajärvi,oldPicea
forest,2Sept.2011,T. Kekki, TK305(TUR).–FranCE, Charente-Maritime,île
deRé,prèsducamping‘LaBonneÉtoile’,underPinus maritima and Quercus
ilex,onleaflitter,26Nov.2006,M. Hairaud, NV2006.11.05(dupl.S);Rhône,
LesHalles,coldeCroix-Régis,25Oct.2006,J. Cavet, NV2006.10.33(dupl.
S);Rhône,Saint-Nizier-d’Azergues,forêtdePramenoux,sousPicea abies,
20Sept.2008,N. Van Vooren, NV2008.09.08(dupl.S).–gErmany, Lower
Fig. 12 Otidea tuomikoskii*.a,b.Apothecia;c.spores;d.paraphyses;e.ectalexcipulum;f.basalmycelium(a:JS.08.68;b–f:KH.11.77).—Scalebars=10µm;
*=allfreshmaterial.—Photos:a.J.Santos.

191
I.Olariagaetal.:AmonographofOtidea
Saxony,Lüneburg,Boitze,Pinus and Abies,Oct.2010,M. Vega privateherb.
s.n.(dupl.S-F256977).–norway, Nord-Trøndelag,Snåsa,BergsåsenNature
Reserve,under Picea and Pinus,2Sept. 2009,K. Hansen & I. Olariaga,
KH.09.130(S).–Spain, Navarre, Orokieta, Loiandi, Picea abies plantation,
17Oct.2008,J.M. Lekuona (ARAN-FungiA5041195).–SwEdEn, Lappland,
3milesNWVuollerim,Bombmurkleskogen,alongtheStoraLuleälven,herb-
rich Picea forest, 19Aug. 2000,M. Karström, MK200065(S); Lappland,
Norrbotten,Messaure,KaltisbäckenNatureReserve,herbrichPicea forest,
3Sept.2011,M. Karström, KH.11.60(S);Medelpad,SödraSillre,Hussborg,
onwood,22Aug.1998,K. Olofsson (S-F256896);Närke,Hidinge,Lekhyt-
tan,KatteMajaskogen,coniferousforestonlimerichsoil,B. Wasstorp,13
Sept.2008,JS.08.77(S);Närke,Snavlunda,ÖSnavlundaNatureReserve,
under Piceain amixedforest,12Sept.2008,L.G. Hellsten & A. Stridvall,
JS.08.68(S);Närke,Vintrosa,KanterbodaskansNatureReserve,onsoil
underconifers,10Sept.2008,A.B. Nilsson, JS.08.60(S);Skåne,Loshult,
Lilla Loshult, Piceaforestwithafewbroadleaftrees(Betula, Quercus),5Sept.
1998,S.-Å. Hanson, SÅH105768(C);Södermanland,Nacka,Kvarnhagen
bySöderbysjön,onsoilinshadow,moistarea,underPicea,26Sept.2008,
J. Santos, JS.08.100(S);Uppland,Björklinge,Drälinge,amongstneedles
andmossesunderpineinconiferouswoods,10Sept.1936,H.G. Bruun &
H. Smith (S-F92983,FungiExs.Suec.3282);Uppland,Trehörningsskogen
NatureReserve,underPicea on rich ground, on needle litter and decayed
wood,1Sept.2011,M. Prieto & I. Olariaga, KH.11.77(S).–USA, Califor-
nia,DelNorteCo.,EarlLakeStatePark,access by Sand Hill Road, 15
Dec.1997,M. Madsen & R. Davis (OSC56826);California,HumboldtCo.,
Trinidad,Nov.1931,H.E. Parks 3749(UPSF-629376);Oregon,BentonCo.,
Corvallis,McDonald-DunnResearchForest,underconifers,23Oct.1996,
E.T. Peterson (OSC56761);Oregon,BentonCo.,Corvallis,westsideofNW
BeechwoodPlace,scatteredclustersofapotheciaonduffandadjacentto
rotting wood under Pseudotsuga menziesii,19Nov.1997,N.S. Weber,NSW
8553(OSC150344);Oregon,DouglasCo.,BureauofLandManagement,
Roseburg District, Swiftwater ResourceArea, under Tsuga heterophylla,
Pseudotsuga menziesii, Polystichum munitum, Berberis nervosa and Holo-
discus tricolor,8Nov.2000,R. Furriel (OSC105550);Oregon,MarionCo.,
Salem District Bureau of Land Management, Cascades ResourceArea,
under Pseudotsuga menziesii, Tsuga heterophylla, Gautheria shallon, Poly-
stichum munitum, Berberis nervosa, Oxalis oregana, Acer circinatum, Alnus
rubra and Rhododendron macrophyllum,12 Nov.1997, K. Dougan(OSC
66350);Washington,Bremerton,26Oct.1942,J.B. Flett(UPSF-629383);
Washington,Eatonville,18Oct.1954,A.H. Smith49143(UPSF-629385);
Washington,Lower NisquallyR.,Mt RainierNationalPark, 2Sept.1948,
A.H. Smith30888(UPSF-629384).
Notes — Otidea tuomikoskii is characterised by the narrowly
ear-shaped apothecia, with high warts on the outside, small
spores, and the excipulum almost always turning yellow in
KOH,togetherwiththeochretoorange-ochrebasaltomentum
indriedspecimens.Otidea nannfeldtii is probably the species
that resembles O. tuomikoskii most, but O. nannfeldtii has
lower warts, lacks orange tones in the basal tomentum, has
paraphyses only rarely with slightly swollen areas, and resin-
ousexudatesontheouterexcipulumthatturnreddishinKOH.
Otidea papillata shares with O. tuomikoskii conspicuous dark
wartsontheoutsideoftheapothecia(seeunderO. papillata).
TheyellowKOHreactionoftheexcipulum,especiallystrongin
the subhymenium and ectal excipulum, has been observed to be
constant,althoughweakinsomecollections.TheKOHreaction
is stronger in recent collections, and can also be macroscopically
observedinfreshapothecia.TheholotypeofO. tuomikoskii is
fromananthill(Harmaja1976),butO. tuomikoskii most often
produces apothecia among needle litter or even on very decayed
wood,inconiferousforests.ItiswidespreadinEurope,whereit
occursinconiferousplantations,andinWesternNorthAmerica
(Peterson1998),andhasbeenfoundinAsia(Caoetal.1990).
Otidea cantharella clade
Apothecia ear-shaped, or cup-shaped and entire, usually clearly
stipitate.Sporesexceeding20µm,biguttulateandwithseveral
additionalsmallguttules(exceptinO. brunneoparva).Paraphy-
sesoftenstronglynotched.AssociatedwithPicea.
Species — Otidea brunneoparva, O. cantharella, O. propin-
quata.
10. Otidea brunneoparva K.Hansen,M.Carbone,Olariaga&
VanVooren,sp. nov. —MycoBankMB537590;ITSbarcode
GenBank:KM010026;Fig.13,14
Etymology.Harmaja(2009a)usedtheepithetbrunneoparva to provision-
allynamethisspecies.Thenameisvalidatedhereandreferstothesmall
sizeandbrowncolouroftheapothecia.
Holotype.SwEdEn,Närke,Knista,Lekhyttan,Kungshall,calcareousold-
growth forest, in thick litter layer, with Picea and Pinus,12Sept.2008,K. Hansen,
KH.08.107(S).Isotype(C).
Apotheciagregarious,12 – 35mmhigh,7–25mmwide,initially
ear-shaped, apex subacute, broadly ear-shaped in the end, sel-
domalmostcup-shaped,split,stipitate.Hymenium dark brown
(6F3–6F7),sometimesolivaceousbrown(5D7,5E7,5E8)or
reddishbrown(7F3 –7F5),sometimespalerinthemargin,pur-
plishochre(5B3,5B4,6D7),whendriedolivaceousbrown(5F4,
5F5,5F7,5E7)ordarkbrown(6E6,7F7,7F8).Receptacle sur-
faceconcolorousorslightlylighter,darkbrown(5F6,6F7,7F8),
slightlyhygrophanous,indryinggoldenbrown(5D6,5D7),when
drieddark brown (6E7,7E7)tocinnamon brown(5D7,6D7,
6E7),furfuraceous,sometimesfinelywartyatthebase,often
longitudinally wrinkled at the base, sometimes almost reach-
ingthemargin.Wartshemispherical,gregarious,concolorous,
sometimesdarkerorpalerbrownthanthebackground.Stipe
4–7×2 – 3mm.Basal tomentum and mycelium white to very
palebrown,whendriedochrebrown.Spores ellipsoid to broadly
ellipsoid, sometimes very slightly inequilateral, with two large
guttules,seldomwith1–2additionalsmallergranules,smooth,
hyaline,(11–)11.5–14 (–15)×6.5–8.5µm(Lm=11.7–13.8µm,
Wm=7.1–8.3µm,Qm=1.6 –1.7;n=6).Paraphyses hooked,
often inrolled, of the same width or slightly enlarged at the
apicesto3– 5.5µmwide,oftenwithclearnotchesorforkedat
apices, when fresh containing refractive, pale yellowish brown
guttules,restricted totheuppermostpart oftheparaphyses;
whendriedpaleyellow.Asci131–195×8–10µm.Apothecial
section700–800µmthick.Subhymeniumc.100–120µmthick,
visible as a darker brown zone, cells cylindrical to swollen,
densely arranged, with scattered brown resinous exudates at
septa.Medullary excipulum of loosely woven textura intricata,
300–500µmthick,hyphaecylindricaltoslightlyswollen,thick-
walled,5–11µmwide,hyalinetoverypalebrown,sometimes
with brown resinous exudates at septa. Ectal excipulum of
textura angularis,80–100 µm, cells thick-walled, yellowish
brown,20–47×12 – 25µm.Surfacewithbroadlyconicalwarts,
25–65µmhigh,composedoffasciculate,shorthyphoidhairs.
Non-wartedpartswithscatteredhyphoidhairs,of2–3subglo-
bosetoelongatedcells,7.5 –10.5µmwide,slightlyconstricted
atsepta,sometimeswithathingelatinoussheath.Resinous
exudates abundant, yellowish to reddish brown, dissolving into
amberdropsinMLZ.Basal mycelium of3–4.5µmwide,very
palebrownhyphae,unchangedinKOH,smoothorwithvery
small,resinousexudates,dissolvinginMLZ.
Specimens examined. Finland, Etelä-Häme, Hämeen, Lammi, Evo,
Kotinenvirginforest, mesic forest of the Myrtillus type, in needle litter of
Pinus sylvestris and Picea abies in basalpart ofan activeanthill,8Sept.
1978, H. Harmaja (S-F249386); Kainuu, Paltamo, Saukkovaara, under
Picea abies in moist spring-fed site, nearly in water, 24Aug.2011, M. Lahti
(TUR-A198582);Koillismaa,Kuusamo,OulankaNationalPark,firstpartof
theKiutaköngästrail,onrichsoilamongPicea and Betulaleaves,25Aug.
2008,M. Carbone(TUR-A198581);ibid.,16Aug.2009(S-F257086,dupl.
TUR-A198579);ibid.,14Aug.2010(TUR-A198580).–SwEdEn,Jämtland,
Östersund,ÄndsjönNatureReserve,onanabandonedanthillinrichPicea
forest,26Aug.2009,K. Hansen & I. Olariaga,KH.09.82(S);Närke,Hidinge,
Lekhyttan,KatteMajaskogen,coniferouslime-richforest,inlitter,13Sept.
2008,B. Wasstorp,JS.08.73(S);Närke,Knista,Lekhyttan,Kungshall,cal-
careous oldgrowth forest, in thick litter layer, with Picea and Pinus,12Sept.
2008,J. Santos,JS.08.66(S);ibid.,JS.08.69(S).
Notes — Otidea brunneoparva is morphologically and geneti-
callyaclearlydistinctspecies.Itismacroscopicallycharacter-

192 Persoonia–Volume35,2015
ised by stipitate, broadly ear-shaped apothecia with dark brown
coloursandsometimesolivaceousshades.Microscopically,the
sporeshapeandsizearediagnostic(Fig.14a),althoughwith
widevariation.Thestronglyinrolledandnotchedapicesofthe
paraphysesareverycharacteristic(Fig.14b)andotherwiseonly
found in a few species like O. propinquata or O. daliensis. The
thick-walled, yellowish brown, angular cells forming the outer
excipulumarediagnostic too (Fig.14c, d);suchthick-walled
cells in the outer excipulum are only otherwise found in O. pro-
pinquata.ThesixITSsequencesofO. brunneoparva,fromfive
different localities in Finland and Sweden, are identical or with
3–8bpdifferences.TheITSnucleotidediversitywithinO. brun-
neoparva,as sampledhere,is 0.74%persite. Phylogenetic
analysesoffourgene-regions(Hansen&Olariaga2015)show
O. brunneoparva forms a distinct monophyletic group with O. pro-
pinquata and O. cantharella.Thesethreespecies arenever-
theless,easilydistinguishedbothmacro-andmicroscopically.
ThedistinctivesporesofO. brunneoparva are only otherwise
found in O. leporina within Otidea. Otidea brunneoparva is
clearly distinguished from O. leporina by the darker coloured
apotheciaandnotchedparaphyses.Otidea bufonia, O. mirabilis
and O. smithiisharedarkbrowncolourswithO.brunneoparva,
butclearlydifferinapothecialshapeandstature(O. brunneo-
parvabeingmoredelicate),narrowerspores(Lm=6.3 –7.3µm,
Qm=1.9– 2.5)ofdifferentshape(fusoidinO. bufonia and O. mira-
bilis)andtheresinousexudatesontheectalexcipulumnotdis-
solvingintoamberdrops.Otidea fusconigra was published as a
provisionalnametoo.ItalsoresemblesO. brunneoparva in the
darkbrownapothecia,andsporesizeandshape(Jamoni2004).
Nevertheless,the paler hymenium colour (‘grey caffelatte’),
the paraphyses without notches and the habitat among alpine
dwarf Salix, suggests O. fusconigraisadifferentspecies.ITS
and LSU sequences of O. fusconigra(collectionGMFN2293),
obtainedbyus,confirmsO. fusconigraisnotconspecific,but
a sister taxon to O. smithii.
Following Cao et al. (1990), O. brunneoparva keys out as
O. olivaceaJ.Z. Cao &L.Fan.Thecup-shaped, darkbrown
apothecia, with olivaceous tinge, suggest these may be closely
related.However,the spores of O. olivacea were described
asconsiderably longer (14 –17 × 8–8.5 µm), almost non-
overlapping with those of O. brunneoparva.Unfortunately,we
were not able to get the type specimen of O. olivacea on loan for
study.ItshouldbenotedthatO. olivacea is a later homonym of
O. olivaceaBucholtz.ThereforeHarmaja(2009b)publishedthe
new name O. olivaceobrunnea for the illegitimate O. olivacea.
Otidea pusillaRahm might be conspecific with O. brunneo-
parva, but the name is not validly published since no type was
indicated(Art. 40.1 ICN) andmore than one gatheringwas
cited(Art.40.2ICN)(McNeilletal.2012),‘collectedoverthree
weeksinthesamesite’.ThedescriptionofO. pusilla agrees with
O. brunneoparva in the dark brown, cup-shaped apothecia and
therelativelybroadspores.Thesporesizegivenintheproto-
loguewas‘15/ 6 –9µ’(Rahm1958),whichwefinddifficultto
interpretduetotheunusuallybroadwidthrange.Unfortunately,
nomaterialcouldbetracedinZH(pers.comm.R.Berndt).
AcollectionnamedbyHarmajaasO. brunneoparva (H6017193)
wasfoundtobemorphologicallyidenticaltoourmaterial.Also
ITSandLSUsequencesconfirmthatHarmaja´sandourmate-
rialareconspecific. ThereforethenameO. brunneoparva is
hereadoptedandvalidatedasanewspecies.
Otidea brunneoparva appears to be rather widespread in the
Scandinavian Picea forests.Whileatleasttwoofourcollec-
tions,asindicated byHarmaja(2009a)grewon anthills, the
rest were found in places with abundant Picea litter, often with
presence of Betula.Twolocalities,includingthetypelocality,
werecalcareoussuggestingO.brunneoparvaiscalciphilous.
Fig. 13 Otidea brunneoparva apothecia.a.KH.08.107,holotype;b.JS.08.66;c.TUR-A198579;d.TUR-A198580.—Photos:b.J.Santos;c,d.M.Carbone.

193
I.Olariagaetal.:AmonographofOtidea
11. Otidea cantharella (Fr.)Quél.,Enchir.Fung.:275.1886
 —Fig.15
Basionym.Peziza cantharellaFr.,Syst.Mycol.2:48.1822:Fr.,loc.cit.
 ≡Flavoscypha cantharella (Fr.)Harmaja,Karstenia14:107.1974.
Neotype designated here: SwEdEn,Jämtland,Östersund,ÄndsjönNature
Reserve,inrichPicea forest, with Hepatica nobilis and Oxalis acetosella,31
Aug.2009,K. Hansen & I. Olariaga, KH.09.125(S);MycoBankMBT178085.
= Peziza caligataNyl.,ExNot.Sällsk.FaunaFl.Fenn.Förh.10:8.1868
‘1869’.
 ≡Otidea caligata (Nyl.)Sacc.,Syll.Fung.8:95.1889.
 ≡Acetabula caligata (Nyl.)Boud., Hist.Classific.Discomyc.Europe:41.
1907.
Holotype.Finland,Uusimaa,Helsinki,1850,W. Nylander(H009215)!
Misapplied names
– Otidea abietinasensuBreitenbach&Kränzlin,Fung.Switzerland 1:82.
1984.
– Peziza propinquatasensuNannfeldt,Ann.Bot.Fenn.3:313.1966.
Apotheciagregariousorcaespitose,23–68mmhigh,13–43
mm wide, initially broadly ear-shaped, in the end broadly ear-
shapedtoalmostcup-shaped,split,stipitate.Hymenium initially
lightbrown(5B5,5B6,5C6),thenochraceousyellow(4A5,4A6,
5A5),ochre-orange(4A7,5B7),sometimeswithpinkishareasor
reddots(6B8),whenwoundedpinkishandmarginbrownishred
(6B8),whendriedorangebrown(6D7,6D8)toreddishbrown
(6C6,6D6).Receptacle surfaceconcolorous,ochrebrown(4B7,
5C7,5B6),slightlyhygrophanous,indryingyellowishochre(4A6,
4A7),sometimes withpalebrownstains, whendriedorange
Fig. 14 Otidea brunneoparva(KH.08.107,holotype).a.Sporesinwater†;b.paraphyses*;c.medullaryandectalexcipuluminwater †;d.wartfromtheectal
excipulum in water†;e.ectalexcipuluminMelzer’sreagent†;f.basalmyceliuminwater†.—Scalebars=10µm;*=freshmaterial;†=driedmaterial.

194 Persoonia–Volume35,2015
brown (6D7, 6D8), furfuraceous to finely warty,sometimes
wrinkledatthebase.Wartshemispherical,gregarious,conco-
lorous,sometimesdarkerthanthebackground,brown.Stipe
4–25×3 – 9 mm. Basal tomentum and mycelium abundant,
whitetoverypalebrown(5A3),verypalebrownwhendried.
Spores ellipsoid and often narrowing toward poles, sometimes
very slightly inequilateral, with two large and several smaller
guttules,smooth,hyaline,(17–)18–21×(9–)10 –11.5(–12)µm
(Lm=17.7–20µm,Wm=10.4 –11.4µm,Qm=1.7–1.8;n=13).
Paraphyses hooked, of the same width or slightly enlarged at
apices,2.5 – 3.5(–5.5)µmwide, withoutorwitha low notch,
seldom forked at apices, when fresh containing small, refractive,
paleyellowguttules;whendriedsmall,refractivegranules.Asci
191–217×11–12µm.Apothecial section900–1200µmthick.
Subhymeniumc.70 –100µmthick,visibleasadarkeryellow
zone,ofcylindricaltoswollencells,denselyarranged.Medul-
lary excipulum of textura intricata, (500 –)700–900µmthick,
hyphaecylindricaltoslightlyswollen,thick-walled,3–12µm
wide, very pale yellow, sometimes with yellow-brown resinous
exudatesatsepta.Ectal excipulum of textura angularis90–120
µm,cellsthin-walled,paleyellow,23 – 55×12 – 25(–30)µm.
Surfacewithconicaltobroadlyconicalwarts,55– 80µmhigh,
formedbyfasciculate,parallel,shorthyphoidhairs,of3–4ovoid
cells,constrictedatsepta,5–8µmwide.Non-wartedpartswith
singlehyphoid hairs, of 2 – 4 subglobose to elongatedcells,
slightlyconstrictedatsepta,7–10µmwide,sometimeswith
agelatinoussheath.Resinousexudatesabundant,yellowto
yellow-brown,dissolvingintoamberdropsinMLZ.Basal my-
Fig. 15 Otidea cantharella*.a,b.Apothecia;c.spores;d.paraphyses;e.medullaryexcipulumshowingresinousexudatesatsepta;f.ectalexcipulum(a:
KH.09.125,neotype;b:KH.11.69;c,d:KH.10.152;e:KH.09.144;f:KH.09.155).—Scalebars=10µm;*=allfreshmaterial.

195
I.Olariagaetal.:AmonographofOtidea
celium of3.5– 4.5(–6)µmwide,hyalinetoverypaleyellowish
brownhyphae,unchangedinKOH,smoothorwithverysmall,
regularly arranged, spheroid, resinous exudates, dissolving in
MLZandpartiallyinKOH.
Specimens examined. Finland, Etelä-Häme,Hattula,Parola,Alppilankal-
lio, Vaccinium myrtillus-type forest, under Picea abies,25Sept.1967,P. Uotila
618(H);Etelä-Häme,Janakkala,Tervakoski,6Sept.1970,P. Uotila 6195
(H);Etelä-Häme, Mustiala,5 Sept.1895, J. Lindroth(H6010923); ibid.,in
pineto,24Aug.1866,P.A. Karsten(H6010922);Etelä-Häme,Mustiala,Tam-
mela,2Sept.1882,P.A. Karsten(H6010829);Perä-Pohjanmaa,Rovaniemi,
Kalkkinulkki,nearoldlimestonequarry,underPicea,23Aug.2011,T. Kekki,
TK211(TUR);Perä-Pohjanmaa,Rovaniemi,Ounasvaara,Picea forest,29
Aug.2011,T. Kekki,TK279(TUR);Perä-Pohjanmaa,Rovaniemi,Välijoki,cal-
ciferous Picea forest,25Aug.2011,T. Kekki,TK236(TUR);Perä-Pohjanmaa,
Tervola,Peura,oldcalciferousPicea forest,5Sept.2011,T. Kekki,TK177
(TUR);Perä-Pohjanmaa,Ylitornio,Kuusikkorommas,calciferousPicea and
Pinus forest,2Sept.2011,T. Kekki,TK301(TUR);Varsinais-Suomi,Vihti,
Nummela,Metsäkulma, on soil,16Sept. 1979,H. Kotiranta(H6010925).
–FranCE, Isère,Villard-de-Lans,BoisBarbu, underPicea abies, 20Sept.
2008,J. Cavet,NV2008.09.16(dupl.S).– hungary,HoheTatrabeiUnter-
Schmecks,Oct.1884, Linhart(UPSF-629314, RehmAscomyceten251b,
as O. leporina f. minor). – italy, Trentino-AltoAdige, Cavelonte, in silvis
coniferis,sempersociaCudoniaeconfusae,Aug.1898,G. Bresadola(UPS
F-629361).–norway,Nord-Trøndelag,Steinkjer,Noem,under Picea,2Sept.
2009,K. Hansen & I. Olariaga,KH.09.129(S);Nord-Trøndelag,Steinkjer,
Strattåsen, in rich Picea forest, 5 Sept. 2009, K. Hansen & I. Olariaga,
KH.09.144(S).–SwEdEn,Blekinge,Rödeby,Spjutsbygd,c.2kmNWfrom
the train station, on needle litter under a Piceainconiferousforest,10Sept.
1946,S. Lundell & S. Wikland(UPSF-146484);Gästrikland,closetoBergby,
mossy Picea forestonacidicground,31Aug.2010,K. Hansen, K. Gillen &
I. Olariaga,KH.10.152 (S);Hälsingland,Kårböle, StOlofs, 26Aug. 2001,
H.-G. Toresson s.n.(S);Hälsingland,southtocrossingbetweenroadsE45
and310,under Picea abies, amongleaflitter,21Aug.2011,J.C. Zamora
& I. Olariaga,KH.11.109(S);Härjedalen,Linsell,Djursvallen,montaneco-
niferousforestwithbirch,onveryrottenlog,13Aug.2000,B. Gahne(UPS
F-125589);Härjedalen,Torkilstöten, under Picea abies among leaf litter,
onacidicground,20Aug.2011,J.C. Zamora & I. Olariaga,KH.11.106(S);
Jämtland,Hammarstrand,Picea mossyforest,26Aug.2009,H. Lindström,
KH.09.83(S);ibid.,KH.09.84(S);Jämtland,SWofMörsil,Sandtjärndalen
NatureReserve,underPicea abies onrichground,7Sept.2009,K. Hansen
& I. Olariaga,KH.09.155(S); Jämtland, Sällsjö surroundings, in young
stand of Picea abies, on rich ground with Betula and Salix,29Aug.2009,
K. Hansen & I. Olariaga,KH.09.104(S); Jämtland, Östersund,Andersön
NatureReserve,Picea forest onrich ground,28Aug. 2009,K. Hansen &
I. Olariaga,KH.09.95(S);ibid.,underPicea and Pinus,KH.09.96(S);ibid.,
KH.09.101(S);Jämtland,Östersund,FillstabäckenNatureReserve,onold
anthill under Picea, K. Hansen & I. Olariaga, 30Aug.2009,KH.09.111(S);
ibid.,KH.09.117(S);ibid.,K. Hansen & X.H. Wang,5Sept.2012,KH.12.99
(S); Jämtland,Östersund,ÄndsjönNatureReserve,inrichPicea forest, with
Hepatica nobilis and Oxalis acetosella,26Aug.2009,K. Hansen & I. Olariaga,
KH.09.78(S);ibid., KH.09.80(S);ibid.,KH.09.98(S);Jämtland,Åre,Kall,
alongStor-Grundsviken,Kallsjön,onoldanthillinlimerichPicea forest, 15
Aug.2008,J. Santos & K. Hansen, JS.08.18(S);Lappland,Jokkmokk,SE
ofVuollerim,Andersviksravinerna(Natura2000area),mixedforest,31Aug.
2011, A. Stridvall,KH.11.111(S);ibid.,KH.11.112(S);Lappland,Jokkmokk,
byKassavare mountain, Köpenhamn, under Picea abies, 1 Sept. 2011,
K. Hansen & I. Olariaga,KH.11.110(S);Lappland,Jokkmokk,‘Nornaskogen’
byÅllojaur,mossyPicea forestonrichground,29Aug.2011,K. Hansen &
I. Olariaga,KH.11.16(S);Lappland,Kuouka,15kmSEMessaure,herb-rich
Picea foreston rich ground, among mosses,3Sept.2011,K. Hansen &
I. Olariaga,KH.11.68(S);ibid.,KH.11.69(S);Lappland,5kmSEVuollerim,
partofAndersviksravinerna,RävabackenNatureReserve,herb-richPicea
forest,21Aug. 2000, M. Karström, MK200061 (S);ibid.,underPicea on
mossy ground, together with Cudonia confusa, 2 Sept. 2011,K. Wiking,
KH.11.59(S);Småland,Ryssby,Gärdsholmen,Björnö,coniferous forest,
6Sept.1930,H.G. Bruun(UPSF-146485);ibid.,ondeepmossinconifer-
ousforest(UPSF-146486);Uppland,Uppsala,Sävja,NorraLunsenNature
Reserve,Picea forest,onsoil,thicklayeroflitterandmosses,28Aug.2008,
J. Santos, JS.08.47(S).–SwitzErland,WallisCanton,Liddes,Palazuit,in
side of brook under Picea abies,17Aug.2008,M. Carbone(MCVE24217).
Notes — Otidea cantharella can be recognised by broadly
ear-shaped apothecia, with yellow to orange tones, often with
awell-developedstipe,andlargespores.Nootherspeciesof
OtideahavethesporesizeofO. cantharella, in combination
withthosemacroscopiccharacters.
Otidea cantharella is associated with Picea abies, and often
producesapotheciaonanthillsorthickneedlelayers.Further-
more, we have often found apothecia of O. cantharella and
Cudonia confusatogether(Fig.15a),andoncewithSpathularia
rufa, both species belonging to the Rhytismatales.Anapparent
closer association in the same fairy ring has also been observed
once.Interestingly,BresadolanotedtheassociationwithC. con-
fusa inoneofhiscollections(UPSF-629361)studiedbyus.The
presence of C. confusa in the same collecting spot may give
afirstplausiblefieldidentificationofO. cantharella.Moreover,
it suggests a possible biotrophic association between O. can-
tharella and C. confusa.
 Nomenclaturalnotes—Thisspecieshasbeenreferredto
as O. caligata (Nyl.) Sacc. (Nannfeldt 1966, Dissing 2000).
Nevertheless,Harmaja(2009a)cametotheconclusionthat
the name O. cantharella must refer to the large-spored species
treated here, with which some authors have later disagreed
(Carbone2010b,VanVooren2011b).Harmaja stressed the
following characters from the protologue supporting the usage
of the name as presented here: ear-shaped, stipitate apothecia,
the yellow colour of Cantharellus cibarius and the occurrence
in PiceaforestsinsouthernSweden.Wehavestudiedmaterial
fromSmåland (UPS F-146485,UPSF-146486),from where
Fries described O. cantharella.It is thereforeverylikelythat
Fries had in mind the species described here when he coined
the name O. cantharella, although he had only seen dried
material, as indicated by the abbreviation ‘v. s.’(vidi siccam,
seendried).Since nooriginalmaterialis known toexist,we
propose a neotype that will attach the name O. cantharella to
the Otideaspeciesdescribedhere.Thisstabilisestheinterpre-
tationproposedbyHarmaja(2009a),alsousedbyMornand&
Courtecuisse(2005), and longbeforeadoptedby Bresadola
(1900:102), and should serve tosettlethe interpretation of
O. cantharella.OurITS-LSUsequencesoffourO. cantharella
collectionsfromSwedenandFranceareidentical.
12. Otidea propinquata (P.Karst.)Harmaja,Karstenia 15:32.
 1976—Fig.16
Basionym.Peziza propinquata P.Karst.,Not.Sallsk.FaunaFl.Fenn.
Forh.10:110.1869.
Lectotype. Finland, Tavastland, Messuby,7 Oct. 1860, P.A. Karsten
(H6010807)!,selectedbyNannfeldt(1966).
= Otidea abietina f.nigraRick,Oesterr.Bot.Z.48:62.1898.
 ≡Otidea abietinavar.nigra (Rick)Sacc.,Syll.Fung.14:746.1899.
Lectotype.auStria, Vorarlberg,anderGamp,im Nadelwald,1700 m,
Sept.1897,Rick (S-F9962)!,indicatedbyNannfeldt(1966).
= Otidea indivisaVelen.,Monograph.Discom.Bohemiae 1:355.1934.
Lectotype.CzECh rEpubliC,KarlštejnN, Oct.1922,J. Fechtner (PRM
149147)!,selectedbyNannfeldt(1966).
Misapplied names
– Pseudotis abietinasensuBoudier, Icon.Mycol. livr.7: nº.131,pl. 333.
1906(preliminarytextwith‘circulaires’).
– Otidea cochleatasensuBreitenbach&Kränzlin,Fung.Switzerland 1:84.
1984.
Apotheciagregarious or caespitose, 7– 20 mmhigh,15–35
mm wide, obconical to broadly cup-shaped, entire, very rarely
split,regularorsometimesundulatein the margin, stipitate.
Hymeniumochrebrown(5C7,6D5,6D6)todarkreddishbrown
(6F6,6F7, 7F7, 7F8),when dried ochrebrown (6B7, 6C7).
Receptacle surface concolorous, orange brown (6F6, 6F7),
slightlyhygrophanous,indryingyellowishbrown(5C6,5C7),
whendried orange brown (6D6, 6D7, 6E7), furfuraceousto
warty,sometimeswrinkledatthebase.Wartshemispherical,
gregarious, concolorous, sometimes darker than the back-
ground,brown.Stipe 7–16 ×2–7mm.Basal tomentum and
myceliumabundant,whitetoverypalebrown(5A3),verypale
brownwhendried. Spores ellipsoid and often narrowing toward

196 Persoonia–Volume35,2015
the poles, sometimes very slightly inequilateral, with two large
andseveralsmallerguttules,smooth,hyaline,(18–)19–21×
10–12.5µm(Lm=19.3 –20µm,Wm=10.9–11.6µm,Qm=1.6 –
1.7;n=4).Paraphyses hooked, of the same width or often
enlargedatapices,3– 5µmwide,oftenwith1– 3notchesor
forked at apices, when fresh containing small, refractive, light
yellowguttules;whendried hyaline. Asci 231– 275 × 12 –16
µm.Apothecial section1100–1500 µm thick. Subhymenium
c.100–120µmthick, of dense textura intricata, visible as a
darkerbrownzone. Medullary excipulum of textura intricata,
600–800µmthick,hyphae4–9µmwide,sometimesslightly
swollen, thin- to thick-walled, very pale brown, sometimes with
brownresinousexudatesatsepta.Ectal excipulum of textura
angularis100 –130µm,cellsratherthick-walled,palebrown,
17–50×10– 40 µm. Surface with conical to broadly conical
warts,50–75µmhigh,formedbyshort,fasciculate hyphoid
hairs, of 2 –3 subglobose to elongated cells, con stricted at
septaornot,8–11µmwide,sometimeswithapalebrownthick
gelatinoussheath.Resinousexudatesabundant,yellow-brown
tobrown,turningbrownishredinKOH,dissolvinginMLZ.Basal
mycelium of3–6µmwide,hyalinetopalebrownhyphae,un-
changedinKOH,smoothornormallywithregularlyarranged,
spheroidtorod-shaped,resinousexudates,dissolvinginMLZ
andpartiallyinKOH.
Specimens examined. CzECh rEpubliC, Praha, Karlštejn, 9Oct.1922,
F. Fechtner(UPSF-629369,syntypeofO. indivisa);Sept.1924(UPSF-629367,
syntype of O. indivisa).–dEnmark,NESjælland,AsserboPlantage,under
Pinus and Picea amongneedles, 7 Oct. 1975,H. Knudsen(C-F-87203).
Fig. 16 Otidea propinquata.a.Apothecia;b.sporesinwater†;c.paraphysesinKOH †;d.wartoftheectalexcipuluminwater†;e.ectalexcipuluminKOH†;
f.basalmyceliuminwater†(a:KH.11.21;b–f:KH.09.99).—Scalebars=10µm;†=driedmaterial.

197
I.Olariagaetal.:AmonographofOtidea
– FranCE, Isère, Lans-en-Vercors,on the ground, on Picea leaflitter, 17
Sept.2008,J. Cavet,NV2008.09.15(dupl.S).–italy,Trentino-AltoAdige,
Sopramonte,adacusabiegnosinsylvisconiferis,Aug.1898,G. Bresadola
(UPSF-629357).–SwEdEn, Jämtland,Sällsjösurroundings,inyoungstand
of Picea abies, onrich ground, 28Aug. 2009,K. Hansen & I. Olariaga,
KH.09.99(S);Jämtland,Östersund,AndersönNatureReserve,under Pinus
and Picea, onrichground,among mosses, 28Aug. 2009, K. Hansen &
I. Olariaga, KH.09.94(S);ibid.,KH.09.103(S);Jämtland,Östersund,Ändsjön
NatureReserve,inrich Picea forest,26Aug.2009,K. Hansen & I. Olariaga,
KH.09.81 (S); ibid., 31Aug. 2009, KH.09.123 (S); Lappland, Jokkmokk,
‘Nornaskogen’ by Ållojaur,mossy Picea foreston rich ground, 28 Aug.
2011, K. Hansen & I. Olariaga,KH.11.21(S);Lappland,Jokkmokk,Ultevis
FjällurskogNature Reserve,Sitoätno, Picea mossy forest on rich ground,
amongleaflitter,31Aug.2011,K. Hansen & I. Olariaga, KH.11.30(S);ibid.,
KH.11.35(S); Lappland, S of Kvikkjobb-Kabla FUR Nature Reserve, by
Kassavaremountain, Köpenhamn, young Picea standbya road,1Sept.
2011, K. Hansen & I. Olariaga, KH.11.53(S); Lappland,Messaure,herb-
rich Picea forest,29Aug.2008,M. Karström,MK0834(S);Lappland,17km
WSWofVuollerim,Slubbojaureskogen,herb-rich Picea forest,8Aug.2002,
M. Karström, MK0222(S);Närke,Knista,Lekhyttan,under Picea, in thick litter
layer,inlimerichforest,12Sept.2008,J. Santos, JS.08.67(S);Uppland,
Uppsala,SätraNatureReserve,onlitteronlimerichsoil,under Picea,23
Sept.2008,J. Santos & K. Hansen,JS.08.98(S).–USA,Washington,Lake
Crescent,underfir,28Oct.1935,A.H. Smith(UPSF-629351).
Notes — Otidea propinquata is easily recognised by the
stipitate, entire, brown apothecia, large spores, and notched
orforkedparaphyses.Otidea daliensis shares brown, entire
apothecia with O. propinquata, but differs in having sessile
apothecia, with purplish brown tones, and basal mycelium that
lacksabundantresinousexudates.
Therehasbeenuncertaintysurroundingthecorrectname of
thistaxon.Anumberofearlyauthorsusedtheepithetabietina
to refer to O. propinquata(Boudier1906,Bresadola1933).In
agreementwith Harmaja (2009a) andCarbone (2010c), we
consider Peziza abietina a nomen confusum.Nannfeldt(1966)
referred to O. propinquata as O. indivisa and argued that the
type of O. propinquatawasconspecificwithO. cantharella (as
caligata).Harmaja(1976)examinedthelectotypeofO. propin-
quata and found it to be clearly distinct from O. cantharella and
O. indivisa,alatersynonym.Thisisconfirmedbyourstudyof
the type of O. propinquata.Otidea propinquata occurs in Picea
forestsoncalcareousground.ItiswidespreadinNorthernFen-
noscandiaandpresentincentralEurope.
Otidea formicarum clade
Apotheciaear- tocup-shaped,split,ochre toreddishbrown.
Sporessmall,9.5–12µmlong.Basalmyceliumwithabundant
yellowresinousexudates.Associatedwithconifers.
Species — Otidea formicarum, O. nannfeldtii, O. subformica-
rum, O. aff.subformicarum, Otideasp.‘b’.
13. Otidea formicarumHarmaja,Karstenia15:31.1976—Fig.
 5f,17
Holotype.Finland, Etelä-Karjala, Miehikkäla, Savanjärvi, on anthill in
spruceforest,26Sept.1970,L. Fagerström (H6003549)!
Apotheciagregarioustocaespitose,8–22mmhigh,5–20mm
wide, broadly ear-shaped, upper margin rounded, then expand-
ing and sometimes in the end becoming cup-shaped and flat-
tened,split, stipitate orsessile. Hymenium yellowish brown
(5C7,5C8)toreddishbrown(6C7,6C8,6D8),whendriedochre
(4A5,4A6)tobrownishochre(5B7).Receptacle surface yel-
lowishbrown (5C6) to reddishbrown (6C6), hygrophanous,
indrying ochre (5B7),when dried ochre(4A5, 4A6), finely
warty,smoothatthebase.Wartsbroadlyconicaltorounded,
gregarious, concolorous, sometimes slightly darker than the
background,brown.Stipe3–6×2 – 4 mm. Basal tomentum
and mycelium abundant,lightyellow-ochre(5A2). Spores ellip-
soid, seldom very slightly inequilateral, with two large guttules,
smooth,hyaline,9.5 –11(–11.5)×(5.5–)6–7µm(Lm=10–10.7
µm,Wm=6–6.9µm,Qm=1.6 –1.7;n=7).Paraphyses curved
tohooked,ofthesamewidthorslightlyenlargedatapices,3–4
µm wide, with up to two slightly swollen areas, occasionally
notched, sometimes when fresh containing small, refractive,
light yellow or green guttules; when dried small, refractive,
hyalinetolightyellowgranules.Asci160– 232×10.5–13µm.
Apothecial section700– 800µmthick.Subhymeniumc.80–100
µm thick, of dense textura intricata, visible as a yellowish brown
zone,cellscylindricaltoswollen,denselyarranged.Medullary
excipulum 400–500µmthick,differentiatedintotwoparts:a)
textura angularisunderneaththesubhymenium,120–200µm
thick,cells6–15µmbroad;b)textura intricata, hyphae some-
timesslightlyswollen,thin-walledtothick-walled,3 –11µmwide,
hyaline to very pale yellow, sometimes with pale yellow resin-
ousexudatesat septa.Ectal excipulum of textura angularis,
70–120µm,cellsthin-walled,lightyellow,17–40×11–25µm.
Surfacewithbroadlyconicalwarts,45– 60µmhigh,formedby
short,fasciculate,hyphoidhairs,of2–3elongatedcells,7–12
µm wide, not or slightly constricted at septa, sometimes with
agelatinoussheath.Resinousexudatesabundant,brownish
yellow,dissolvingintoamberdropsinMLZ.Basal mycelium of
3–5µmwide,hyalinetoverylightyellowhyphae,unchanged
inKOH,withregularlyarranged,verysmall,spheroidresinous
exudates,dissolvinginMLZ,partiallyandmoreslowlyinKOH.
Specimens examined. Finland, Etelä-Häme,Lammi,Evo,Vahtervehmas,
Kotinenvirginforest,6Sept.1988,H. Harmaja(H6003551);Etelä-Häme,
Loppi,Topeno,Piimästennummi,onahuge anthill,under Picea,16 Sept.
2011, S. Huhtinen11/65(TUR);Perä-Pohjanmaa,Rovaniemi,Välijoki,calcife-
rous Picea forest, onanthill,25Aug.2011,T. Kekki,TK223(TUR);Uusimaa,
Elimäki,Villikkala,Lääksynmäki,mesicheathspruceforest,onanthill,22Oct.
2005, U. Nummela-Salo & P. Salo(H6003550);Varsinais-Suomi,KoskiTl.,
Hongisto,onoldanthillunderPicea abies, with Betulasp.,Salix caprea and
Pinus sylvestris,8Aug.1998,M.-L. & P. Heinonen(TUR124728);Varsinais-
Suomi,Lieto,Suopohja,SEofPäivärinne,alongtheroadbythehouseEsko-
Ukura, on old anthill under Picea abies,2Oct.2009,K. Ruottinen (TUR-A
183242). – FranCE, Haute Savoie, Thorens-Glières, plateau des Glières,
under Picea,ontheground,22Sept.2006,L. Francini,NV2006.09.11(S).
–norway,Nordland,Grane,HolmvassdalenNatureReserve,onoldanthill
under Picea,29Oct.2009,J. Lorås(S-F244372).–SwEdEn,Dalarna,Stora
Tuna,betweenFalubäckenandÖvreMorbyggefäbod,abundantinthelower
partofanactiveanthill,22Sept.1963,R. Morander(UPSF-146725);Närke,
Askersund,Orkarebäckens NatureReserve,calcareous forest, onan ac-
tive anthill under Picea,11Sept.2008,J. Santos, JS.08.63(S);ibid.,under
Picea inlitter,JS.08.62(S);Uppland,Bondkyrka,Gottsundabergen,1Oct.
1927,J.A. Nannfeldt (UPS F-146706);Uppland,Bondkyrka,Nåsten,S of
Läbyvadstation,amongconiferousneedles,16Sept.1932,J.A. Nannfeldt
(UPSF-146709);Uppland, Älvkarleby,Billuddens Nature Reserve,under
Pinus and Picea on sandy ground, at the base of a Pinus,15 Sept.2011,
J.C. Zamora & I. Olariaga,KH.11.104(S);Ångermanland,Ullanger,Håll,in
theeasternslopeofMtMoberget,c.0.5kmSWofp.35,29,ontheground,
14Sept.1974,R. Moberg(UPSF-146735).
Notes — Otidea formicarum is characterised by relatively
small, reddish brown, broadly ear-shaped apothecia, small
spores,andbyitshabitat,often occurring on anthills. For a
comparison with the sister species O. subformicarum see under
thatspeciesbelow.Otidea nannfeldtii resembles O. formicarum,
but differs in often having more narrowly ear-shaped and paler
coloured apothecia, sometimes with pink tones in the hyme-
nium.DriedapotheciaofO. nannfeldtii have, on the contrary,
darker colours than O. formicarum.Otidea pseudoleporina and
O. formicarum share apothecial shape, but O. pseudoleporina
can be distinguished by the ochre-orange colour, and compara-
tivelynarrowerspores(Qm=1.7–1.9).
TheoriginaldescriptionofO. formicarum was based on several
collectionsmadeonanthillsin Finland (Harmaja 1976), and
morerecent material has been cited from anthills(Harmaja
2009a,b),mainlyassociatedwithPicea.Mostofthematerial

198 Persoonia–Volume35,2015
examined by us was also from anthills, but at least one of our
Swedishfinds(KH.11.104)wasfromPinus needle litter, which
shows the habitat of O. formicarum partly overlaps with the
habitat of O. subformicarum.
14. Otidea nannfeldtiiHarmaja,Karstenia15:31.1976—Fig.
5a,d,18
Holotype.Finland,Ahvenanmaa,Lemland,Nåtö,spruceforestnearÖver-
gård,17Sept.1972,C.-A. Haeggström (H6002902)!
= Otidea angustaHarmaja,Karstenia48:35.2009.
Holotype.Finland,Varsinais-Suomi,Lohja,Jalassaari,Ahtiala,EofHeimo
house, mixed somewhat moist rich woods with Picea, Betula, Corylusetc.
onsomewhatcalcareoussoils,23Aug.1965,H. Harmaja (H6010804)!
Misapplied names
– Otidea papillatasensuVanVoorenetal.,Bull.Mycol.Bot.Dauphiné-Savoie
188:52.2008.
Apotheciagregarioustocaespitose,8– 35mmhigh,5 –15mm
wide, initially long, narrowly ear-shaped, sometimes expanding
andbecomingbroadlyear-shaped,split, stipitate or sessile.
Hymenium ochre (4A5), orangish ochre (5B4, 5C4) or pale
brown(6D6,6D7),sometimeswithpinktonesorentirelypink-
ish(6A4), when dried orange-ochre (5A5)to reddish brown
(6C6).Receptacle surfacebrown orange(5B4,5C4)topale
brown(6D6–7),slightlyhygrophanous,indryingochre(4A5),
whendriedreddishbrown(7D7,7E7),finelywarty,smoothat
thebase.Wartsconical,gregarious,concolorous,sometimes
distinctlydarker than thebackground, brown. Stipe4–8×
Fig. 17 Otidea formicarum(KH.11.104)*.a.Apothecia;b. spores;c.paraphyses;d.ectalexcipulum;e. resinousexudatesoftheectal excipulum;f.basal
mycelium.—Scalebars=10µm;*=allfreshmaterial.

199
I.Olariagaetal.:AmonographofOtidea
2–3mm.Basal tomentum and mycelium abundant, white to
lightyellow(5A2)orochre(5A3). Spores ellipsoid, sometimes
slightly inequilateral, with two large guttules, smooth, hyaline,
(9 –)9.5 –10.5(–11.5) × 5.5– 6.5(–7) µm (Lm=9.8 –10.9 µm,
Wm=5.5–6.6µm,Qm=1.6–1.9;n=10).Paraphyses curved
to hooked, of the same width or slightly broader at apices,
2.5–5µmwide,withoutnotches,rarelywithuptotwoslightly
swollen areas or forked at apices, when fresh containing small,
refractive,lightyellowguttules;whendriedhyalinetolightyel-
low.Asci137–190×8 –10µm.Apothecial section650–900µm
thick.Subhymeniumc.80–100µmthick,visibleasayellowish
brownzone,ofcylindricaltoswollencells,denselyarranged.
Medullary excipulum of textura intricata,400–600µmthick,
differentiatedinto two parts:a)textura angularis underneath
thesubhymenium,100 –150 µm thick,cells13–32× 12 – 21
µm;b)textura intricata, 400 – 450µmthick,hyphae4 – 9 µm
wide, thin-walled to slightly thick-walled, hyaline to very pale
yellow,sometimeswithpaleyellowresinousexudatesatsepta.
Ectal excipulum of textura angularis, sometimes of a textura
prismatica,80–120µm,cellsthin-walled,hyalinetolightbrown,
16–40× 11–22 µm. Outerpartofconicalto broadly conical
warts,45 – 85µmhigh,formedbyshort,fasciculate,hyphoid
hairs,of2 – 3(–4)subglobose to elongatedcells,constricted
atsepta,5– 9µmwide,sometimeswithagelatinoussheath.
Resinous exudates abundant, yellow brown, dissolving into
amberdropsinMLZ,reddishbrowninKOH.Basal mycelium
of3– 5.5µmwide,septate,hyalinetoverylightyellowhyphae,
unchangedinKOH,withverysmall,regularlyarranged,sphe-
roid,yellowresinousexudates,dissolvinginMLZ,partiallyand
moreslowlyinKOH.
Fig. 18 Otidea nannfeldtii.a.Apothecia; b.spores*;c. paraphyses*;d. ectalexcipulum*;e.hyphoidhairs withgelatinoussheath*;f.amber dropson the
outermostectalexcipulumcellsinMelzer’sreagent†(a– e:KH.10.302;f:S-F249387).—Scalebars=10µm;*=freshmaterial;†=driedmaterial.

200 Persoonia–Volume35,2015
Specimens examined. Finland, Varsinais-Suomi,Lohja,Jalassaari,Ah-
tiala,Alho,veryclosetotheAhtialaNatureReserve,belowasprucetreein
mixedforestonfairlycalcareoussoil,29Sept.1978,H. Harmaja (S-F249387,
ex H6017194asO. lohjaënsis nom.prov.).–FranCE, HautesAlpes,Provence-
Alpes-Côte-d’Azur,LaBâtie-Montsaléon,LesChariotsduBuech,ausoldans
lalitièred’aiguillesdepins(P. sylvestris),26Oct.2008,N. Van Vooren,NV
2008.10.01(dupl.S).–italy, Abruzzo,Pietracamela(TE),PratidiTivo,on
soil in mixed forest, mainly with Larix but also Picea and Pinus,2Oct.2009,
B. De Ruvo (S-F257096);Calabria,Celico(CS),ContradaColamauci,under
Pinus and Picea abies,16Sept.2009,C. Lavorato, CL091116-17(dupl.S);
Calabria,MoranoCalabro(CS),Campotenese,underPinus sylvestris,7Dec.
2009,C. Lavorato, CL091207-01(dupl.S).–SwEdEn,Gotland,OllajvsNature
Reserve,close toLjugarn,under Picea and Pinus on calcareous ground,
31Sept.2010,K. Hansen, K. Gillen & I. Olariaga, KH.10.302(S);ibid.,under
Picea and Pinusonrichcalcareoussoil,27Sept.2011,H. Tuovila & S. Huh-
tinen,KH.11.115(S);Lappland,Jokkmokk,‘Nornaskogen’byÅllojaur,under
Picea among mosses, K. Olofsson, 1Sept.2011,KH.11.45(S);Lappland,
Messaure,3 miles NW Vuollerim,herbrichPicea forest, 1 Sept. 2005,
M. Karström,MK0536(S); Lappland, VästraTjetnekbäcken,9 km E of
Jokkmokk,underPicea abies, K. Olofsson,1Sept.2011,KH.11.112 (S);
Uppland,Stockholm,Enebyberg,Rinkebyskogen,onsoilunderPicea and
deciduoustreesin side offootpath,29Sept. 2008, J. Santos, JS.08.103
(S).–USA, Oregon,DouglasCo.,ThielsenCreek,underconifers,13Oct.
2010, R. Helliwell, rh101310(OSC).
Otidea cf. nannfeldtii — dEnmark, LøvenholmSkov,Langsø,25kmWof
Grenaa,30Sept.1968,H. Folkmar(C-F-48295),(C-F-48296),(C-F-48297).
Notes — Otidea nannfeldtii is characterised by ochre to light
brown, narrowly ear-shaped apothecia, small spores and resin-
ous exudates on the ectal excipulum turning reddish brown in
KOH.Itresemblesotherspecieswithear-shapedapotheciaand
small spores, such as O. formicarum, O. papillata, O. pseudo-
leporina and O. tuomikoskii.ThemostsimilarspeciesisO. for-
micarum(seeunderO. formicarum).Otidea tuomikoskii is sepa-
rated from O. nannfeldtii by higher and more densely placed
warts on the apothecial outer surface, along with the yellow
reactionoftheexcipuluminKOH.Otidea pseudoleporina has
a brighter ochre-orange hymenium, and the resinous exudates
of the outer excipulum only partly convert into reddish grey
heterogeneousdropsinKOH.
Otidea nannfeldtii, as treated here, shows some phylogenetic
structure(Fig.1;butseeHansen&Olariaga2015).Recently
O. angusta was described as distinct from O. nannfeldtii,
basedonafewsubtlecharacters,i.e.tallerandslightlythicker
fleshed apothecia, with very faintly brownish basal mycelium,
paraphyses with shorter and thinner apical cells, and maybe
smallerspores (Harmaja 2009a).At that time only a single
collectionof each species was known.Basedon studies of
additionalcollections, we did not find correlations between
any morphological characters and the subgroups supported
byourmultiplemolecularphylogenies.Therefore,wepropose
O. angusta be placed in synonymy with O. nannfeldtii.Support-
ing this, our study of the holotype of O. angusta revealed larger
spores(9.5–11×5–5.8µm)thancitedintheprotologue(8 – 9.8
×4.5– 5.2µm),andbroaderparaphyses(2 – 3µm)withlonger
terminalcells(upto67µm).Wehavealsoobservedbothtall,
narrow and rounded apothecia of O. nannfeldtii in the same
spots, and we consider the apothecial shape to vary during
development.Otidea lohjaënsis,proposedbyHarmaja(2009a)
asa provisional name,is also suggestedto be conspecific
with O. nannfeldtii. A collection identified by Harmaja as
O. lohjaënsis (S-F249387, ex-H6017194) showed excipular
resinousexudatesdissolvingintoamber drops in MLZ (Fig.
18f),contrarytothekeycharactergivenforO. lohjaënsis (i.e.
notrespondingtoMLZ).ThevariationintheITSregionwithin
O. nannfeldtii as recognised here is high, but displays no length
variation(alignment790bplong).TheITSsequencesofthe
Finnish holotypes of O. angusta and O. nannfeldtiishow26bp
differences.Atthesametime,however,theITSsequenceof
theSwedishKH.10.302shows33and32bpdifferencesfrom
the O. nannfeldtii and O. angustaholotypes,respectively.The
NorthAmericanrh101310shows14and20bpdifferencesfrom
the two holotypes, O. nannfeldtii and O. angusta,respectively.
ThecollectionsshowingnoLSUsequencevariation(Fig.1),
showalsono,oronly1–2bpdifferencesintheITSregion.No
variation was found between the holotype of O. angusta and
theFinnish S-F249387 (identified as theprovisional O. loh-
jaënsisbyHarmaja).
Otidea nannfeldtiiwaspreviouslyknownonlyfromSouthWest
Finland(Harmaja1976,2009a).Herewereportitfromother
areasinEurope,andforthefirsttimefromNorthAmerica.
15. Otidea subformicarumOlariaga,VanVooren,M.Carbone
 &K.Hansen,sp. nov.—MycoBankMB809252;ITSbar-
 codeGenBank:KM010054;Fig.19,21
Etymology.ReferringtoitssimilaritytoO. formicarum.
Holotype.Spain,Huesca,Bielsa,ErmitadeNªSeñoradePineta,1270m,
42.638039,0.180532(decimalformat),underPinus sylvestris and Abies alba
oncalcareousground,13Oct.2012,J.C. Campos & J. Herranz(S-F242696).
Apotheciagregarioustocaespitose,13– 32mmhigh,10–20mm
wide, broadly ear-shaped, soon becoming deeply cup-shaped,
split,marginsometimeslobate,sessileorstipitate.Hymenium
orangebrown(6C6,6C8)todarkreddishbrown(6D7,6D8),
slightlyhygrophanous,whendried orange brown (7A8). Re-
ceptacle surfaceorange brown(6B7,6C7)to reddishbrown
(6D8),hygrophanous,indryingorangeochre(6A6,6A7),when
driedochrebrown(5B6),brown(6C8)orreddishbrown(7D8),
finelyfurfuraceous,smoothatthebase.Wartspresent near
themargin,flatandrounded,concolorous,brown.Stipe3–4
×2–3mm.Basal tomentum and myceliumpaleyellow(4A3).
Spores ellipsoid, slightly inequilateral, with two large guttules,
rarely with one to several smaller granules, smooth, hyaline,
10.5–12× 6 – 6.5µm(Lm=11.1–11.7µm,Wm=6.1–6.7µm,
Qm=1.7–1.9;n=4).Paraphyses broadly hooked, sometimes
curved,sometimesenlargedattheapicesto2.5 –3µmwide,
without notches, sometimes with a brown matter covering the
apices,whenfreshcontainingsmall,paleyellowguttules;when
dried heterogeneous, pale yellow. Asci 184– 237 × 11–11.5
µm. Apothecial section 800 –1200 µm thick. Subhymenium
c.100–150µmthick,ofdensetextura intricata, with tendency to
textura angularis towards the medullary excipulum, visible as a
darkerorangebrownzone,cells2.5–6(–15)µmwide,withscat-
teredbrownresinousexudates.Medullary excipulum300–700
µm thick, of textura intricata, hyphae thin-walled to slightly
thick-walled,3–12µmwide,verypaleyellowtopalebrownish
yellow,sometimeswithbrownresinousexudatesatsepta.Ectal
excipulum of textura angularis,(85 –)100 –120µm,cellsthin-
walled,verypalebrown,(9 –)17– 22(–35)×(6 –)11.5– 23µm.
Surfacewithnarrowlytobroadlyconicalwarts,45–65µmhigh,
formedbyshort,fasciculate,hyphoidhairs,of2–3cylindricalto
ovoidcells,7–10µmwide,notorslightlyconstrictedatsepta.
Non-wartedareaswithsinglehyphoidhairs,of2–3cylindrical
toovoidcells,(5–)7–12µmwide,notorslightlyconstrictedat
septa,sometimeswithagelatinoussheath.Resinousexudates
abundant, pale yellowish brown, dissolving into amber drops
inMLZ,turning slightly darker and partlydissolvinginKOH.
Basal mycelium of2– 5.5µmwide,hyalinetoverylightyellow
hyphae,unchangedinKOH,withrefractive,paleyellowdrops
onthesurface,dissolvinginMLZ.
Specimens examined.italy,Cosenza,Calabria,Colamauci,Celico,under
Pseudotsuga menziesii, C. Lavorato,28Sept.2005, CL050928-30(dupl.
S-F256978).–Spain, Canary Islands, La Palma, under Pinus canariensis,
26Nov.2008,J. Fernández Vicente, P. Iglesias, F. Hidalgo, J.R. Undagoitia,
S. Lequerica & R. Martínez (S-F256979); La Rioja, Clavijo, under Pinus
sylvestris, on the ground, C.M. Pérez del Amo & R. Gil,3Jan.2009,private
herb.CMP1179,RM1095(dupl.S-F256980);Madrid,Bustarviejo-Canencia,
PuertodeCanencia,3Oct.1979,E. Álvarez(AH44526).

201
I.Olariagaetal.:AmonographofOtidea
Other specimens examined.Otidea aff.subformicarum — mExiCo, Sa la-
zar,ParqueNacionalMiguelHidalgo,Abiesforest,23Sept.2007,M. Her-
nández(FH301035); Veracruz,Cofre dePerote,Camino delos Conejos,
Los Lescados, montane forest of Pinus teocote, P. montezumae, Arbutus
jalapensis,18Sept.2007,M.E. Smith(FH301036).
Notes — Otidea subformicarum is closely related to O. formi-
carumbasedonbothmorphologicalandmolecularcharacters.
Both have broadly ear- to cup-shaped apothecia and small
spores,andareassociatedwithconifers.Diagnosticfeatures
of O. subformicarum are the orange-brown to reddish brown
apothecial colours, and especially the long and narrow spores
comparedtorelatedspecies.Otidea formicarum has shorter
andcomparativelymoreroundedspores(Fig.20).AllO. sub-
formicarum apothecia were collected under Pinus, Pseudotsuga
menziesii or Abies.TheecologyofO. formicarum partly over-
laps(seefurtherunder that species). Otidea subformicarum
appears to have a southern European distribution, whereas
O. formicarumisknownonlyfromFennoscandiaandtheAlps.
Otidea subformicarum forms a strongly supported clade in
theITS-LSUphylogeny(Fig.2).ThefourITSsequencesof
O. subformicarum, from Spain and Italy, are nearly identical
(onlyS-F256979differsin1bp),exceptforasmallvariation
intheminisatellites.TheITSnucleotidediversityis0.71%in
the four minisatellites unique to O. subformicarum.Thefive
ITSsequencesof O. formicarum, from Finland, Norway and
Sweden,are likewise almost identical (onlyJS.08.63differs
in2bp).TheITSsequencesofO. subformicarum and O. for-
micarumshowmanynucleotidedifferences(40/747bp).Thus
theinterspecificITSnucleotidediversityismuchhigher(5.35%)
thanthe intraspecific diversity(in O. subformicarum 0.07%
without the minisatellites and in O. formicarum0.10%).
TwocollectionsfromMexico,FH301035andFH301036(here
referred to as O.aff.subformicarum),belongtotheO. formi-
carum clade based on our molecular phylogenetic analyses
(Fig.1,2).AlthoughanalysesofthecombinedITS-LSUdataset
did not resolve the relationships among these three lineages
(Fig.2)andtheLSUphylogenyplaceFH301035asastrongly
supported sister taxon to two collections of O. subformicarum,
ourthree-andfour-genephylogenies(Hansen&Olariaga2015)
suggest the two European species share a most recent com-
monancestor,andtheMexicancollectionsdivergedearlier.The
sporesizeand shape of theMexicancollectionsconformto
those of O. subformicarum(Fig.20),buttheapothecialcolours
differinthesecollections.Freshmaterialhasnotbeenavailable
tous,butthephotooffreshapotheciaofFH301035showsegg-
yellow apothecia, clearly distinct from O. subformicarum.Photos
ofapotheciaofFH301036haveadarkpurplishbrowncolour,
Fig. 19 Otidea subformicarum apothecia.a. S-F242696,holotype; b.S-F256980;c.S-F256979;d.CL050928-30.—Photos:b.C.PérezdelAmo; c.J.
FernándezVicente;d.C.Lavorato.
9.7 10.2 10.7 11.2 11.7
O. formicarum
O. subformicarum
O. aff. subformicarum
O. pseudoleporina
O. nannfeldtii
O. sp. ‘b’
Lm (µm)
6.9
6.7
6.5
6.3
6.1
5.9
Wm (µm)
Fig. 20MeansporelengthandwidthincollectionsofspeciesintheO. for-
micarum clade and O. pseudoleporina, based on 20 spores from each
collection.

202 Persoonia–Volume35,2015
which has not been observed in O. subformicarum.Basedon
these colour differences and molecular characters, we suggest
that those collections belong to an additional undescribed spe-
cies,possiblyendemictoNorthAmerica.Furthercollectionsare
needed to get insights into the variation and delimitation of this
taxonforitsformaldescription.
Outside the O. formicarum clade, a few species have similar
spores to O. subformicarum. These differ by a combination
of other features: Otidea tuomikoskii by narrowly ear-shaped
apothecia,a receptaclesurfacewithhighwarts(55 –177µm
high),anectalexcipulumthatturnsyellowinKOHandtypically
orangeochrebasaltomentum;andO. nannfeldtii by apothecia
often having yellow tones, sometimes pink stains, and above
allbytheshorterspores(Lm=10–10.7µm).
Otidea unicisa clade
Apotheciawithochreyellowtones.Basaltomentumwithochre
tones.Sporesellipsoid,withwarts±ridgesorspinose.Resinous
exudates on the outermost ectal excipulum cells dissolve and
exudebrightyellowpigmentinKOH.
Species — Otidea kaushalii, O. unicisa, O. yunnanensis.
16. Otidea kaushalii(J.Moravec)K.Hansen&Olariaga,comb.
nov.—MycoBankMB810994;Fig.22
Basionym.Sowerbyella kaushaliiJ.Moravec,Mycol.Helv.2:94.1986.
 ≡Aleurina kaushalii(J.Moravec)W.Y.Zhuang&Korf, Mycotaxon29:
312.1987.
 ≡Otideopsis kaushalii(J. Moravec) J.Moravec,Mycol.Helv.3: 138.
1988.
Fig. 21 Otidea subformicarum(S-F242696,holotype).a.Spores*;b.paraphysesandasci*;c.ectalexcipuluminwatershowingreddishbrownresinousexu-
dates†;d.ectalexcipuluminMelzer’sreagentshowingamberdrops†;e.basalmyceliuminwater†;f.basalmyceliuminMelzer’sreagent†.—Scalebars=10
µm;*=freshmaterial;†=driedmaterial.

203
I.Olariagaetal.:AmonographofOtidea
Holotype.india,WestBengal,Darjeeling, Batasi,onsoilanddecayed
woodinangiospermforest,alt.7600f,6Sept.1979,R. Kaushal,PAN18169.
Isotypesherb.J.Moravec(CUP61814,C-F-60847!).
Apothecia16–65mmhigh,12 – 62mmwide,broadlyear-shaped,
split,stipitate.Hymenium dirty grey to very faintly incarnate,
greyishyellow,orange-ochre(5A6)whendried.Receptacle sur-
face darkorangebrown(6E7,6E8)whendried,denselywarty.
Wartsconical,acuteorblunt,denselygregarious,darkerthan
thebackground, dark reddishbrown. Stipe 3 – 32 mmlong,
3–10mmwide.Basal tomentum and myceliumochre(5A4).
Spores ellipsoid to slightly subfusoid, inequilateral, with one or
two large and/ or a few small guttules, with thin, often curved,
spines,upto1(–1.5)µmhigh,denseratthepoles,sometimes
unitedin shortridges,hyaline,14 –17×7–9µm(Lm=14.9–
15.7µm,Wm=7.7–7.9µm,Qm=1.9–2;n=2).Paraphyses
curvedto hooked,sometimesenlargedat apices,2.5–5µm
wide, when dried containing small, yellowish refractive gran-
ules.Apothecial section700–800µmthick.Asci188 – 213×
11–13µm. Medullary excipulum of textura intricata, hyphae
thick-walled,hyaline to very pale yellow.Ectal excipulum of
textura globulosa-angularis,70 –90µmthick,of3 – 4celllay-
ers,cellsthin-walled, paleyellowishbrown,12 – 46 ×13–33
µm.Surface with conicalwarts,denselyplaced, 40–140µm
high,50–137µmwide,formedbyglobosetoelongatedcells,
8–11.5µmbroad.Resinousexudatesabundant,reddishbrown,
amorphous, and/ or many yellowish brown, crystal-like, oblate
Fig. 22 Otidea kaushalii.a.Apothecia;b.sporesinwater†;c.sporesinCottonBlue†;d.paraphysesinKOH†;e.ectalexcipulumandwartswithabundant
crystal-like, resinous exudates in water†;f.close-upofcrystal-like,oblatespheroidexudatesonoutermostectalexcipulumcellsinwater†(a– d,insertiononf:
T.Læssøe6236;e,f:PAN18169,isotype).—Scalebars=10µm;†=driedmaterial.—Photos:a.T.Læssøe.

204 Persoonia–Volume35,2015
spheroid,6–8×7.5 –11µm,striatebodies,withaconstricted
centre,notdissolvinginMLZ,dissolving and exuding bright
yellowpigmentinKOH.Basal mycelium of 4.5–6µm wide,
thin-walled to slightly thick-walled, very pale yellow hyphae,
unchangedinKOH,withregularlyarranged,spheroid,yellow
resinousexudates,dissolvinginMLZ,unchangedinKOH.
Specimens examined. malaySia,SabahKinabalu,onrottenwood,2Mar.
1999,T. Læssøe 6236(C,dupl.BORH).
Notes — Otidea kaushalii is closely related to O. yunnanen-
sisbasedonouranalysesoftheLSU(Fig.1)andmorphology.
AlreadyMoravec(1988)statedthis,whenhecombinedO. kaus-
halii in Otideopsis.Thedistinctlyspinysporeornamentationin
thesetwospeciesappearsidentical;seeSEMphotographsof
the spores of the holotype of O. kaushaliiinMoravec(1986)
and of O. yunnanensisinLiu&Zhuang(2006).Bothspecies
have a receptacle surface with densely placed, high, dark brown
warts and a lighter coloured hymenium, i.e. greyish yellow,
creamtoyellow.Otidea kaushalii is distinguished by the smaller
spores, and possibly by the ectal excipulum of large, yellowish
brownglobosetoangularcells(Fig.22e).Wereportherefor
thefirsttimeasofaruniquetypeofcrystal-like,oblatesphe-
roid, striate exudates on the outermost ectal excipulum cells
of O. kaushalii (Fig.22e,f)thatmightbeconfinedtoO. kaus-
halii within Otidea.Theseareabundantinboththeisotypeand
theMalaysianmaterial.AthirdcollectionofO. kaushalii has
beenreportedbyZhuang&Korf(1987)fromXizang,China.The
MalaysianandChinesecollectionshavesmallerapotheciawith
shorteroralmostnostipe(apotheciaupto19×18mm;stipe
3×3mmintheMalaysianmaterial,Fig.22a)ascomparedto
Fig. 23 Otidea unicisa.a.Apothecium;b.apothecia†;c.sporesinwater †;d.sporesinCottonBlue †;e. paraphysesinwater†; f.ectal excipulumin water†
(a:JK12082101;b:H7003343;c– f:KH.06.06).—Scalebars=10µm;†=driedmaterial.—Photos:a.J.Karakehian.

205
I.Olariagaetal.:AmonographofOtidea
theholotype(apotheciaupto65×62mm;stipe32×10mm).
Theapotheciaof O. yunnanensis have also been described
withvaryingstipelength(6–25×4–6mm).
17. Otidea unicisa(Peck)Harmaja,Karstenia26:44.1986.—
 Fig.23
Basionym.Peziza unicisa Peck, Rep. (Annual)NewYorkState Mus.
Nat.Hist. 26: 81. 1874.
 ≡Sowerbyella unicisa(Peck)J.Moravec,CzechMycol.47:266.1994.
Holotype.USA,NewYork,LewisCounty,Croghan,FeltHouse,ground
inwoods,Sept.(NYSf3283).
Misapplied names
– Otidea grandissensuKanouse,Mycologia41:672.1949;Liu&Zhuang,
Fung.Diversity23:188.2006.
Apotheciagregarioustocaespitose,12– 25mmhigh,12–35mm
wide, initially broadly ear-shaped, then becoming cup-shaped,
split,stipitateorsessile.Hymenium ochraceous yellow, some-
timeswithpinktinges,yellowishochre(5A5,5B5),orange-ochre
(5A6)whendried.Receptacle surface ochraceous yellow, when
drieddarkorangebrown(6E7,6F7),ochre-brown(5A4)towards
the base, warty,occasionally with shallow ribs at the base.
Wartsrounded,gregarious,concolorousorslightlydarkerthan
thebackground,reddishbrown.Stipe4–8×2–4mm.Basal
tomentum and myceliumabundant, yellowish ochre(5A4).
Spores ellipsoid to slightly fusoid, inequilateral, with two large
guttules, often with a few smaller guttules, with small, low warts,
oftenirregularridges,denseratthepoles,hyaline,(13 –)14 –
15.5(–16.5)×6.5–8.5µm(Lm=14.6–15.2µm,Wm=7.1– 8µm,
Qm=1.8– 2.1;n=3).Paraphyses curved to hooked, sometimes
enlargedatapices,2.5–4µmwide,sometimeswith1–2low
notches,whendriedcontainingrefractive,paleyellowgranules.
Asci181–197×8–11µm.Apothecial section750 – 900µmthick.
Subhymeniumc.70 –120µmthick,ofcylindricalcells,densely
arranged.Medullary excipulum of textura intricata,500 –700µm
thick,hyphaethin-walled,3.5–9µmwide,hyalinetoverypale
yellow,withoutresinousexudatesatsepta.Ectal excipulum of
textura angularis,70– 95µmthick,cellsthin-walled,hyalineto
lightyellow,11–30×6.5–18µm.Surfacewithconicalwarts,
35–85µm high,formedbyfasciculate,short, hyphoidhairs,
ofglobose toelongatedcells,constricted atsepta,6 –11µm
wide.Resinousexudatesabundant,darkyellowishtoreddish
brown, dissolving in part and converting into reddish particles
in MLZ, turning brighter yellow in KOH. Basal mycelium of
4.5–6µmwide,thin-walledtoslightlythick-walled,hyalineto
verypaleyellow hyphae, unchanged in KOH, withregularly
arranged, spheroid, yellow resinous exudates, dissolving in
MLZ,unchangedinKOH.
Specimens examined. USA, Massachusetts,Carlisle,GreatFarm,on
woodydebris,8July2006,L. Millman, KH.06.06(FH);Massachusetts,Car-
lisle,TowleConservationLand,15July2006,L. Millman (FH301030);Massa-
chusetts,Purgatory Charm,9 July2003,Z. Wang, ZWGeo65-Clark (S);
Michigan,CutR.,MackinacCo.,infrondosewoods,10Aug.1949,A.H. Smith
33020(UPSF-629380);Michigan,NofHessel,MackinacCo.,infrondose
woods,15Aug.1949,H. Imshaug 3458(UPSF-629377);Michigan, Luce
Co.,TahquamenonFallsStatePark,21Aug.1951,A.H. Smith39070(UPS
F-629382);Michigan,WofDetouronM134,ChippewaCo.,inbeech-maple
woods,14Aug.1949,H. Imshaug3348(UPSF-629379);NewHampshire,
Shelburne (UPS F-629427); North Carolina, Macon Co., near summit of
StandingIndianMountain,onduffandburiedwoodof Betula,1Aug.1969,
H.H. Burdsall2605(dupl.H7003343);WestVirginia,onlightloamandleaf
mold,15July1896,L.W. Nuttall,Ellis868(UPSF-630023);WestVirginia,
MonongahelaNationalForest,DollySodsWilderness,WildlifeTrail(TR560),
on soil among leaf litter under Fagus grandifolia, Acer pennsylvanicum, and
Betulasp.,21Aug.2012,J. Karakehian,JK12082101(FH,dupl.S).
Notes — Otidea unicisa is easily recognised by the spore or-
namentationoflow,delicatewartsandshortridges.Thedried
specimens are typically bicoloured, with dark brown outside
andorange-ochrehymenium.
ThesporeornamentationofO. unicisa was for a long time over-
looked(Harmaja1986).Theornamentationisclearlyvisibleat
1 000×,especiallyatthepoleswhereitismoreprominent.We
agree with Harmaja (2009a) that what Kanouse (1949) de-
scribed under the name O. grandis is O. unicisa.Thebicoloured
apothecia with brown outside and orange hymenium, along
withtheornamentedspores,andsporesizesintherangeof
O. unicisa,supportthisview.Liu&Zhuang(2006)alsogavea
collection with the typical spore ornament of O. unicisa under
the name O. grandis (forSEMoftheornamentseetheirf.8).
BothstudiesmentiontheBoudierplate328(n°.134,1905;as
O. grandis),whichshowsbicolouredapothecia.Otidea unicisa
has not been recorded outside Eastern North America, and
inouropiniontheBoudierplate328showstypicalO. bufonia
apothecia.Corroboratingthis,twoBoudierspecimenslabelled
O. grandis(UPSF-629342 and likelythecollectionused for
plate328:PC0093644),studiedbyus,areinfactO. bufonia.
For further comments on O. grandis see Excluded, dubious
andimperfectlyknowntaxa.
18. Otidea yunnanensis(B. Liu&J.Z. Cao)W.Y.Zhuang &
C.Y.Liu,Fung.Diversity23:188.2006
Basionym.Otideopsis yunnanensisB.Liu&J.Z.Cao,ShanxiUniv.J.,
Nat.Sci.Ed.4:70.1987.
Holotype.China,Yunnan,DulongRiver,LangTuan,ongroundinforest,
30Aug.1982,D.C. Zhang(HKAS12150).
Otidea yunnanensis was described when the monotypic genus
Otideopsiswaserected.ItwasseparatedfromOtidea because
of the ornamented spores and paraphyses with fused apical
portion(Liu & Cao1987). Upon re-examinationof the type
material,Liu&Zhuang(2006)discoveredthattheparaphyses
hadfreeapices.Theyalsoconcludedthatthesporeswerewith
two large and several small guttules, and not multi-guttulate as
originallydescribedbyLiu&Cao(1987).OurLSUphylogeny
confirmsthat O. yunnanensis is deeply nested within Otidea
(Fig.1).ThespeciesiseasilyrecognisedwithinOtidea by the
largesporeswithfine,curvedspines.Differentsporesizeshave
however,beenreportedfromtheholotype:18–20×8–10µm
(includingornament,Liu&Cao1987)and15–21×9.7–10.5
µm(excludingornament,Moravec1988),andincluding one
additionalcollection: 16.5 – 20 × 7.6–10µm (Liu &Zhuang
2006).TheverywidesporemeasurementsgivenbyMoravec
overlap somewhat with the spore length of the closely related
O. kaushalii (forfurthercomparisonsseeunderO. kaushalii).
Otidea bufonia-onotica clade
Apotheciadark brownorochreyellow.Basaltomentum with
brownoryellowtones,especiallywhendried.Sporesfusoidor
ellipsoid,smooth.Resinousexudatesontheectalexcipulum
convertingintoreddishparticlesormeltingintoamberdrops.
Species — Otidea brevispora, O. bufonia, O. fusconigra ad
interim, O. mirabilis, O. onotica, O. purpurea, O. smithii.
19. Otidea brevispora(W.Y.Zhuang)Olariaga&K.Hansen,
comb. & stat. nov.—MycoBankMB808974
Basionym.Otidea onoticavar. brevisporaW.Y.Zhuang,Mycotaxon94:
368.2006‘2005’.
Holotype. China, Yunnan,Baoshan, 24 July 2003, Z.L. Yang (HKAS
43003)!
Otidea onoticavar.brevispora was distinguished from O. ono-
ticabasedonshorterspores(Zhuang2006).Ourstudyofthe
O. brevisporaholotypeconfirmedthesmallerspores(9.5 –10.5
×5.5– 6µm;Lm=9.9µm,Wm=5.6µm,Qm=1.75).Otherwise
O. brevispora shares all macro- and microscopic features with

206 Persoonia–Volume35,2015
O. onotica, including the yellow reaction of the basal mycelium
inKOH.OurMLanalyses,withanLSUGenBanksequenceof
the Chinese holotype, show O. onotica var. brevispora is a sister
taxon to a clade of O. onoticaspecimens(Fig.1).Basedonthis
andthesmallersporesweconsiderittobeadistinctspecies.
20. Otidea bufonia(Pers.)Boud.,Hist.Classif.Discomyc.
 Europe:52.1907.—Fig.4b,c,5b,24
Basionym.Peziza bufoniaPers., Mycol. Eur.1: 225. 1822: Fr.,Syst.
Mycol.2:54.1822.
 ≡Geopyxis bufonia(Pers.)Sacc.,Syll.Fung.8:73.1889.
Lectotype designated here: FranCE,insylvulaVincennes,Aug.1816(L
0116690/911.81.97,Persoonherbarium)!;MycoBankMBT178089.
= Peziza umbrinaPers.,Observ.Mycol.2:77.1799.
 ≡Scodellina umbrina(Pers.)Gray,Nat.Arr.Brit.Pl.1:668.1821.
 ≡Peziza cochleatavar.umbrina(Pers.)Fr.,Syst.Mycol. 2:50. 1822:
Fr.loc.cit(‘a umbrina’).
 ≡Otidea umbrina(Pers.)Bres.,FungiTrident.Ser.2,fasc.11–13:68.
1898.
Lectotypedesignatedhere:Sowerby,Col.Fig.Engl.Fung.1:t.5.1797
(asPeziza cochleata);MycoBankMBT200088.
= Peziza pseudobadiaCooke,Mycographiapart4:176.1877.
 ≡Aleuria pseudobadia (Cooke)Gillet,Champ. FranceDiscomycetes:
38.1879(‘pseudo-badia’).
 ≡Geopyxis pseudobadia(Cooke)Sacc.,Syll.Fung.8:69.1889(‘pseudo-
badia’).
Holotype.FranCE,Mérignac,surunmur,1814(K(M)195314,exherb.
Cooke).
= Otidea pedunculata Velen.,Monogr. Discomyc. Bohemiae 1: 354.
1934.
Lectotype designated here: CzECh rEpubliC,Hrusice,Aug.1924,J. Vele-
novský(PRM147622)!;MycoBankMBT200086.
Fig. 24 Otidea bufonia*.a,b.Apothecia;c.spores;d.paraphyses;e.ectalexcipulum;f.basalmycelium(a:KH.09.172;b:JS.08.55;c – f:KH.09.171)—Scale
bars=10µm;*=allfreshmaterial.—Photos:b.J.Santos.

207
I.Olariagaetal.:AmonographofOtidea
Misapplied names
– Otidea grandissensuBoudier,Icon.Mycol. livr.6:n°.134,pl.328.1905
(preliminarytextwith‘circulaires’).
– Peziza cochleatasensuSowerby,Col.Fig.Engl.Fung.1:t.5.1797.
Apotheciagregarious,rarelycaespitose,15–45mmhigh,17–
32mmwide,initiallyear-shaped,thensoonexpandingandbe-
comingdeeplycup-shaped,split,stipitateorsessile.Hymenium
initiallyorangebrown(6C6)sometimesolivaceousbrown(4D6),
thendarkorangebrown(6F6,7E8),whendriedgreyishbrown
(5E3,5E4), slightly purple.Receptacle surface dark brown
(6E4–6E7),sometimespalerustybrown(6D8,6E8)orpurplish
brown(6E3)orwitholivaceoustones,slightlyhygrophanous,
indryingslightly paler,whendried darkorangebrown(6E5,
6F4),warty,seldomslightlywrinkledatthebase.Wartsconi-
cal to flattened, gregarious, dark brown, sometimes distinctly
darkerthanthebackground.Stipe5–14×7–10mm,oftenhol-
lowandfeltyinside.Smellweak;tastemild.Basal tomentum
and mycelium abundant,brownishwhitetolightbrown(6B3).
Spores narrowly fusoid, rarely ovoid, inequilateral, with two
large guttules, very rarely with a third small guttule, smooth,
hyalinetopaleyellowish,(12–)13–16.5(–18)×6 –7.5(–8)µm
(Lm=12.4–16.1µm,Wm=6.3 –7.3µm,Qm=1.9–2.5;n=8).
Paraphyses hooked, a few curved, of the same width or slightly
enlarged at apices, 3.5 –5 (–7) µm wide, without notches or
rarely with a notch on the underside, when fresh containing
small,refractive, light brownishyellow guttules; when dried
lightbrownish yellow. Asci 143 –172 ×10–12 µm. Apothe-
cial section 1000 –1300 µm thick. Subhymeniumc.70–100
µm thick, visible as a darker zone, composed of cylindrical
to swollen cells, densely arranged, with scattered brownish
resinousexudates at septa.Medullary excipulum of textura
intricata,500–700µmthick,hyphaethick-walled,3.5– 9 µm
wide, hyaline to light brown, some covered with strikingly dark
brown,striateresinousexudates.Bigcrystal-likeaggregates
sometimes present among the hyphae, dissolving in KOH,
visiblein MLZ.Ectal excipulum of textura angularis,80 –100
µmthick,cellsthin-walled,brownish,16– 31×11–25(–30)µm.
Surfacewithbroadlyconicalwarts,40– 80µmhigh,formedby
short,fasiculatehyphoidhairs,of3 –4subglobosetoelongated
cells,constrictedat septa,6–11µmwide,sometimes witha
gelatinoussheath.Resinousexudatesabundant,darkbrown,
partly dissolving and converting into small reddish particles in
MLZ,partlydissolvinginKOH.Basal myceliumof3.5–4.5µm
wide, hyaline to brown hyphae, with oily, light brown drops on
thesurface,sometimescrystalloidandrod-shaped.
Specimens examined. CzECh rEpubliC, Prague-Eastdistrict,infraKlo-
kočná,incarpineto,5Oct.1931,J. Velenovský(PRM150074,syntypeof
O. pedunculata).–dEnmark, Fyn,JuelsbergSkov,NofNyborg,alongroad-
side in Fagus forest, 28Sept.1986,D. Boertman(C-F-47790);EJylland,
LøvenholmSkovene,EldrupSkov,ingrassalongroad,23Sept.1979,E. An-
dersen (C-F-86688); E Jylland, VejleNørreskov, at roadside on rich soil
under Populus tremula, Alnus, Corylus, Betula, Fraxinus, Fagus, etc.,13
Sept.1993, J. Vesterholt(C-F-20453);NEJylland,RubjergKnude,under
Abies,18Sept.1989,C. Lange & J. Vesterholt(C-F-25955);NW Jylland,
NystrupPlantage nearThisted,1Sept.1972,K. Toft(C-F-48018);WJyl-
land, Bordrup Plantage, in Picea and Pinus plantation along forest road on
sandyground,17Sept.2011,M. Sasa(C-F-94240);Lolland,FavrstedSkov,
among mosses, on calcareous soil, by forest-road under Fagus,4Oct.2007,
K. Hansen & B. Kullman,KH.07.37 (S);Sjælland, FeldskovennearSorø,
14Aug.1974,M. Lange(C-F-47995);N Sjælland,Gribskov,underFagus
in theforest,19Sept.1971,H. Dissing(C-F-48019);ibid.,8Oct.1973,P.M.
Pedersen(C-F-48015);Sjælland,HorserødHegn,6kmWofHelsingør,along
roadside,17Sept.1964,H. Dissing(C-F-48000).–Finland,Varsinais-suomi,
Lohja,Virkkala,Pähkinäniemi,herb-richdeciduousforestoncalcareous,mull
soil, with Corylus avellana, Betula pendula, Populus tremula, Picea abies
and Pinus sylvestris,28Aug.1997, J. Vauras12503F(TUR-A,dupl.S).–
FranCE, Fontainebleau, on the ground under Quercus and Abies,Oct.1876,
E. Boudier(PC0093644,asO. grandis);ForêtdeChâtellerault(L0111782,
Herb.Persoon,as Peziza umbrina);Landes,Contis,underPinus pinaster
on sandy soil, 6 Dec. 2009, J.L. Teres,KH.09.249 (S); Loire-Atlantique,
Bourgneuf-en-Retz,5Nov.2009,G. Moyne,NV 2009.11.01(S);Picardie,
Compiègne,Oct.1892,E. Boudier(UPSF-629342,asO. grandis);Saône-et-
Loire,Robin,onthegroundunderCorylus and Quercus, J.-P. Dechaume,NV
2008.09.12(dupl.S);Vendée,Jard-sur-Mer,induneforestwithdominance
of Pinus pinaster and Quercus ilex, G. Ouvrard,NV2006.11.07(S);Vosges,
Rambervillers,forêtdeSaint-Hélène,underAlnus inapeatbog,5Oct.2006,
M. Hurtu,NV2006.10.12 (S). – italy,Caorle (VE), Brussa,Vallevecchia,
under Pinus pineaclose tothesea, 4Nov.2001,E.Campo(S-F257089);
Ceva(CN),9kmfromthecity onthe roadtoViola,onthe sideof atrack
under Quercus pubescens, Castanea sativa and Corylus avellanea,16Sept.
2005, M. Carbone(S-F257088);DeivaMarina (SP), under Quercus ilex,
5Nov.2008,M. Carbone(S-F257090);Piemonte,VignoleBorbera(AL),Fraz.
Varianosuperiore,insoilunderQuercus pubescens and Castanea sativa,19
Oct.2010,M. Carbone (S-F257087).–Spain, Barcelona, entre Sant Feliu de
Codines y Castellterçol, forest with Quercus ilex and Pinus sylvestris, on cal-
careous ground, 14Oct.1976,C. Montoliu (AH44527);Huesca,Javierregay,
under Quercus rotundifolia and Q. humilis humus,5Dec.2009,F. Prieto &
A. Gonzálezs.n.(S);Navarre,Arbizu,inmixedforest,14Nov.2009,J. Martín
(ARAN-FungiA5048005);Navarre,Etxaurimendatea,underQuercus ilex,
2Dec.2006,J.M. Lekuona(ARAN-FungiA5053019);Navarre,Lete,under
Quercus faginea and Q. rotundifolia oncalcareous ground,19Dec.2009,
J.L. Teres & P.M. Pasaban, KH.09.248(S);Valladolid,TudeladeDuero,
PuenteHierro,underQuercus ilex, J. Santos(S-F22110).–SwEdEn,Gotland,
nearVisby,Värnhem,underFagus and Quercus robur on rich ground, with
Hepatica nobilis,22Sept.2009,E. Bohus-Jensen, K. Hansen & I. Olariaga,
KH.09.189(S);Skåne,Degeberga,Mörkavad, broadleaf forest, 27 Sept.
2001, S.-Å. Hanson,SÅH2001-253(C);Skåne,Helsinborg,Gyhult,broadleaf
forest dominated by Fagus and Quercus, on the ground, 13Oct.1994,S.-Å.
Hanson,SÅH16457(C);Uppland,Stockholm,Enebyberg,Rinkebyskogen,
on rich soil in deciduous forest, under Corylus, but also Tilia, Quercus and
Betula,1Sept.2008,J. Santos,JS.08.55(S);ibid.,17Sept.2008,JS.08.79
(S);Uppland, Uppsala, Hågadalen-NåstenNatureReserve,Predikstolen,
under Quercus robur, Picea abies, Corylus and Salix, on rich bare ground,
17Sept.2009,K. Hansen & I. Olariaga,KH.09.171(S);ibid.,KH.09.172(S);
Uppland,Vaksala,nearTörnby,Ekbacken,grassygroundwithbarepatches
of soil, close to a Quercus robur tree,13Sept.2010,K. Gillen & I. Olariaga,
KH.10.199(S);Öland,Algustrum,Hönstorp,500mSOofthevillage,grazed
mixeddeciduousforest,19Sept.1993,T. Knutsson, TK93-209(S);Öland,
Långlöt,Åstad,c.75mfromBjörkerumsvägen,c.550SoftheT-crossing
inÅstad,underCorylus, 6Sept.1998,T. Knutsson,TK98-208(S).–USA,
Michigan,CheboyboganCo.,ColonialPt,BurtL.,inbeech-maplewoods,
9Aug. 1951, A.H. Smith 37560 (UPS F-629510); Michigan, Emmet Co.,
Pellston, in beech-maple woods, 11Aug. 1951, A.H. Smith 37654 (UPS
F-629511);Minnesota,ItascaStatePark,onsoilinmesicdeciduouswoodsof
redoak,buroak,birch,hazelnut,Ostrya,28July2010,R.A. Healy,RH1218
(MIN933332);Minnesota,LakeAlexanderSNA,onsoilinmixedwoodswith
canopy or red oak, Ostrya,whitebirch,16Aug.2011,R.A. Healy,RH1393
(MIN9333323).–without loCality(L0111780, Herb.Persoon, asPeziza
umbrina);(L0111781,Herb.Persoon,asPeziza umbrina).
Notes — Otidea bufonia is macroscopically characterised
by the cup-shaped apothecia, with a dark brown outside and a
brownbasaltomentum,especiallyindriedspecimens.Micro-
scopically, the narrowly fusoid spores and the presence of hy-
phae with striate resinous exudates in the medullary excipulum
areimportantdiagnostic characters. Otidea mirabilis is very
similar to O. bufonia(seeunderO. mirabilis).
ThehymeniumcolourshowsahighvariabilityinO. bufonia(Fig.
24a,b);wehaveobservedorangebrown,olivaceousordark
browntones withinasingle collection.Also thecolourof the
receptacle surface varies and can be with rusty to purplish
brownorolivaceoustones.Ifpurplishtonesarepresentitmay
be confused with O. mirabilis.Thepigmented,striate,crystal-
like exudates on the hyphae of the medullary excipulum are
considered a unique feature for O. bufonia (Korf & Zhuang
1991,Harmaja2009a,Parslow&Spooner2013),andhave
been used to separate O. bufonia from O. mirabilis(Carbone
etal.2010,VanVooren2010).Ourresultsstrengthenthisview.
Wehaveobservedstriateexudatesinallthematerialexamined
(Fig.4b, c).Thesizeandabundance ofthestriate exudates
are variable, and often restricted to the outermost part of the
medullaryexcipulum (Fig. 4b). InO. mirabilis we have only
observedbiflabellate,crystal-likeexudates(Fig.4d).

208 Persoonia–Volume35,2015
An exceptional O. bufonia specimen with deviant spore char-
acterswasdiscovered(NV2009.11.01),inwhichthe spores
were ovoid and considerably shorter than in the rest of the
material.ThiscollectionisnestedwithintheO. bufonia clade,
alongwithtypicalcollections.
Nomenclatural notes — A specimen from the Persoon her-
barium(911.81.97,L0116690)labelledasO. bufonia by Per-
sooncanbeconsideredtypematerial(Harmaja2009a,b),being
collected prior to the publication of O. bufonia.Wedesignateit
hereasthelectotypetostabilisethename.Itincludesasingle
dark brown apothecium, with uniseriate, smooth, subfusoid
spores, 14 –16 × 5.8–7 µm, curved paraphyses, numerous
dark brown, striate exudates on the hyphae of the medullary
excipulum, and outermost ectal excipular cells with dark brown
resinousexudates.
Otidea umbrina has been recognised as a separate species
(Medardi1995, Dissing2000),based mainlyondifferenthy-
meniumcolour.TheoriginaldescriptionofO. umbrina and the
Sowerbyplateto whichPersoon(1799)referred,agree with
our concept of O. bufonia.ConfirmingHarmaja(2009a),three
of the four collections of O. umbrina studied by us and kept in
Persoon´sherbariuminLbelongtoO. bufonia.Thusweregard
O. umbrina as a synonym of O. bufonia in accordance with other
authors(Dennis1978).ThenameO. bufonia is sanctioned at
specificrankandthushaspriorityoverO. umbrina, which is
sanctionedonlyatvarietyrank.ThenameO. grandis has been
used for O. bufoniacollectionswithbicolouredhymenium,i.e.
Fig. 25 Otidea mirabilis*.a,b.Apothecia;c.spores;d.paraphyses;e.ectalexcipulum;f.ectalexcipulumwartinMelzer’sreagent,showingreddishreaction
ofresinousexudates(a:KH.09.188;b:KH.10.294;c– f:KH.10.308).—Scalebars=10µm;*=allfreshmaterial.

209
I.Olariagaetal.:AmonographofOtidea
orange-brown or olivaceous hymenium in contrast to a dark
brownoutside(Boudier1905,VanVooren2010).Fortheinter-
pretation of O. grandis see Excluded, dubious and imperfectly
knowntaxa.WesynonymiseO. pedunculata with O. bufonia
based on the study of two syntypes, and lectotypify it with
PRM147622.Thelectotypeshowsdarkbrown,splitapothe-
cia,fusoidspores (13.5 –15.5 ×5.5–6.5µm,from 8 spores)
and importantly striate exudates on some of the hyphae of the
medullaryexcipulum.Harmaja(2009b)studiedtheholotypeof
P. pseudobadia and found it is a later synonym of O. bufonia.
Only one collection of P. pseudobadia couldbelocatedinKew
(K(M)195314);itisfromMérignacasgivenintheprotologue,
butfromawall,whereasCookewrote‘ontheground’.Inany
case, one of the apothecia corresponds well to the illustration
intheprotologue,andweconsiderthistheholotype(andthe
materialstudiedby Harmaja,althoughnoannotationby him
couldbefound;pers.comm.B.Aguirre-Hudson).
21. Otidea mirabilisBolognini&JamoniinJamoni,Funghie
 Ambiente85–86:56.2001.—Fig.4d,25
Holotype.italy,Piemonte,Alagnavalsesia(VC),Vald’Otro,onsoilunder
Picea and Larix,alt. c.1500m,3Sept. 1999,D. Bolognini,GMFN 1951.
Isotype(S-F256929)!
Misapplied names
– Otidea leporinasensuZhuangproparte,Mycotaxon96:367.2005.
Apotheciagregariousorcaespitose,18–62mmhigh,9–52mm
wide, initially ear-shaped, short or elongated, soon expanding
andbecoming deeply cup-shaped,split, stipitate orsessile.
Hymeniuminitiallyreddishochre(5B7,6B7)orsometimesoli-
vaceousbrown(5F5,5F6),thendarkreddishochraceousbrown
(5D8)todarkpurplebrown(6F4– 6F7),whendriedochraceous
brown(5D6)topurplishbrown(6E5,6E6).Receptacle surface
dark purple brown with bluish lilaceous shades (7F2, 7F3),
fadingawaywithtimetoochraceousdark-brown(5D7),then
darkpurplebrown(6F4– 6F7),sometimeswithgreyishochre
(4A3,4B3)patchesinunexposedparts,hygrophanous,indry-
inglightpurplebrown(5D2,6D2)toochrebrown(5B4),when
driedpurplebrown(6D4,6F3,6F4),warty,oftenwrinkledatthe
base.Wartsbroadlyconicaltohemispherical,obtuse,gregari-
ous, dark brown, sometimes distinctly darker than the back-
ground.Stipe7–17×3–7mm.Smellweak,vaguelyresin-like;
tastemild,slightlybitterintheend.Basal tomentum and my-
celiumabundant,whitishtolightbrown(5A3). Spores narrowly
fusoid, inequilateral, with two large guttules, very rarely with
additional1–3smallguttules,smooth,hyalinetopaleyellow-
ish,(13 –)13.5 –16(–17)×6–7(–7.5)µm(Lm=14.1–15.4µm,
Wm=6.3–6.9µm,Qm=2.1–2.3;n=9).Paraphyses curved to
hooked,ofthesamewidthtoslightlyenlargedatapices,2.5–5
µm wide, without or occasionally with a few low notches towards
the apex, when fresh containing small, refractive, light brown-
ish yellow guttules; when dried refractive, hyaline guttules.
Asci178–204×9–11µm.Apothecial section1000–1300µm
thick.Subhymeniumc.100 –150µmthick,visibleasadarker
zone,composedofirregularly,denselyarranged,globosecells,
withscatteredbrownishresinousexudates at septa. Medul-
lary excipulum of textura intricata,600–900µmthick,hyphae
4–11(–15)µmwide,sometimesslightlyswollen,thintothick-
walled, hyaline to light brown, seldom with brown resinous
exudates at septa or biflabellate exudates, sometimes forming
cross-like aggregates. Ectal excipulum of textura angularis,
80–100µm thick, cellsthin-walled to slightly thick-walled,
brownish,8–30×8 –18µm.Surfacewithbroadlyconicalwarts,
35–55µm high,formedbyshort,fasciculate, hyphoidhairs,
of 2 –4 subglobose to elongated cells, constricted at septa,
6.5–10(–17) µm wide, sometimeswithagelatinous sheath.
Resinousexudatesabundant,darkbrown,partiallydissolving
andturningreddishinMLZ.Basal myceliumof3 – 6µmwide,
light yellow to light brown hyphae, seldom with oily brownish
drops on the surface, with abundant small dark brown resinous
exudates,roundedtoirregular.
Specimens examined. dEnmark,Bornholm,RøPlantage,onneedlelayer
under Picea,30Sept.2001,C. Lange,KH.01.09(C).–Finland, Koillismaa,
Kuusamo,Oulanka NationalPark,Ampumavaara,southofthemainroad
toLiikanen,14Aug.2010,M. Carbone(S-F257083).–FranCE,Isère,Lans-
en-Vercors,1300m,on theground,on litter,under Larix, 20Sept.2008,
J. Cavet,NV2008.09.14(dupl.S).–india,Uttarakhand,Kalika,Nainital,on
loosehumussoil,22Sept.1973,S. Chandes(UPSF-630072).–italy, Friuli
VeneziaGiulia,Tarvisio(UD),ValSaisera-Valbruna,oncalcareoussoilunder
Picea abies, Larix decidua and Fagus sylvatica,27 Sept. 2010,G. Dose
(S-F257092).–norway,Rana,St.Alteren,7kmWofMoiRana,4Sept.
1972,H. Dissing(C-F-87187).–SwEdEn,Gotland,Ala,Näsmyr,underPicea
and Pinus sylvestris oncalcareousground,30Sept.2010,E. Bohus-Jensen,
K. Hansen, K. Gillen & I. Olariaga,KH.10.294(S);Gotland,Ljugarn,forest
closetoKaupungsFridhem,underPicea abies on calcareous ground, under
acliff, 27 Sept.2010,K. Hansen, K. Gillen & I. Olariaga, KH.10.279 (S);
Gotland,Lojstahed,Russpark,grazedforestwith Pinus sylvestris, on calcare-
ousground,2Oct.2010,K. Hansen, K. Gillen & I. Olariaga,KH.10.308(S);
Gotland,nearVisby,Rävhagen,underPinus sylvestris, with small Quercus
robur, Helianthemum sp.,Melampyrum pratense, Gallium verum, on sandy
andacidifiedsoil,22Sept.2009,E. Bohus-Jensen, K. Hansen & I. Olariaga,
KH.09.188(S);Gotland,Tofta,SmågårdenaturskogNatureReserve,Tofta
strand, under Pinus sylvestrisoncalcareousground,28Sept.2010,K. Hansen,
K. Gillen & I. Olariaga,KH.10.285(S);ibid.,KH.10.288(S).
Otidea cf. mirabilis — SwEdEn,Öland,Gräsgård,Solberga,StoraAlvaret,
1300mSofthevillage,17Oct.1993,T. Knutsson,TK93-276(S).
Notes — Otidea mirabilis is characterised by dark brown
apothecia, purple to lilaceous-bluish shades in the receptacle
surface of especially young apothecia, fusoid spores, and when
present, biflabellate, crystal-like exudates in the medullary ex-
cipulum.Otidea mirabilis and O. bufonia are strongly supported
assisterspeciesinourmultigenephylogeny(Hansen&Olari-
aga2015).Thorough morphological comparisons show that
O. bufonia differs from O. mirabilis in lacking lilaceous-bluish
shades, and having brown striate exudates on some hyphae of
themedullaryexcipulum.Otidea smithii is distinguished from
O. mirabilis by the shorter spores with lower Qm(1.9–2.0).
Otidea mirabilis was described with emphasis on the lilaceous
tones of the outside, in contrast to the paler ochre to olivaceous
hymenium(Jamoni2001).Partofourmaterialshowsthiscol-
our pattern, but collections with dark brown hymenium have
alsobeen observed.Someauthorshave notedtheabsence
of pigmented resinous exudates in the medullary excipulum of
O. mirabilis(Carboneetal.2010,VanVooren2010).Wehave,
however,observedbrowncrystal-likeexudatesinKH.10.308
andtheholotype.Theyareflabellateandformingcross-likeag-
gregates(Fig.4d),clearlydifferentfromthosepresentinO. bu-
fonia (Fig.4b,c).
Coniferous forests on calcareous ground are the typical habitat
of O. mirabilis.TwoChinesecollectionsassignedtoO. leporina
byZhuang(2006)andsequencedbyLiu&Zhuang(2006)are
resolved in the O. mirabiliscladeinourLSUtree(Fig.1).Based
onmorphologywereportitalsofromIndia(UPSF-630072).
22. Otidea onotica(Pers.)Fuckel,Jahrb.NassauischenVer-
eins Naturk. 23–24:330.1870.—Fig.26
Basionym.Peziza onoticaPers.,Syn.Meth.Fung.:637.1801:Fr.,Syst.
Mycol.2:48.1822.
 ≡Scodellina onotica(Pers.)Gray,Nat.Arr.Brit.Pl.1:668.1821.
 ≡Peziza leporina var.onotica (Pers.)P.Karst.,BidragKannedomFin-
landsNaturFolk19:41.1871(‘P. leporina*P. onotica’).
Lectotypedesignatedhere:Sowerby,Col.Fig.Engl.Fung.1:t.79.1797
(as Peziza leporina). Epitype designated here: SwEdEn, Gotland, Ollajvs
NatureReserve,closetoLjugarn,calcareousgroundin mossy Picea forest,
27Sept.2010,K. Hansen, K. Gillen & I. Olariaga, KH.10.284(S);MycoBank
MBT178083.
= Peziza roseaSchumach.,Enum.Pl.2:416.1803.

210 Persoonia–Volume35,2015
Misapplied names
– Otidea concinnasensuBoudier,Icon.Mycol. livr.26:n°.552,pl.325.1909
(preliminarytextwith‘circulaires’).
– Peziza leporinasensuSowerby,Col.Fig.Engl.Fung.1:t.79.1797.
Apotheciagregarioustocaespitose,25–100mmhigh,14–80
mm wide, initially long and narrowly ear-shaped, then soon
expanding, and becoming deeply cup-shaped, split, stipitate
or sessile. Hymenium initially light yellow (4A3 –4A5), then
ochraceousyellow(4A6,4A7,5A6,5A7,5B6,5B7),insome
partslightorange(5A3,5A4),oftenwithpinktonesorentirely
pinkish(6A4),sometimes with red dots,whendriedorange-
ochre (5A5, 5A6) to reddish ochre (6B7, 6B8). Receptacle
surfaceochraceousyellow(4A6,4A7),slightlyhygrophanous,
in drying slightly paler, when bruised sometimes brownish ochre
(5B7)in themargin,whendriedbrownishochrewithorange
tinge(5B7,5B8),slightlywarty,sometimeswrinkledatthebase
whenold.Wartsbroadlyconical,gregarious, concolorous or
sometimesdistinctlydarkerthanthebackground,brown.Stipe
13–27×4–14mm.Smellweak;tastemild.Basal tomentum
and mycelium abundant, whitish to light yellow (4A2, 5A2).
Spores ellipsoid to broadly ellipsoid, inequilateral, with two large
guttules,smooth,hyaline,(11–)12–13.5(–14)×(5.5–)6–7µm
(Lm=12.1–13.3µm,Wm=6.2 – 6.8 µm,Qm=1.8–2;n=10).
Paraphyses curved to hooked, of the same width or slightly
broaderatapices,2.5–4.5µmwide,withoutnotchesorrarely
with a slightly swollen area on the underside, when fresh con-
tainingsmall,refractive,lightyellowguttules;whendriedhyaline
topaleyellowguttules.Asci138– 233×9.5–12µm.Apothecial
section850–1400 µm thick. Subhymenium c. 100 –120 µm
thick, of dense textura intricata,visibleasadarkerzone,cells
Fig. 26 Otidea onotica*. a, b.Apothecia; c. spores; d. paraphyses; e. ectal excipulum; f. basal mycelium (a: KH.10.284, epitype; b: KH.09.132; c – f:
KH.09.165).—Scalebars=10µm;*=allfreshmaterial.

211
I.Olariagaetal.:AmonographofOtidea
cylindricaltoswollen,denselyarranged.Medullary excipulum
of textura intricata, 400–600µmthick,hyphae5–7(–12) µm
wide,thick-walled,hyaline.Ectal excipulum of textura angularis,
80–110µmthick,cells13 – 55×11–28µm,thin-walled,light
yellow.Surfacewithconicalwarts,85–105µmhigh,formedby
short,fasciculatehyphoidhairs,of2 – 3subglobosetoelongated
cells,constrictedatsepta,11–14µmwide.Resinousexudates
abundant,yellow,dissolving intoamberdropsinMLZ. Basal
myceliumof3.5 – 6µmwide,hyalinetoverylightyellowhyphae,
turning yellow in KOH, with very small, regularly arranged,
spheroid,resinousexudatesonthesurface,dissolvinginMLZ,
partiallyandmoreslowlyinKOH.
Specimens examined. CzECh rEpubliC, SouthBohemia,Netolice,Sept.
1922,Hampl(PRM148341).–dEnmark, Amager,Kongelunden,soilalong
forestpath,21July1998,K. Hansen, T. Læssøe & C. Lange, KH.98.107(C);
Møn,FanefjordSkov,underconifersoncalcareousinfluencedsoil,30Sept.
2008,H. Knudsen(C-F-89691);Sjælland,Geelskov,12kmNofCopenhagen,
under Fagus,18Sept.1963,L. Hansen & A. Kjøller(C-F-47985).–FranCE,
Landes, Contis phare, under Pinus pinaster onsandy soil, 9Sept.2009,
J.L. Teres (ARAN-FungiA8200131C).–italy, Piemonte,Vinadio(CN),San
Bernolfo, Picea abies and Abies alba forest, with presence of Fagus sylvatica,
29 Sept. 2008, M. Carbone (MCVE 23277). – norway,Nord-Trøndelag,
Gjøråsvika,Leksvik,underPicea onrichground,amongmosses,3Sept.
2009,K. Hansen & I. Olariaga,KH.09.132(S);underCorylus and Picea on
richground,KH.09.136(S).–Spain,Gipuzkoa,Aia,Amezketalardikobidegu-
rutzea,underAbies alba,3Oct.2009,J.L. Teres (ARAN-FungiA3033701A);
Madrid,SierradeGuadarrama,deciduousforest,amongQuercus pyrenaica
leaves,4Oct.1981,G. Moreno(AH2528);Navarre,Arbizu,mixedforest,14
Nov.2009,J. Martín(ARAN-FungiA5048003);Navarre,Orokieta,Loiandi,in
Picea abies plantation,10Oct.2009,Aranzadi ZE(ARAN-FungiA5041174-
2).–SwEdEn,Skåne,Vittskövle,Segesholm,Herremöllan,broadleafforest,
27 Sept. 2001, S.-Å. Hanson, SÅH 2001-266 (C); Uppland, Stockholm,
NorraJärvafältet,HanstaNatureReserve,onrichgroundunderPicea abies
with Corylus avellana,15Sept.2009,K. Hansen & I. Olariaga,KH.09.164
(S);ibid.,KH.09.165(S),KH.09.166(S);Uppland,Uppsala,forest-covered
hill NNE of Naturicum, coniferous forest with Quercus and Corylus,4Oct.
2011, R. Sundin, KH.11.108(S);Uppland,Uppsala,Norra LunsenNature
Reserve,Lunsentorpet,onrichorganicsoil,underdeadstandingPicea,28
Aug.2008,J. Santos,JS.08.48(S);Uppland,Uppsala,Vänge,Fibyurskog
NatureReserve, oldPiceaforest,inthe middleofa path,17 Sept.2009,
K. Hansen & I. Olariaga,KH.09.175(S).–USA, Minnesota,CedarCreek
ESR,onsandysoilinoaksavannah,canopyofpinoak,hazelnut,18July
2011, R.A. Healy,RH1121(MIN933307);Minnesota,LakeAlexanderSNA,
on soil, mixed woods, canopy of red oak, paper birch, Ostrya,29July2010,
R.A. Healy,RH1222(MIN933311);Minnesota,WildRiverStateParkMany,
on moist, sandy soil in mesic deciduous woods with oak, 23 July 2010,
R.A. Healy,RH1199(MIN933309);Oregon,BentonCo.,Philomath,Wood
CreekRoad,18Nov.1996,E.T. Peterson(OSC56759);Washington,Lower
TahomaCreek,MtRainierNationalPark,27Aug.1948,A.H. Smith30764
(UPSF-629424); Washington,NisquallyR.,Mt Rainier NationalPark,30
Aug.1948,H.A. Imshaug2116(UPSF-629438).
Notes — Otidea onotica is one of the most common and well-
knownspeciesofthegenus.Itischaracterisedbyratherlarge,
ochraceous yellow apothecia, often with a pinkish tinge, dots
orstainsinthehymenium,andsporesofunusualsizewithin
Otidea. The presence of pinkish tones varies considerably.
Apothecia ranging from entirely pink to completely devoid
of pinkish tones have been observed in the same locality, in
KH.09.132andKH.09.136,respectively.Thesetwocollections
haveidenticalITSsequences(GenBankaccessionsKM010103
andJN942772).Aconsistentcharacter,herereportedforthe
firsttime,istheyellowreactionofthebasalmyceliuminKOH.
Otidea unicisa resembles O. onotica macroscopically, but is
distinguishedbyornamentedspores.
 Nomenclaturalnotes—Carbone(2009)revisedthenomen-
clature of O. onotica.HeconsideredtheplatebySowerby(1797,
as Peziza leporina)theonlyelementseenbyPersoon,and
thus, the holotype. However, Persoon did indirectly refer to
two elements by giving the habitat in the protologue as beech
forests(faginetis)andcitingSowerby’splate,whichshowsan
oakleaf.Thus,theplateisnottheholotype;i.e.theoneelement
onwhichtheauthorbasedthename(Art.9.1ICN;McNeillet
al.2012).Theuseofthetermholotypecouldbecorrectedto
lectotypeaccordingtoArt.9.9,butthisarticleisnotapplicable
sinceCarbone’sholotypeindicationdoesnotfulfilArt.7.10of
theICN(itdoesnot include the phrase ‘designated here’or
equivalent)(McNeilletal.2012).Carbone(2009)selectedin
additionanepitypefromPersoon´sherbarium.Butasanepi-
type must refer to the type it interprets and there was no validly
selectedtypein 2009,theepitypificationbyCarbone is also
notvalid.WethereforetypifyO. onotica here, by designating
as lectotype the Sowerby plate, supported by a newly collected
epitypewithmultiplegenesequencesandcolourphotographs.
Theselection of an epitypefrom Sweden is justified bythe
namebeingsanctioned(Friessawandstudiedlivingmaterial
as indicated by ‘v. v. ’),andourITSsequencesofO. onotica
(4, 5, 6, 7, 8, 9) from Denmark, Italy,Norway and Sweden
(includingtheepitype)beingidentical.TheLSUsequencesof
O. onotica(4,5)fromDenmarkand Swedendifferonly2 bp
fromtheothercollections.
23. Otidea purpurea (M.Zang)Korf&W.Y.Zhuang,Mycotaxon
 22:507.1985
Basionym.Acetabula purpurea M.Zang,ActaBot.Yunnan.1:101.1979.
Holotype.China,Tibet,Zayu,onthegroundinforestwithPinus yunna-
nensis,1Sept.1976,M. Zang 670(HKAS5670)!
Notes — Otidea purpurea is probably closely related to
O. mirabilis. It is also characterised by dark brown apothecia
with lilaceous tones, but it clearly differs in the smaller spores
(8.8–10×4.5 – 5.2µm).Twoadditionaltaxa,O. subpurpurea
and O. bicolor, should be compared with O. purpurea.Otidea
subpurpurea likewise has violaceous tones on the outside,
buthaslargersporesandasci(Zhuang&Yang2008).Otidea
bicolorhas darkbrownapothecia andsmallspores (Zhuang
2010),andisprobablyalsocloselyrelatedorconspecificwith
O. purpurea,basedonthesporesize.WetreathereO. purpu-
rea in the O. bufonia clade, because of the dark brown apothecia
andbrownbasaltomentum.
24. Otidea smithiiKanouse,Pap.MichiganAcad.Sci.,Part1.
 24:28.1939‘1938’—Fig.27
Holotype.USA,California,CrescentCity,18Nov.1937,A.H. Smith 8843
(MICH14408)!
Apotheciagregariousorcaespitose,32 –70mmhigh,13–40
mm wide, initially narrowly ear-shaped, short or elongated,
finallyexpanding,atlatestagesbecomingdeeplycup-shaped,
split,normallyshortlystipitate.Hymeniumpurple-brown(6D5)
to dark purple-brown (7E5), then ochraceous brown (5D8),
whendriedverypalepurplishbrown(5A2).Receptacle surface
darkpurple-brown(7E5),sometimeswithlilaceousshadesor
lighterochre-brown(6C5)patchesinunexposedparts,slightly
hygrophanous,indryingdarkochraceousbrown(6D7),when
dried reddish brown (6E5, 6E6), warty,sometimes wrinkled
atthebase.Wartsconical,obtuseoracute,gregarious,dark
brown,sometimesslightlydarkerthanthebackground.Stipe
12–26×5–12mm.Basal tomentum and mycelium abundant,
whitishtobrown(5B2,5C2). Spores ellipsoid, sometimes fu-
soid, inequilateral, with two large guttules, smooth, hyaline,
12–14(–14.5)×6 –7.5µm(Lm=12.5 –13.6µm,Wm=6.4–7.1
µm, Qm=1.9–2.0;n=5).Paraphyses curved to hooked, of the
samewidthtoslightlyenlargedatapices,3–5µmwide,some-
timeswith1– 3notchesorwithalownotchneartheapex,when
driedcontainingrefractive,paleyellowguttules.Asci175– 251×
9–11µm.Apothecial section800–1300µmthick.Subhymenium
c.140–180µmthick, of dense textura intricata, visible as a
darkerzone,hyphaeintermixedwithafewsubglobosecells,
withscatteredbrownishresinousexudates at septa. Medul-
lary excipulum of textura intricata,700–950µmthick,hyphae

212 Persoonia–Volume35,2015
Fig. 27 Otidea smithii(ECV3345).a.Apothecia;b.sporesinwater†;c.paraphysesinwater†;d.ectalexcipuluminwater †.—Scalebars=10µm;† = dried
material.—Photos:a.E.Vellinga.
Fig. 28 Otidea borealis(S-F242694,holotype).a,b.Apothecia;c.close-upofapotheciashowingawartyexternalreceptacleanddentatemargin;d.Young
narrowlycup-shapedapothecium.—Photos:M.Carbone.

213
I.Olariagaetal.:AmonographofOtidea
4–14(– 21)µmwide,sometimesslightlyswollen,thin-walledto
slightly thick-walled, light brown, seldom with light brown resin-
ousexudatesat septa.Ectal excipulum of textura angularis,
70–100µmthick,cellsthin-walledtoslightlythick-walled,pale
brown,13–30×8–15µm.Surfacewithconicalwarts,34–71
(–102)µmhigh,formed by short, fasciculate, hyphoidhairs,
of 2 –5 subglobose to elongated cells, constricted at septa,
6–14µmwide,sometimeswithagelatinoussheath.Resinous
exudates rather abundant on the outside, dark brown, partly
dissolvingandturningreddishinMLZ.Basal myceliumof3– 5
µm wide, light to darker brown hyphae, sometimes with pale
brown drops on the surface, with abundant small, rod-shaped
toirregular,brownresinousexudates.
Specimens examined. Canada, BritishColumbia,GoldstreamPark,Vic-
toria,ondecayedwoodundermatureDouglasfir,21Sept.1968,J. Ginns
1212(UPSF-629486).–USA,California,AlamedaCo.,Berkeley,LeConte
Avenue, in lawn under Betula and Cedrus, 25 Oct. 2005, E. Vellinga,
ECV3345(S);California, Del Norte Co., California StatePark,LakeEarl
WildlifeArea,accessbySandHillRoad,withAbies grandis, Picea sitchen-
sis, Polystichum munitum, Rubus spectabilis, Vaccinum ovatum,15 Dec.
1997,M. Madsen & R. Davis (OSC56823);Oregon,BentonCo.,Philomath,
WoodsCreekRoad,15Nov.1997,E.T. Peterson (OSC56811);Washington,
PierceCo.,MtRainierNationalPark,LowerTahomaCreek,30Oct.1996,
E.T. Peterson (OSC56753);ibid.,18Oct.1997(OSC56799);Washington,
LewisCo.,Gifford-PinchotNationalForest,CampCreekFallsTrail,onlitter
with Acer circinatum, A. macrophyllum, Tsuga heterophylla, Berberis nervosa,
Polystichum munitum, Linnaea borealis, Rubus sp., Goodyera oblongifolia,
6Nov.1997, E. Hathaway & E. Millian (OSC56830).
Notes — Otidea smithii is closely related to O. bufonia and
O. mirabilis, as shown by morphological and molecular charac-
ters.Thesethreespeciessharedarkbrownapothecia,brown
basal tomentum and dark brown resinous exudates in the ectal
excipulum.Otidea smithii is distinguished by shorter spores with
a lower Qmvalue,andtypicallynarrower,ear-shapedapothecia.
Theunique,biflabellateorstriateexudates in the medullary
excipulum, as seen in O. bufonia and O. mirabilis(Fig.4c,d),
respectively, are not present in O. smithii.Onlypigmentedresin-
ous exudates at the hyphal septa in the medullary excipulum
are rarely found in O. smithii (asinFig.4a).Otidea smithii is so
faronlyknownfromWesternNorthAmerica(Kanouse1949,
Peterson1998).OurITSsequenceofO. smithii(JN942771)
isidenticaltotheITSsequenceoftheholotype(AF072065).
Otidea concinna clade
Apothecia cup-shaped, split or sometimes entire, often with bright
yellowtones.Sporessmall,10–12.5µmlong,smooth.Para-
physestypicallystraightandclaviformatapices.Basalmyce-
liumwithscatteredreddishoryellowresinousexudates.
Species — Otidea borealis, O. caeruleopruinosa, O. concinna,
O. flavidobrunneola, O. lactea, O. minor, O. oregonensis, O. phlebo-
phora, O. rainierensis, O. sinensis, O. tianshuiensis, O. sp. ‘a’.
25. Otidea borealisM.Carbone,Olariaga,K.Hansen&Van
 Vooren,sp. nov.—MycoBank MB809250; ITS barcode
 GenBankKM010023;Fig.28,30
Etymology.Referringtoasupposedlyborealdistribution.
Holotype.Finland,Koillismaa,Kuusamo,Juuma,Jäkälävuoma,western
partofJäkälävuomagorges,manyapotheciainmoistsoilamongmosses
at brookside, under Picea abies, 16Aug.2010,M. Carbone(S-F242694).
Isotype(TUR-A198578).
Apotheciagregarioustocaespitose,12–22mmhigh,8–15mm
wide, initially narrowly ear-shaped, upper margin subacute to
rounded,thenbroadlyear-shaped,split,stipitate.Hymenium
pale yellowish ochre (4A2, 5A2), when dried greyish ochre
(4A2, 5A2). Receptacle surface ochraceous yellow (4A5,
5A5),yellowish brown(5B5,5B6)in someparts,sometimes
brown(5D7)inthemargin,hygrophanous,whendriedreddish
brown(6D5,6D6),warty,seldomslightlywrinkledatthebase.
Marginsometimes finely dentateand darker.Wartsbroadly
conical,gregarious,lighttodarkbrown.Stipe well developed,
3–12×2–6mm.Basal tomentum and mycelium white to very
paleyellow(3A2),whendriedverypaleyellow(3A2).Spores
broadly ellipsoid, symmetrical, with two large guttules, smooth,
hyaline,(10–)10.5–11.5× 6–7µm (Lm=10.5 µm,Wm=6.5
µm, Qm=1.7;n=1).Paraphyses straight to curved, very few
hooked,2–2.5µmwide,atapicesupto3 – 4.5(–5)µmwide,
withoutnotches,terminalsegment33 – 47µmlong,narrowly
claviform to claviform, when dried containing slightly refractive,
hyalinegranules.Asci167–198×9–10.5µm.Apothecial sec-
tion1000µmthick.Subhymeniumc.100µmthick,ofdense
textura intricata,cells2– 4µmthick.Medullary excipulum600
µm thick, of textura intricata, hyphae thin-walled to slightly
thick-walled,4–7µmwide,hyaline,withoutresinousexudates.
Ectal excipulum of textura prismatica,80µmthick,cellsthin-
walled,hyaline,unchangedtoverypaleyellowinKOH,10 – 21
×8.5 –11µm.Surfacewith broadlyconicalwarts,63–90µm
high,formedbyfasciculate,short,hyphoidhairs,of3–6sub-
globoseto ovoid cells, constricted atsepta,6–12 µm wide.
Non-wartedparts withsingle,2–5-celled hyphoidhairs,with
cylindricaltoclaviformuppercell,45 –73×8–10µm.Resinous
exudatesgoldenbrown,partlydissolvinginMLZ(amberdrops
notobserved),brightyellowandpartlydissolvinginKOH.Basal
mycelium of3–4.5µmwide,verypaleyellowhyphae,bright
yellowin KOH, with spheroid torod-shaped, reddish yellow
resinousexudates,dissolvinginMLZ,partiallydissolvingand
turningbrightyellowinKOH.
Other specimens examined.Otidea sp.‘a’ — SwEdEn, Lappland, Arvids-
jaurs,LillånAllmänningreservat,8kmSVJärvträsk,herb-richPicea forest
oncalcareousground,12Sept. 2009,M. Karström,MK0942(S);ibid., 10
Sept.2010,MK1081(S).
 Notes—WithintheO. concinna clade, O. borealis is distin-
guished by broadly ear-shaped apothecia, with a pale yellowish
ochre hymenium and a darker ochre yellow outer surface with
smallbrownwarts.Microscopically,thesporesareproportion-
ally broader than in closely related species, and the mycelium
atthebaseoftheapotheciaturnsbrightyellowinKOH(tested
indriedspecimenonly;Fig.30f).
At least four species of the O. concinna clade have bright
yellow outer apothecial colours, similar to O. borealis.Otidea
concinna appears morphologically most similar, but besides the
citrine-yellow outer receptacle, it differs in the narrower spores
(Fig.29)withhigherQmvalue.Otidea phlebophora and O. minor
are distinguished by the often entire apothecia with ribs or
anastomosingveinsatthebaseanddistinctlynarrowerspores.
Otidea oregonensis differs from O. borealis in typically having
5
5.2
5.4
5.6
5.8
6
6.2
6.4
6.6
6.8
7
10.1 10.6 11.1 11.6 12.1 12.6
O. oregonensis
O. minor
O. concinna
O. borealis
O. sp.
‘a’
O. phlebophora
O. kauffmanii type
O. rainierensis type
O. oregonensis
O. concinna
Lm (µm)
Wm (µm)
Fig. 29Meansporelengthandwidthincollectionsofspeciesinthecoreof
the O. concinna clade,basedon20sporesfromeachcollection.

214 Persoonia–Volume35,2015
obconical apothecia, sometimes with strongly rugulose base,
citrine-yellow colour in the outer receptacle when young, nar-
rowerspores (Fig. 29),andthemycelium atthebaseof the
apotheciadoes notturnyellow inKOH.TheNorthAmerican
O. rainierensis(= O. kauffmanii )appearstohavesomeyellowish
apothecialpigment(seefurtherunderO. rainierensis).Never-
theless, the spores of O. rainierensis are longer than the spores
of O. borealis(Fig.29).Furthermore,ourITS-LSUphylogenies
(Fig.3)suggestthatthesespeciesaredistinctfromO. borealis.
TwoSwedishcollections,MK0942andMK1081,constitutean-
otherundescribedspecies(Otidea sp.‘a’)nestedwithinthe
O. concinna group.ItresemblesO. borealis in spore shape
(onlyslightlylonger inOtidea sp.‘a’:11–12(–13) ×6–7µm;
Lm=11.2–11.6µm,Wm=6.6– 6.8µm,Qm=1.6–1.7;n=2)and
yellowreactionofthebasalmyceliuminKOH.ThetwoOtidea
sp.‘a’collectionsdifferfromO. borealis in the brownish ochre
receptacle surface, without yellow tones, and ochre to pinkish
ochrehymenium.Also our analyses of the ITS-LSU regions
support O. sp.‘a’andO. borealisasdistinctspecies(Fig.3).
Since only dried material has been available to us, we postpone
a formal description of O.sp.‘a’untilfreshandmoreabundant
material,withgoodcolourphotographs,becomesavailable.
Fig. 30 Otidea borealis(S-F242694,holotype)†.a.Sporesinwater;b.paraphysesandasciinKOH;c.ectalexcipuluminwater,withresinousexudatesonthe
outside;d.ectalexcipulumturningbrightyellowinKOH,resinousexudatespartlydissolving;e.basalmyceliuminwater;f.basalmyceliuminKOH.—Scale
bars=10µm;†=alldriedmaterial.

215
I.Olariagaetal.:AmonographofOtidea
26. Otidea caeruleopruinosaHarmaja,Karstenia48:37.2009
 —Fig.31
Holotype.Finland,Varsinais-Suomi,Lohja,Jalassaari,20Sept.1978,
H. Harmaja(H6010805)!
Apotheciagregarioustocaespitose,31– 60mmhigh,17– 50
mm wide, initially ear-shaped, upper margin rounded, then cup-
shaped,split,marginsometimeslobulate,stipitateorsessile.
Hymeniumyellowishbrown(5C6),whendriedyellowishbrown
withafaintolivaceoustint(4D6)ororange-ochre(6B6,6C6).
Receptacle surface upper half concolorous with hymenium or
slightlyduller(5C5,5C6),lowerhalfmorewhitish(pubescent
pruinose),slightlyhygrophanous,whendriedconcolorouswith
hymenium towards the margin, below cream, or pale reddish
brown(6C6),furfuraceoustoveryfinelywarty,seldomslightly
ribbedatthe base. Wartsbroadlyconical, gregarious, paler
orbluish.Stipe not well developed, rooting, up to 11 ×5mm.
Basal tomentum and mycelium abundant, when fresh ochra-
ceouswhite,when dried light ochre (5A2). Spores narrowly
ellipsoid to ellipsoid, often inequilateral, with two large guttules,
smooth,hyaline,(10.5 –)11–12.5×5.5– 6.5µm(Lm=11.2–11.8
µm,Wm=5.9 – 6.1µm,Qm=1.9;n=3). Paraphyses straight
or curved, or broadly hooked, of the same width or broader at
apices,2–4(– 5)µmwide,withoutnotches,terminalsegment
26–78µmlong,whenfreshcontainingfew,small,weaklyrefrac-
tive,paleguttules;whendriedrefractive,hyalinegranules.Asci
144–200×9–11µm.Apothecial section700 – 950µmthick.
Subhymeniumc.70 – 80µmthick,of2.5 –5µmwidecylindrical
cells,withscatteredswollen,12 –18µmwidecells,denselyar-
ranged.Medullary excipulum of textura intricata,550–700µm
thick,hyphaethintothick-walled,3– 9µmwide,intercalatedwith
swollencells,12 –18µmdiam,hyaline,withoutresinousexu-
dates.Ectal excipulum of textura prismatica-angularis,100–120
µm,cellsthin-walled,palebrown,sometimesyellowinKOH,
10–32× 10 – 22µm.Surfacewithconicalto broadly conical
warts,60–80 µm high, formedbyshort,fasciculatehyphoid
hairs, of 2 –4 ovoid to cylindrical cells, 6 –10 µm wide, not
constrictedatsepta.Non-wartedpartswithsingle2–7-celled
hyphoidhairs,ofglobosetocylindricalcells,20–33 × 9 –17
µm.Resinous exudates abundant, yellow toreddish brown,
dissolvinginMLZ,brightyellowandpartlydissolvinginKOH
andtheouterexcipulumturningbrightyellow.Basal mycelium
of3.5 – 6(–8)µm widehyphae,with spheroidtorod-shaped,
reddishyellow resinous exudates,dissolving in MLZ,more
slowlyandpartiallydissolvinginKOH,turningyellow.
Specimens examined. Spain,Girona,Ripollés,Setcases,underCorylus
avellana, Betula verrucosa and Buxus sempervirens, oncalcareoussoil,26
Aug.2010,M. Tabarés & S. Santamaría, MT10082601(dupl.S).–SwEdEn,
Södermanland,Nynäshamn,Herrhamra,onsoilunderFagus, in narrow for-
estareaalongtheroad,18Sept.2013,I.-L. Walter,KH.13.48(S);Uppland,
Älvkarleby,Vfromthetrainstation,24Sept.1950,G. Fåhraeus & G. Stenlid
(UPSF-146664).
 Notes—ThemaindiagnosticcharactersofO. caeruleoprui-
nosa are cup-shaped, split apothecia, whitish to pale cream hy-
menium,greyoutsideandrathersmallspores.Harmaja(2009a)
described O. caeruleopruinosa with strong emphasis on the
bluish shades of the warts on the apothecial outside, a charac-
ter that has not been observed in our Swedish, or the Iberian
material(VanVoorenetal.2011).Athirdgatheringfromthe
Iberiansitedoesnotshowbluishtoneseither(M.Tabarés,pers.
comm.).
Fig. 31 Otidea caeruleopruinosa.a.Apothecia;b.apotheciumshowingbasaltomentum†;c.sporesinwater†;d.paraphysesinwater †;e.ectalexcipulumin
water†(a:MT11080205;b– e:H6010805,holotype).—Scalebars=10µm;†=driedmaterial.—Photos:a.M.Tabarés;b.J.Kearey.

216 Persoonia–Volume35,2015
Otidea caeruleopruinosa belongs to the O. concinna clade, and
it shares several characters with the other members of the
clade, such as cup-shaped apothecia, small spores, and an
ectal excipulum of textura prismatica to textura angularis. It is
morphologically most similar to O. flavidobrunneola(seeNotes
underthatspecies).
27. Otidea concinna(Pers.)Sacc.,Syll.Fung.8:96.1889—
 Fig.32
Basionym.Peziza concinnaPers.,Mycol.Eur.1:221.1822:Fr.,Syst.
Mycol.2:49.1822.
 ≡Helvella scutellata Schaeff., Fung.Bavar.Palat.Nasc. 4:101. 1774
(‘Elvela’).
Lectotypedesignatedhere:Schaeffer,Fung.Bavar.Palat.Nasc. 2:t.150,
f.1.1763(‘Elvela tertia’).Epitype designated here: SwEdEn, Uppland, Stock-
holm, Naturhistoriska riksmuseet, in front of the Botany building, under a
big Quercus robur tree, by a row of Populus,20 Sept.2009, K. Hansen &
I. Olariaga,KH.09.183(S);MycoBankMBT178084.
= Helvella pyxidata Schaeff.,Fung.Bavar.Palat. Nasc. 4: 111.1774
(‘Elvela’).
 ≡Peziza marsupium var.pyxidata(Schaeff.)Pers.,Syn.Meth.Fung.2:
640.1801(‘ß pyxidata’).
 ≡Scodellina pyxidata (Schaeff.)Gray,Nat.Arr.Brit.Pl.1:669.1821.
Misapplied names
– Flavoscypha cantharellasensuHarmaja,Karstenia14:107.1974.
Apotheciagregarioustocaespitose,17–71mmhigh,12– 66mm
wide, initially broadly ear-shaped, then soon expanding and be-
coming deeply cup-shaped, split, often broader above, very
rarelyentirewhenyoung,stipitateorbroadlysessile.Hymenium
paleyellowishochre(3A3,4A5)topalegreyishochre(5A2),
Fig. 32 Otidea concinna*.a, b.Apothecia; c.spores;d. paraphyses;e.ectal excipulum;f.basal mycelium(a,c, e,f:KH.09.183, epitype;b:KH.09.217;
d:KH.09.176).—Scalebars=10µm;*=allfreshmaterial.

217
I.Olariagaetal.:AmonographofOtidea
sometimes with rose stains or spots, when dried light yellowish
grey(2A2)toverypaleyellowishochre(3A2,3A3).Receptacle
surface brightcitrineyellow(3A5– 3A7),sometimeswithagreen-
ishhue(2A6),slightlyhygrophanous,indryingslightlypaler,
fadingtobrownishochre(4B5)inage,whendriedyellow(3A7),
furfuraceous, sometimes warty in young apothecia, sometimes
shallowlywrinkledat thebase.Wartsflattobroadlyconical,
leavingareticulumamongthem,concolorous.Stipe not well
developed,rooting,7–8×4–5mm.Smellsweet,fruity,weak;
tastemild.Basal tomentum and mycelium abundant, white to
verypalecream(4A2),sometimeswithverypalepurplishtones,
whendriedwhitetoverypaleyellow(4A2).Spores narrowly el-
lipsoid to ellipsoid, inequilateral, with two large guttules, smooth,
hyaline,(10–)10.5 –12×(5–)5.5–6.5µm(Lm=10.8–11.2µm,
Wm=5.6–6µm, Qm=1.8 –2; n=8). Paraphyses straight to
bent, seldom broadly hooked, at apices claviform to abruptly
capitate,2.5– 6(–8.5)µmwide,withoutnotches,terminalseg-
ment19–66µmlong,whenfreshcontenthyalinehomogenous
orof small,refractive,hyalinetopaleyellowgranules; when
driedrefractive, hyaline granules. Asci 176 –196 ×10–11.5
µm.Apothecial section1200 – 2100 µm thick.Subhymenium
c.60–90µmthick,composedofcylindricalcells,2–4µmwide,
withscatteredswollencells,9 –13µmwide,denselyarranged.
Medullary excipulum 600 –1000(–1500) µm thick, of textura
intricata,hyphaecylindricaltoslightlyswollen,3 –11(–15)µm
wide,thintothick-walled,hyaline,withoutresinousexudates.
Ectal excipulum of textura prismatica-angularis, 80–100 µm
thick, cells thin-walled, hyaline to very pale yellow, sometimes
light reddish in KOH, 12 –35 × 6 – 22 µm. Surface with low
broadlyconicalwarts,60 –70µmhigh,formedbyfasciculate,
parallel,short,hyphoid hairs, of4–6ovoidcells, constricted
atsepta,5 – 9.5µmwide.Non-wartedpartswithsingle,2– 6-
celled, hyphoid hairs, uppermost cell narrowly claviform to
subcapitate,20–45×4.5–7µm,sometimeswithagelatinous
sheath.Resinousexudatesabundant,yelloworreddish,dis-
solvingintoamberdropsinMLZ,turningbrightyellowinKOH.
Basal mycelium of cylindrical to slightly swollen, hyaline to very
paleyellowhyphae,3– 4.5(–8.5)µmwide,yellowinKOH,with
scattered, spheroid to rod-shaped, yellow or reddish resinous
exudates,dissolvinginMLZ,partiallyinKOH.
Specimens examined. dEnmark, NE Sjælland, JægersborgDyrehave,
8Sept.1979,H. Knudsen(C-F-87186);NWJylland,NystrupPlantage,‘Kridt-
stien’,oncalcareoussoilwithAbies,14Sept.1985,T. Læssøe(C-F-81617).
–EStonia,Saaremaa,Abruka,adterram,16Sept.1966,K. Kalamees(UPS
F-629562).–Finland, Varsinais-Suomi, Turku,IspoinenKatariinanlaakso,
Wpartofthereserve, in rich, essentially deciduous woods with Corylus, Quer-
cus, Tilia,etc.onbareclayeymullsoil,22Aug.1977,H. Harmaja(S-F249360,
ex-H6015773).–FranCE, Saône-et-Loire,LaGrande-Verrière,Senavelle, on
the ground under Pseudotsuga menziesii,12Sept.2008,J.-P. Dechaume,NV
2008.09.13(dupl.S).–Spain, Huesca,Javierregay,humusofQuercus rotun-
difolia and Q. humilis,5Dec.2009,F. Prieto & A. González,KH.09.250(S);
Huesca,SallentdeGállego,Lanuza,14Oct.2006,I. Olariaga (BIO-Fungi
13002);Huesca,YebradeBasa,closetoStaOrosiachapel,inFagus syl-
vatica forest,11Oct.2009,P. Siljeström(ARAN-FungiA8700091).–SwEdEn,
Gotland,Ala,Stenstugu,nearabigQuercus robur tree in meadow, on rich
ground,amonggrass,26Sept.2009,E. Bohus-Jensen, K. Hansen & I. Ola-
riaga,KH.09.217(S);Gotland,Lojsta,Lojstaslot,underCorylus on a chalk-
richslope,19Sept.2000,T. Knutsson,TK2000-078(S);Närke,Havsta,Brun-
torpskärret,underconifers,10Aug.2008,B. Wasstorp,JS.08.59(S);Skåne,
Degeberga,Segesholm,onasteepslopeonbareground,underFagus and
Ulmus,23Sept.2010,K. Hansen, K. Gillen, I. Olariaga,KH.10.256(S);
Skåne,Fjälkstad,Balsberget,onrichgroundamongleaflitterunderFagus,
20Sept.2010,K. Hansen, K. Gillen & I. Olariaga,KH.10.212(S);Skåne,
Kristianstad, N. Lingenäset, Storskogen, on ground in broadleaf forest,
18Sept.1996,S.-Å. Hansons.n.(C);Uppland, Uppsala,Carolinaparken,
under Corylus avellana, 7Sept.2010, S. Ryman, K. Hansen, K. Gillen &
I. Olariaga,KH.10.180(S); Uppland, Uppsala, Hågadalen-NåstenNature
Reserve,Predikstolen,underQuercus robur, Ulmus glabra, Sorbus aucu-
paria, with Hepatica nobilis, Geum urbanum, Convallaria majalis, on rich
ground,19Sept.2009,K. Hansen & I. Olariaga,KH.09.176(S);ibid.,6Oct.
2010,KH.10.182(S).
Notes — Otidea concinna is characterised by cup-shaped,
split apothecia, citrine yellow receptacle surface, along with para-
physespredominantlystraightandclaviformatapices.Otidea
minor and O. phlebophora share with O. concinna a yellow
receptacle surface, but they have anastomosing ribs at the
apothecialbaseandnarrowerspores(Fig.29).Otidea orego-
nensis is distinguished from O. concinna by the often obconical
apothecia,sometimeswithbluntribsatthebase.
Thebrightyellowreceptaclesurfaceisoneofthemaindiagnos-
tic characters of O. concinna, but we have observed the yellow
colour can disappear with age, and the outer surface becomes
brownishochre(Fig.32b)thusmakingitdifficulttorecognise
thespecies.Otidea rainierensis is a North American taxon that
shows an apothecial shape similar to O. concinna.Neverthe-
less, bright citrine yellow tones are absent in O. rainierensis,
and our molecular data support it as distinct from O. concinna
(Fig.1,3).ForfurthercommentsseeunderO. rainierensis.
Nomenclatural notes — No authentic material has been
locatedinPersoon’sherbariuminL.Wethereforeselectone
elementinSchaeffer’splate(t.150,f.1)asthelectotype,be-
causeitmostcloselyresemblePersoon’sdescription,showing
two or more large, convolute, caespitose, sessile, cup-shaped
apothecia with a citrine (although light) outer surface. Per-
soon(1822)collectedthespeciesin‘sylvulaVincennesprope
Parisios’andSchaeffer’splateisfromBavaria,Germany.Fries
sanctionedthename,butdidnotseematerial(‘v. ic.’,seenfrom
icones).Nevertheless,weselectanewlycollectedepitypefrom
Sweden(Fig.32a,c,e,f),associatedwithQuercus(thehabitat
describedbyPersoon)andbackedbysequencesofmultiple
generegions(ITS,LSU,PRB1,RPB2andEF1)ofO. concinna
collectionsfromsouthernEurope(Spain)andfromtheepitype
beingidentical(Hansen&Olariaga2015),acknowledgingthe
speciesiswidelydistributedinEurope.
28. Otidea flavidobrunneolaHarmaja,Karstenia48:38.2009
 —Fig.33
Holotype. Finland, Varsinais-Suomi,Lohja, Jalassaari,Ahtiala, Alho,
AhtialaNatureReserve,rich,predominantlydeciduous(Quercus, Corylus
etc.)woodsoncalcareoussoils,20Sept.1978,H. Harmaja(H6010806)!
Apothecia gregarioustocaespitose,15–70mmhigh,8–50mm
wide, initially narrowly to broadly ear-shaped, then cup-shaped,
split,marginsometimeslobulate,shortly stipitate or sessile.
Hymeniumcream or pale yellowish (4A2, 4A3), whendried
orange-ochre(5B6).Receptacle surface pale brownish ochre
(5A3,5A4),slightly hygrophanous, when dried darkreddish
brown(5E5),furfuraceoustoveryfinelywarty,seldomshallowly
wrinkledat the base.Wartsflattened, gregarious, concolor-
ous.Stipenotwelldeveloped.Basal tomentum and mycelium
abundant,whitetopalecream(4A2)whenfresh,ochre(5A4)
toorange-ochre(6A4)whendried. Spores narrowly ellipsoid
to ellipsoid, inequilateral, with two large guttules, very rarely
withathirdsmallguttule,smooth,hyaline,9.5–11×(4.5–)5 – 6
µm(Lm=10–10.6µm,Wm=5.1– 5.5µm,Qm=1.9–2;n=5).
Paraphyses curved, a few straight or broadly hooked, of the
samewidthorslightlybroaderatapices,3–4µmwide,without
notchesorslightlysinuousunderside,uppermostcell40–76µm
long,whenfreshcontainingyellowguttules;whendriedhyaline
guttules or granules. Asci 127–170 × 9 –10 µm. Apothecial
section900–1900µmthick.Subhymeniumc.100µmthick,of
dense textura intricata,visibleasayellowishzone,cellscylin-
dricaltoslightlyswollen.Medullary excipulum 600 –1300µm
thick, of textura intricata,hyphae5–13µmwide,thick-walled,
sometimes with a thinner outer wall, hyaline, without resinous
exudates.Ectal excipulum of textura prismatica, sometimes of

218 Persoonia–Volume35,2015
a textura angularis,100–120µm,cellsthin-walled,yellowish,
distinctlycitrineyellowinKOH,18–38×11–13µm.Surfacewith
broadlyconicalwarts,40 –70µmhigh,formedbyfasciculate,
shorthyphoidhairs,of3–4ovoidtoelongatedcells,constricted
atsepta,5– 9µmwide,sometimeswithagelatinoussheath.
Non-wartedpartswith2– 3-celledhyphoidhairs,withclaviform
uppermostcell, more rarelycylindrical, 33 – 55 × 8–13µm.
Resinousexudatesabundant,yellowishbrown,partiallydis-
solvingandturningslightlyreddishinMLZ,partlydissolving,
brightyellowandappearinggelatinousinKOH.Basal myce-
lium of4– 6µmwide,hyalinetoverypaleyellowhyphae,with
scattered swollen septa, with rounded to rod-shaped, yellow
resinousexudates,slowlydissolvinginKOH,quicklyinMLZ.
Specimens examined. Finland, Uusimaa,Nurmijärvi,parishcentre,under
Quercus, 16 Sept.1987, P. Askola 2220(H);Uusimaa,Nurmijärvi,parish
centre, clayey soil under Quercus,11Aug.1988,P. Askola2334(H);Uusi-
maa,Nurmijärvi,parishcentre,themarginoftheparkbythevicarage,under
Quercus,18Aug.1988,P. Askola2360(H).–norway,Nord-Trøndelag,Leks-
vik,Gjøråsvika,onrichground,onslopeunderCorylus and Picea, 3Sept.
2009,K. Hansen & I. Olariaga,KH.09.153(S).–SwEdEn, Uppland, Bondkyrka,
Vårdsätranaturpark,17Aug.1927,H. Svensson (UPSF-146554);Uppland,
Uppsala,infrontoftheprison,onbaregroundunderdeciduoustrees,14Sept.
1938,R. Gustafsson (UPSF-146718).
Notes — Otidea flavidobrunneola is macroscopically charac-
terised by cup-shaped, split apothecia, especially with age very
pale yellowish brown outside and cream-white to pale yellow
hymenium.Microscopically,thesmallsporesandtheyellow
Fig. 33 Otidea flavidobrunneola.a.Apothecia;b.apotheciumshowingbasaltomentum†;c.sporesinwater†;d.paraphysesinwater †;e.ectalexcipulumin
water†;f.ectalexcipuluminKOH†;g.basalmyceliuminwater †(a,c:KH.09.153;b,d – g:H6010806,holotype).—Scalebars=10µm;†=driedmaterial.—
Photos:b.J.Kearey.

219
I.Olariagaetal.:AmonographofOtidea
ectalexcipulumturningbrighteryellowinKOHarediagnostic.
Theapotheciaandbasaltomentumbecomecharacteristically
darker upon drying: the outside turns dark brown, the hyme-
niumandthebasaltomentumorangeochre(Fig.33b).Otidea
caeruleopruinosa and O. flavidobrunneola are morphologically
verysimilar,butdiffer inthesporesizes(Lm=11.2–11.8µm
vs10.1–10.6µm) and colours of the basal tomentum in dried
specimens.AccordingtoHarmaja(2009a),O. flavidobrunneola
is reminiscent of O. bufonia, but the latter has a darker brown
basaltomentumandlarger,narrowlyfusoidspores.Wereport
herethefirstfindsoutsideFinland.
29. Otidea lactea J.Z.Cao&L.FaninCaoetal.,Mycologia
 82:735.1990
Holotype.China,Heilongjiangprovince,YichunCity,onground(rotten
wood?)under broadleaftrees, 6Sept.1987,J.Z. Cao(HMAS 61359,ex-
MHSU1803).
 Notes—WewerenotabletogetthematerialofO. lactea
on loan, but it appears to be a distinct species based on the
entire, cup-shaped, cream white apothecia and paraphyses
withbroadly clavatetosubgloboseapices(Caoetal.1990).
Also,inourLSUphylogeny(includingaGenBanksequenceof
theholotype,DQ443447fromLiu&Zhuang(2006)),O. lactea
formsadistinctsisterlineage(ML71%,PP99%)toacladeof
seven collections of O. minorfromEurope(ML95%,PP98%,
Fig.1).Otidea minor differs from O. lacteainthe(mostoften)
split apothecia, with a yellow outer surface and often more
narrow, straight to bent, subclaviform to claviform paraphyses
apices.Otidea lactea has the characteristic broad apices of
the paraphyses of the O. concinna clade, which however, in
O. lacteabecomeshookedwithage(Caoetal.1990).Theholo-
typewasoriginallydepositedinMHSU(Caoetal.1990),but
latertransferredtoHMAS(confirmedbyHong-MeiLu,HMAS).
30. Otidea minor(Boud.)Olariaga&K.Hansen,comb. & stat.
nov.—MycoBankMB808975;Fig.34
Basionym.Otidea cantharellavar. minorBoud., Icon.Mycol.livr.23:n°.
411.1909(preliminarytextwith‘circulaires’).
 ≡Flavoscypha cantharellavar.minor(Boud.)Häffner,Rheinland Pfäl.
Pilzj.4:36.1994.
Lectotypedesignatedhere:Boud.,Icon.Mycol. livr.23:n°.411,pl.326.
1909;MycoBankMBT178086.
Misapplied names
– Otidea cantharellasensuLundell&Nannfeldt,FungiExs.Suec. 1–2:93.
1934.
– Flavoscypha cantharellasensuDennis,Brit.Ascomyc.:pl.8D.1978.
Apothecia gregarious to caespitose, 8 –33 mm high, 4 – 41
mm wide, broadly ear-shaped or shallowly cup-shaped, often
elongatedononeside,normallysplit,oftenstipitate.Hymenium
lightochre(3A2,3A3)toyellowishgrey(4A3),sometimeswith
weakrosestainsorspots,whendriedyellowishochre(4A4,
4A5)to ochre(5A4).Receptacle surface citrineyellow(2A6,
2A7),slightlyhygrophanous,indryingslightlypaler,fadingto
brownishochre(4B5,4B6)inage,whendriedreddishochre
(6B6–6D6)toreddishbrown(5B4,5B5),slightlyfurfuraceous
to slightly warty, some apothecia with low ridges coming from
thestipe,restrictedtothebaseorreaching1/2–1/3oftheapo-
thecium,sometimesinterconnectedbylowveins.Wartsminute,
conical,concoloroustobrownish.Stipe often well developed,
cylindrical,2–12 ×2–5mm.Basal tomentum and mycelium
abundant, white, seldom with purplish tones, very pale yellow
(4A2)whendried. Spores narrowly ellipsoid, often inequilateral,
withtwolargeguttules,smooth,hyaline,(10–)10.5–12.5(–13)
×(4.5–)5–5.5 (–6.2)µm(Lm=10.7–12.3µm,Wm=5 – 5.6µm,
Qm=2.1–2.3;n=10). Paraphyses straight to bent, seldom
curved,subclaviformtoclaviformatapices,3– 8µmwide,with-
outnotches,uppermostcell20 –71µmlong,whenfreshwith
homogeneous content or seldom containing slightly refractive,
light yellow granules at apices, often with a yellow refractive
body(Fig.34d); whendriedsmall,slightlyrefractive, hyaline
granules.Asci 153 –169 × 9 –10.5 µm. Apothecial section
750–1000µmthick.Subhymeniumc.100µmthick,ofdense
textura intricata,hyphae 3 – 6µmwide.Medullary excipulum
of textura intricata,500–700µmthick,hyphae4 – 9(–13)µm
wide, sometimes slightly swollen, thin to thick-walled, hyaline
toverypaleyellow,withoutresinousexudates.Ectal excipulum
of textura prismatica-angularis,80 –150µm,cellsthin-walled,
verypaleyellow,unchangedinKOH,10–33×6–15µm.Sur-
facewithlowflattenedwarts,35–53µmhigh,formedbyshort,
fasciculate,hyphoidhairs,of2–3ovoidcells,constrictedat
septa,6–9µmwide.Non-wartedpartswith2–4-celledhyphoid
hairs,withsubclaviformtoclaviformuppermostcell,43–58×
7.5–13µm.Resinous exudates abundant, yellow orreddish
tobrownishyellow,dissolvingintoamberdropsinMLZ.Basal
myceliumof3 –7 µm wide, verypaleyellow,slightly swollen
hyphae,notchanginginKOH,withveryscatteredamorphous
orrod-shaped,yellowresinousexudates,dissolvinginMLZ.
Specimens examined. dEnmark,Jylland,NystrupPlantage,Kridtstien,on
calcareous soil under Abies,1Nov.2007,T. Læssøe, TL-13332(C);NJylland,
RoldSkov,BuderupholmBjergskov,byfenceofCypripedium, calcareous soil
alongroadside,deciduousforest,14Sept.1998,K. Hansen, KH.98.84(C);
Jylland,Vorsø,under Salix caprea,25Sept.1981,T. Læssøe,TL-0684(C);
ibid.,onbaseofSalix caprea,8Sept.1982,T. Læssøe,TL-0754(C);Sjæl-
land,Geelskov,10kmNofCopenhagen,3Aug.1950,M. Lange(C-F-47967).
–Finland, Helsinki,Toukola,Koreankatu,Acer, Betula, Populus tremula, Salix
caprea, Sambucus, Sorbus, Aegopodium podagraria, Urtica dioica,8Sept.
1992,R. Saarenoksa 24592 (H);Varsinais-Suomi,Lohja,Pähkinäniemi,very
rich,somewhatdrygrass-herbforestwithcalcareousground,1Aug.1997,
U. Nummela-Salo & P. Salo 4051 (H);Varsinais-Suomi,Lohja, Virkkala,
NE-slopeofPähkinäniemi, veryrich,somewhatdrygrass-herbforestwith
calcareous soil, nearby Corylus avellana, Populus tremula, Betula pendula,
12Sept.2006,U. Nummela-Salo & P. Salo 10724(H).–italy, Calabria, Acri
(CS),CrocediGreca,14Sept.1995,C. Lavorato, CL950914-01(dupl.S).
– SwEdEn, Gotland, Fårö,Avanäset, under Pinus sylvestris, on humus,
sandysoil,27Sept.2011,J.C. Zamora,KH.11.103(S);Gotland,Lojstahed,
Russpark,grazedforestwithPinus sylvestris,oncalcareousground,2Oct.
2010, K. Hansen, K. Gillen & I. Olariaga,KH.10.311(S);Småland,Stenbro-
hult,StockanäsSSVofStenbrohultskyrka,underPyrus, Salix and Prunus
domestica,9Aug. 2011, G. Aronsson (UPS F-548414); Södermanland,
Södertalje,Mörkö,Oaxen, in rich soil in deciduous forest, Corylus avellana,
Salix caprea, with Lactarius citriolens,10Sept.1994,P. Höljer (H7003652);
Uppland, Uppsala, the park in front of the prison, on bare soil, among needles
etc.underAbies,16Aug.1932,S. Lundell(S-F108335,FungiExs.Suec.
93); Öland, Högby,Horns kungsgårds Nature Reserve, under Corylus,
6Aug.2000,T. Knutsson,TK2000-057(S).
Notes — Otidea minor is recognised macroscopically by
apothecia with a yellow outside, and shallow ribs and veins
atthebaseofatleastsomeapothecia.Otidea concinna and
O. oregonensis can be distinguished by the broader spores
with a lower Qm(Fig.29).InfreshmaterialofO. minor, some
paraphyses had a strikingly, yellowish refractive body in the
upper part, a so far unique feature within Otidea.Toassessits
taxonomic value, this feature has to be checked in additional
freshmaterialandincloselyrelatedspecies.
Otidea minor has been confused with O. phlebophora due to the
presenceofribsorveinsatthebaseoftheapothecia(Lundell&
Nannfeldt1934,Dennis1978).Harmaja(2009a)proposedthe
provisional name O. subconcinna for O. minor as circumscribed
here.After examining two collections annotatedby him, we
considertheFinnishmaterialtobeconspecificwithourSwed-
ishfinds,based on both morphological and moleculardata.
AlthoughHarmaja(2009a)didnotdirectlycompareO. minor
(asO. subconcinna)andO. phlebophora, he stated in the key
that O. phlebophora has ‘other tingesof yellow’and mostly
straightparaphyses. Wecouldnotconfirmthesedifferences
inthematerialweexamined.Instead,O. minor can be distin-

220 Persoonia–Volume35,2015
guished from O. phlebophora by the presence of at least some
apothecia without ribs, and ribs when present shallower and
lessanastomosing,aswellasnormallysplitapothecia.Inspite
of their morphological similarity, O. minor and O. phlebophora
are not sister species in our four-gene phylogeny, but both
are deeply nested within the O. concinnaclade (Hansen &
Olariaga2015;seealsoFig.3).Boudier´splateunderOtidea
cantharellavar.minor shows shallow ribs at the base of some
apothecia, agreeing with our material, and as considered by
Harmaja(2009a).NooriginalmaterialcouldbelocatedinPC,
andwethereforedesignateBoudier’splateasthelectotype.
31. Otidea oregonensisK. Hansen & Olariaga,sp. nov.—
 MycoBankMB808973;ITSbarcodeGenBankKM010048;
 Fig.35
Etymology. Named after Oregon, the area where most of the specimens
ofthisspecieshavebeencollected.
Holotype.USA,Oregon,DouglasCo.,UmpquaNationalForest,Diamond
LakeRangerDistrict,under Pseudotsuga menziesii and Abies concolor,7Nov.
2010, J. Moore, Moorefun58(OSC).Isotype(S).
Apotheciagregarioustocaespitose,12 – 48mmhigh,23 – 80
mm wide, shallowly to deeply cup-shaped, sometimes elon-
gated on one side or obconical, split, seldom entire, sometimes
stipitate.Hymeniumgreyishwhite(2A2)topaleochre(4A2–
4A4),sometimeswithrosestainsorspots,whendriedpale
ochre(4A2–4A4). Receptacle surface bright citrine yellow
Fig. 34 Otidea minor(KH.10.311)*.a.Apothecia;b.apotheciumshowingveinedouterreceptacle;c.spores;d.paraphysesshowingacrystallizedbody;e.ectal
excipulum;f.ectalexcipuluminMelzer’sreagent,resinousexudatesmostlywashedaway,showingfreehypoidhairs.—Scalebars=10µm;*=freshmaterial.

221
I.Olariagaetal.:AmonographofOtidea
(2A6–2A8),slightlyhygrophanous,fadingtobrownishochre
(4B5,4B6)inage,whendriedbrownishochre(4A4,4B4,5B4),
furfuraceous, sometimes some apothecia wrinkled-veined at
thebase,seldomwithshortribsreachingup1/3tothemar-
gin,partly coveredbywhitetomentum fromthebase.Warts
sometimespresenttowardthebase,minute,flat,concolorous.
Stipesometimeswelldeveloped,cylindrical,3 –18×3–11mm.
Smellmild.Basal tomentum and mycelium abundant, white,
whendriedverypaleyellow(4A2). Spores narrowly ellipsoid
to ellipsoid, sometimes inequilateral, with two large guttules,
rarelywith a fewsmall granules, smooth, hyaline, 10 –11.5
× 5.5 –6 (– 6.5) µm (Lm=10.4–11.3 µm, Wm=5.6 –5.9 µm,
Qm=1.8 –2; n=7). Paraphyses bent to curved, sometimes
straight,subclaviformtocapitateatapices,3– 6(–8)µmwide,
withoutnotches, uppermostcell28–70µmlong,whendried
containingsmall, weakly refractive,hyaline granules. Asci
171–203×9 –10.5µm.Apothecial section1000 –1400(–2000)
µmthick.Subhymenium100–140µmthick,ofdensetextura
intricata,hyphae2–4.5µmwide,withscatteredswollen,upto
12µmwidecells. Medullary excipulum 650–850(–1400) µm
thick, of textura intricata,hyphae5–9(–17)µmwide,some-
times swollen, thin- to thick-walled, hyaline, without resinous
exudates.Ectal excipulum of textura prismatica to textura
angularis,70–100µmthick,cellsthin-walled,verypaleyellow,
brighteryellowinKOH,16 –37×8 –14µm.Surfacewithlow
warts,35–70µmhigh,formedbyshort,fasciculate hyphoid
hairs,of2–4ovoidcells,constrictedatsepta,5 – 9µmwide.
Non-wartedpartswith2–4-celled,hyphoidhairs,withclaviform
Fig. 35 Otidea oregonensis.a.Apothecia;b.apotheciashowingfadedcolourintheouterreceptacle;c.sporesinwater†;d.paraphysesinwater †;e.ectal
excipulum in water†;f.ectalexcipuluminKOH†(a,c – f:Moorefun58,holotype;b:rh139).—Scalebars=10µm;†=driedmaterial.—Photos:a.J.Moore;
b.R.Helliwell.

222 Persoonia–Volume35,2015
uppermostcell,33–55×7–12µm.Resinousexudatesoften
abundant,yellow,dissolvingintoamberdropsinMLZ,brighter
yellowinKOH.Basal mycelium of3–5(– 8)µmwide,sometimes
slightlyswollen,verypaleyellowhyphae,notchanginginKOH,
with very scattered, spheroid to rod-shaped, yellow resinous
exudates,dissolvinginMLZ.
Specimens examined. USA, Oregon,JacksonCo.,RogueRiverNational
Forest, under Pseudotsuga menziesii, Abies concolor, Pinus ponderosa,
2Dec.1999,B. Schroeter (OSC 72950); ibid., 27 Oct. 1990, D. McKay,
NSW6354(OSC132740,dupl.S);Oregon,JosephineCo.,BureauofLand
Management,Medford District,GrantsPass ResourceArea,LittlePickett
Creek, under Pseudotsuga menziesii, Calocedrus decurrens, Lithocarpus
densiflorus, Quercus chrysolepis,19Dec.2000,R. Meyer (OSC108041);
Oregon, Marion Co., near Breitenbush Hot Springs Community,27 Oct.
1996,J. Trappe(OSC56745);Oregon,Umpqua NF,Diamond Lake RD,
under Pseudotsuga menziesii,2Nov.2010,J. Moore, Moorefun31(S);ibid.,
old growth forest with Abies concolor, Pseudotsuga menziesii,2Nov.2010,
R. Helliwell, rh139(OSC);Washington,LewisCo.,GiffordPinchotNational
Forest,CampCreeksFallsTrail,6Nov.1997,M. Castellano (OSC56829).
Notes — Otidea oregonensis is characterised by a citrine
yellow receptacle surface, often a wrinkled to veined, or shal-
lowlyribbedapothecialbase,andrelativelybroadspores.The
apothecia are typically obconical cup-shaped, with a narrow
base.Otidea oregonensis was treated as O. rainierensisKa-
nousebyPeterson(1998),primarilybasedonthepresenceof
swollenapicesoftheparaphyses.Also,oneoftheparatypes
of O. rainierensis (EGS2179),sequencedbyPeterson(1998),
is nested within the O. oregonensis lineagein our ITS-LSU
phylogeny(Fig.3).Nevertheless,oursequencesoftheholotype
of O. rainieriensis, from four different gene regions, show it
belongs to another lineage, well separated from O. oregonensis
(Hansen&Olariaga2015).Kanouse´sconceptof O. rainier-
ensis wasthereforemixed.TheITS-LSUphylogeny,including
a larger sampling of the O. concinna clade, likewise resolves
O. oregonensis asadistinctspecies.
Otidea phlebophora and O. minor, so far only known from
Europe, resemble O. oregonensis in the cup-shaped apo-
theciawith yellow outsideand swollen paraphyses.Otidea
phlebophora differs macroscopically from O. oregonensis in
predominantly entire apothecia, with always a strongly veined-
ribbed base reaching up 1/2 to the margin, while only small
veins are present in some apothecia of O. oregonensis.Based
on the material examined here, the slightly narrower spores in
O. phlebophora (Qm=2–2.1vsQm=1.8 – 2inO. oregonensis)
are a constant and reliable character to separate these species
(Fig.29). Otidea minor tends to have smaller, shallower and
more broadly cup-shaped apothecia than O. oregonensis.The
spores of O. minor have a higher Qmvalue(2.1–2.3)thatdoes
not overlap with the Qm of O. oregonensis.
32. Otidea phlebophora(Berk.&Broome)Sacc.,Syll.Fung.
 8:97.1889—Fig.36
Basionym.Peziza phlebophora Berk.&Broome,Ann.Mag.Nat.Hist.,
ser. III, 18:122. 1866.
 ≡Flavoscypha phlebophora(Berk.&Broome)Harmaja,Karstenia 14:
107.1974.
Lectotype designated here: England,North Somerset, Brislington,16
Sept.1853(K(M)144045,exHerb.C.E.Broome)!;MycoBankMBT178087.
Apotheciagregarious,8 –12mmhigh,3– 40 mmwide,shal-
lowly to deeply cup-shaped, sometimes elongated on one side,
entire,seldomsplit,oftenstipitate.Hymeniumpaleochre(4A2,
4A3),sometimeswithrosestainsorspots,whendriedyellowish
Fig. 36 Otidea phlebophora (JV06-385).a.Apothecia; b.sporesin water†; c.paraphysesin water†; d.ectalexcipulum inwater †.—Scalebars=10µm;
†=driedmaterial.—Photos:a.J.Vesterholt.

223
I.Olariagaetal.:AmonographofOtidea
ochre(4B4,5B6,5C6)toreddishbrown(6C5,6C6).Recepta-
cle surfacebrightcitrineyellow(3A7,2A7),sometimeswitha
greenishhue(2A6),slightlyhygrophanous,fadingtobrownish
ochre(4B5)inage,whendriedyellowishbrown(5B6,5C6)to
reddishbrown(6C6,6D6),furfuraceous,withhighridgesinthe
basalpartofallapotheciareachingup1/2–1/3tothemargin,
interconnected by veins, partly covered by white basal tomen-
tum.Wartsminute,flattorounded, appressed, concolorous.
Stipeoftenwelldeveloped,cylindrical,1–10×1.5–3mm.Basal
tomentum and myceliumabundant,white,verypaleochre(5A2)
whendried. Spores narrowly ellipsoid, sometimes inequilateral,
withtwolargeguttules,smooth,hyaline,(9.5–)10–11.5(–12)
×(4.5–)5 – 5.5(–6.2) µm (Lm=10.2 –11µm, Wm=4.9 –5.1
µm, Qm=2–2.1;n=5).Paraphyses straight to bent, cylindri-
caltoclaviform at apices, 2 – 3.5 µm wide, without notches,
uppermostcell23 – 46µmlong,whendriedcontainingsmall,
ratherrefractive,hyalineguttules. Asci 130 –153 × 8 – 9 µm.
Apothecial section650 –1000µmthick.Subhymenium80 –100
µm thick, of dense textura intricata, hyphae 2 –3 µm w ide.
Medullary excipulum of textura intricata,450–650µmthick,
hyphae4 –7(–14)µmwide,sometimesswollen,thick-walled,
hyalineto verypaleyellow,withoutresinousexudates.Ectal
excipulum of textura prismatica-angularis,80–100µmthick,
cellsthin-walled,hyalinetoverypaleyellow,unchangedinKOH,
15–21×6–11µm.Surfacewithlowflattenedwarts,40 – 60µm
high,formedbyshort,fasciculatehyphoidhairs,of3–4ovoid
cells,constrictedat septa, 8 –12µmwide.Non-wartedparts
with2 – 3-celledhyphoidhairs,withclaviformuppermostcell,
30–60×7–13µm.Resinousexudatesabundant,yellow,dis-
solvingintoamberdropsinMLZ.Basal myceliumof3.5 –7µm
wide, slightly swollen, very pale yellow hyphae, not changing
inKOH,withveryscattered, spheroid to rod-shaped, yellow
resinousexudates,dissolvinginMLZ.
Specimens examined. dEnmark, NEJylland, RubjergKnudePlantage,
under Abiesinlargefairyring,28Aug.2006,J. Vesterholt & L. Vesterholt,
JV06-385(C);NESjælland,RavnsholtsHegn,underPicea abies,30Aug.
1999,B.W. Pedersen (C-F-71506).– Finland, Varsinais-Suomi,Lohja,
Jalassaari,Alho, by theAhtialamanor, in rich forestwithQuercus robur,
Corylus avellana, Prunus padus, cultivated Larix sibirica and Betula,30Aug.
1967,H. Harmaja (H6010675).–SwEdEn,Dalarna,Husby,Husbyvärdhus,
on lawn under Quercus, Tilia, Larix and Acer, 31Aug.1958,R. Morander
(UPSF-144691);Skåne,Helsinborg,JordbodalenbyHarlyckan,onsandy
ground in deciduous forest under Quercus,13July1995,S.-Å Hanson, SÅH
30601(C);Uppland,Djurö,Runmarö,Södersunda,inthecity,ontheground
under Syringa,18Sept.1949,G. Haglund & R. Rydberg (S-F108338,UPS
F-144689);Uppland,Uppsala,Sunnersta,Almlund,calcareous,humus-rich
clay,23Aug.1986,J. Nitare(UPSF-119845).–UK,Northampshire,King´s
Cliffe,onsoil,1853,M.J. Berkeley(K(M)144046,syntypeofP. phlebophora);
NorthSomerset,Brislington,TheBeeches,Sept.1853,C.E. Broome(K(M)
194582,syntypeofP. phlebophora).
Other specimen examined. Otidea integra — italy, Sopramonte,1892,
G. Bresadola(S-F108342).Localitynotspecified,insilvismixtis,1892,G. Bre-
sadola(PC124965).
Notes — Otidea phlebophora is primarily characterised by
highanastomosingribsandveinstowardstheapothecialbase.
Diagnosticcharactersareinaddition,thepredominantlyentire
apothecia,withacitrineyellowoutside.Foracomparisonwith
O. concinna, O. minor and O. oregonensis see Notes under
thosespecies.Harmaja(1986)elevatedO. integra to species
rank based on smaller apothecia and broader, curved paraphy-
ses.ThelikelyoriginalspecimenofO. integracollectedin1892
andkeptinBresadola’sherbarium(S-F108342),showscurved
paraphysesup to 5 µmbroad, the same asHarmaja (1986,
2009a)gaveforO. phlebophora.TheparaphysesintheO. phle-
bophora material examined by us were straight to bent, and
otherwise similar to the original material of O. integra.Inspiteof
themorphologicalsimilarity,theITS2sequenceofO. integra(281
bpobtained)isdifferentfromtheITSsequencesofthematerial
assigned to O. phlebophora by us, and we preliminary accept
O. integraasaseparatespecies.ThepositionofO. integra is
withoutsupportinourITS-LSUphylogeny(Fig.3).
 Nomenclaturalnotes—Harmaja(1974:107)indicatedalec-
totype of O. phlebophora atK,buthegavenocollectionnumber
andthetypificationwasnotachieved.Wehavestudiedthree
offoursyntypesatKewandselectheretherichestcollection
containingten apotheciaasthelectotype.Itconformsto the
current interpretation of the name, with several entire apothecia
(i.e.withoutasplit),aribbed-veinedbaseseenontwoofthe
apothecia(the otherswiththebasegluedtothecardboard),
spores in the range 9.5 –11.7 × 4.7– 5.9 µm (Lm=10.6 µm,
Wm=5.1µm),paraphysesstraightandenlargedatapices,and
abundant yellow resinous exudates in the ectal excipulum that
dissolveintoamberdropsinMLZ.
33. Otidea rainierensisKanouse,Mycologia41:674.1949
Holotype.USA,Washington,PierceCounty,LowerTahomaCreek,MT
RainierNationalPark,23Aug.1948,A.H. Smith 30553(MICH14410)!
= Otidea kauffmanii Kanouse,Mycologia41,6:673.1949.
Holotype.USA,Michigan,Lakeland,18July1915,C.H. Kauffman (MICH
14409)!
Kanouse(1949)onlyhaddriedmaterialofO. rainierensis and
describedthe outsideoftheapothecia as‘ochraceousbuff’,
‘cinnamonbuff ’to‘woodbrown’andthehymeniumas‘avel-
laneous’,‘vinaceousbuff’to‘drabgrey’.ForO. kauffmanii she
had notes on fresh material and she separated it from O. rain-
ierensis based on the presence of yellow tones in the apothecia
(outside‘chamois’to‘ochraceous’,hymenium‘creambuff’).We
believe O. rainierensis does have yellow apothecial tones, as
weobservedsmallyellowresinousexudates(inwater)inthe
type collection of O. rainierensis, and as observed in closely
allied species the yellow colour can fade and almost disappear
withage(Fig.32b).Differentspore sizes were also used to
distinguishthetwospecies.ThesporesoftheholotypeofO.
kauffmanii are however, larger than noted in the protologue,
11.5–12.5 × 5.5–6.5 µm, Lm=11.9µm, Wm=6 µm, Qm = 2
(spores8–10 (–12)×5–6 (–7)µmintheprotologue),andthus
overlapping with the spores of the holotype of O. rainierensis,
11–12×6.5–7µm,Lm=11.7,Wm=6.7µm,Qm=1.7.Wesug-
gest O. kauffmanii and O. rainierensis constitute a single spe-
cies, based on molecular and morphological study of the type
material(Fig.3).OurITSsequencesoftheholotypesdifferby
4bp.SeealsocommentsonO. microspora under Excluded,
dubiousand imperfectly knowntaxa. Otidea rainierensis is
characterised by a smooth apothecial base, long and relatively
broad spores, compared to the rest of the species in the O. con-
cinna clade, and by paraphyses with abruptly enlarged, broadly
clavatetogloboseapices.
EXCLUDED, DUBIOUS AND IMPERFECTLY KNOWN
TAXA
Cochlearia(Cooke)Lambotte,Mém.Soc.Roy.Sci.Liege,ser.
2.14:323.1888
 Nom.illegit.Art.53.1,non CochleariaL.,Sp.Pl.2:647.1753(Cruciferae).
Basionym. Peziza subg. Cochlearia Cooke, Mycographia part 6: 252
(‘IndexSystematicus’).1879.
 Notes—Eckblad(1968)selectedPeziza cochleata as the
type species for the genus Cochlearia considering it a synonym
of Otidea.However,Rifai(1968)hadalreadyselectedPeziza
aurantia as the type species, herewith making Cochlearia a
synonym of Aleuria.He felttoomuchdoubthadsurrounded
the identity of P. cochleata and it better not be selected as the
typespecies.

224 Persoonia–Volume35,2015
Otidea abietina(Pers.)Fuckel,Jahrb.NassauischenVereins
Naturk.23–24:330.1870‘1869–1870’
Basionym. Peziza abietinaPers.,NeuesMag.Bot.1:113.1794:Fr.,Syst.
Mycol.2:47.1822.
≡ Discina abietina(Pers.)Rehm,Rabenh.Krypt.-Fl.,ed.2,3:977.1896.
 ≡Pseudotis abietina(Pers.)Boud.,Icon.Mycol.listprél.600sp.:3(un-
numberedpage).1904.
 Notes—Nannfeldt(1966)statedthattheoriginalmaterial
belongs to Peziza badia,butHarmaja (2009a)andCarbone
(2010c)reportedtwocollectionsinPersoon’sherbariumunder
O. abietina representing O. propinquata and a third collection
O. bufonia.IftypifiedwithelementsbelongingtoeitherO. pro-
pinquata or O. bufonia, the name O. abietina would take priority
overany ofthose.Otidea abietina is the type species of the
genus Pseudotis(Boud.)Boud.TheidentityofPseudotis will
thus remain open until O. abietina isclarifiedortypified.Another
way to typify O. abietina would be to select an element belong-
ing to Peziza badia.However,thischoiceshouldbe studied
more thoroughly, since it would make Pseudotis available as
a genus name for the Peziza depressa-Ruhlandiella lineage, if
Pezizaissplitintosmallergenerainthefuture(Hansenetal.
2005).ForthetimebeingweregardO. abietina as a nomen
ambiguum,asseveralothers(Harmaja2009a,Carbone2010c,
Parslow&Spooner2013).
Otidea albaVelen.,Monograph. Discom. Bohemiae 1: 354.
1934
Holotype.CzECh rEpubliC,Karlštejn,Sept.1924,Fechtner(PRM149788)!
 Notes—Theholotype specimen has the typical oblong
spores of the O. alutaceacomplex,(13.5–)14.5–16.5(–17.5)
×6.5–7.5µm(Lm=14.9µm,Wm=6.9,Qm =2.1).Judgingfrom
thesporesize,O. alba might represent either O. alutaceas.str.
or the O. alutaceaclade3b.Itshouldbeconsideredinfuture
studiesofthecomplex.
Otidea aurantia(Pers.)Massee,Brit.Fungus-Fl.4:448.1895
Basionym.Peziza aurantia Pers.,Observ.Mycol.2:76.1800:Fr.,Syst.
Mycol.2:49.1822.
 Notes—This is the type species of Aleuria, A. aurantia
(Pers.:Fr.)Fuckel.
Otidea aurantiavar. atromarginata (W. Phillips & Plowr.)
Massee,Brit.Fungus-Fl.4:449.1895
Basionym.Peziza aurantia var. atromarginata W.Phillips&Plowr.,Gard.
Chron.17:191.1882.
 Notes—Thebloodreddiscwithshort,obtuse,darkbrown,
3–4septatehairs,givingthemarginadarkappearancesuggest
this may be a species of Melastiza.Thesporesaretuberculate
withthread-likeappendages.
Otidea aurantiavar.stipitata (W.Phillips)Massee,Brit.Fungus-
Fl.4:448.1895
Basionym.Peziza aurantiavar.stipitataW.Phillips,Man.Brit.Discom.:
57.1887.
 Notes—Thesmall,bright scarlet apothecia with a‘stem
equallingthe height of the cup,4 mm’and the ornamented
spores suggest this is a species of Aleuria or Sowerbyella.Itwas
described as a variety of Aleuria aurantia(asPeziza aurantia
Oed.).Ramsbottom(1914)citedO. aurantiavar.stipitata as a
synonym of Sowerbyella rhenana(asAleuria rhenanaFuckel),
butthetypematerialispresumablylost(Spooner&Yao1995)
andnomoderninterpretationcanbeprovided.
Otidea auriculariiformisHenn.,Hedwigia36:232.1897
Holotype.brazil, A. Glaziouno.20181(S-F9965,exHerb.Sydow)!
 Notes—ThisspeciesbelongstothegenusPhillipsia, Sar-
coscyphaceae. The large spores, (30.5 –)31.5– 36.5(– 37) ×
12.5–14.0µm(Lm=33.2µm,Wm=13.1µm,from13spores),
areellipsoid,inequilateralinprofileview,smoothorwithfaint
cyanophobic, parallel, longitudinal ridges. Asci seem thick-
walled,with an internal eccentricthickened apical pad. The
medullary excipulum is of interwoven hyphae, running mostly
parallel with the outer surface and the ectal excipulum is a
narrow band of textura prismatica, with the long axes of the
cellsparallelto theexterior.Thetropicaldistributionand the
substrate, suggested in the diagnosis to be wood, are typical
for Phillipsia.
Otidea cinerascensVelen.,Novit.Mycol.:152.1947
Holotype.CzECh rEpubliC,Moravia,Žarošice,Aug.1940,V. Vacek(PRM
151779).
 Notes—Thegrey-ochraceousapotheciaand sporeswith
parallel sides suggest O. cinerascens belongs to the O. aluta-
ceacomplex.Typenotstudiedbyus,butannotatedin2009by
B.SpoonerasO. alutacea (aphotographofthecollectionand
annotationprovidedbyJanHolec,PRM).
Otidea cochleata(L.)Fuckel,Jahrb. Nassauischen Vereins
Naturk.23–24:329.1870.‘1869–1870’
Basionym.Peziza cochleataL., Sp.Pl.4: 183.1753:Fr.,Syst.Mycol.,
Index:129.1832.
 ≡Cochlearia cochleata (L.)Lambotte,Fl.Mycol.Belgique1:323.1880.
 Notes—The interpretation of the original description is
difficult,butO. cochleata has been treated as a taxon in the O.
alutacea group(e.g.Lundell&Nannfeldt1938,Dissing2000,
Mornand&Courtecuisse2005,Zhuang2006),probablyfollow-
ingBulliard(1791:plate154asPeziza cochleata).Partofthe
Bulliardplate(f.b)hasnowbeenselectedasthelectotypefor
O. alutacea(Carbone2010a).ThenameO. cochleata should
be considered in future revisions of the O. alutaceacomplex.
For a review of the nomenclatural history of O. cochleata see
Carbone(2010a)andParslow&Spooner(2013).
Otidea darjeelensis(Berk.)Sacc.,Syll.Fung.10:4.1892
Basionym.Peziza darjeelensisBerk.,Hooker’sJ.Bot.KewGard.Misc.
3:202.1851.
 Notes—Rifai(1968)statedthatthetypespecimenofP. dar-
jeelensisatKewhasiodinepositiveasciandechinulatespores,
and does not belong to Otidea.TwocollectionsofP. darjeelensis
arepresentinKewthatmayrepresentoriginalmaterial:india,
Sikkim, J.D. Hooker(K(M)177412,exHerb.Berkeley);and
india,Sikkim (K(M)177413,exHerb. Cooke).Noannotation
labelbyRifaiwasfound(B.Aguirre-Hudson,pers.comm.),but
hemostlikelystudiedthecollectionfromBerkeley’sherbarium,
becausehelistedJ.D.Hookerasthecollector.Thecollection
K(M)177413mightbeapartofK(M)177412,becauseCooke
(1876, f. 215) illustrated P. darjeelensis from specimens in
Berkeley’sherbarium.Bothcollectionsshouldbestudiedand
alectotypeselected.
Otidea dochmia(Berk.&M.A.Curtis)Sacc.,Syll.Fung.8:
95.1889
Basionym.Peziza dochmia Berk.& M.A.Curtis,J. Linn.Soc. Bot.10:
364.1869.
 ≡Phillipsia dochmia(Berk.&M.A.Curtis)Seaver,N.Amer.Cup-fungi,
Operc.:184.1928.

225
I.Olariagaetal.:AmonographofOtidea
 ≡Aurophora dochmia (Berk.&M.A.Curtis)Rifai,Verh.Kon.Ned.Akad.
Wetensch.,Afd.Natuurk.,sect.2,57:52.1968.
 Notes—ThisisthetypespeciesofthegenusAurophora.
Rifai(1968)distinguishedAurophora from Phillipsia by its fan-
shaped apothecia and the presence of a gelatinous matrix in
themedullaryexcipulum.
Otidea domingensis(Berk.)Sacc.,Syll.Fung.8:97.1889
Basionym.Peziza domingensis Berk.,Ann.Mag.Nat.Hist.,ser.II,9:201.
1852.
 Notes—Thisisthe type species of thegenusPhillipsia,
P. domingensis(Berk.)Berk.(seeHansenetal.1999).
Otidea doratophora(Ellis&Everh.)Sacc.,Syll.Fung.8:96.
1888
Basionym.Peziza doratophoraEllis&Everh.,J.Mycol.1:90.1885.
 Notes—Thesmallsporesandasci,alongwiththepointed
paraphyses, suggest this taxon does not belong to Otidea.Cash
(1953)proposedO. doratophora is a synonym of Ionomidotis
irregularis(Schwein.)E.J.Durand(asMidotis irregularis).
Otidea euplecta(Cooke)Sacc.,Syll.Fung.8:97.1889
Basionym.Peziza euplectaCooke,Mycographiapart3:125.1876.
Holotype.USA,Alabama,Peters4560(K(M)161851,exHerb.Berkeley
as Peziza phlebophoravar.)!
 Notes—Thetypeisinapoorcondition:oneapotheciumim-
mature;theotherinfected.But the species likely belongs to
Sarcoscypha, Sarcoscyphaceae.Theasciareinamyloid,thick-
walled and with an eccentrically placed, thickened operculum,
sporesareellipsoid,slightlyinequilateral,19 –21×10.5 –11µm,
smooth,andparaphysesstraight,filiform,branchingabove.The
excipulum is composed of interwoven hyphae that give rise on
theoutsidetoshallowpustules.
Otidea felina (Pers.) Bres., Fungi Trident.ser. 2, fasc. 14:
103.1900
Basionym.Peziza felinaPers.,Mycol.Eur.1:223.1822.
Holotype. FranCE,prope Pariseos, sylvula Meudon (L0116774,Herb.
Persoon).
 Notes—VanVooren&Carbone(2012)revisedtheholotype
and demonstrated that it belongs to the O. alutacea group.
Parslow&Spooner(2013)consideredO. felina a synonym of
O. alutacea.FurtherstudiesontheO. alutacea group should
consider the name O. felina,whichmightbeepitypifiedforan
unequivocalinterpretation.
Otidea fibrillosaMassee,Brit.Fungus-Fl.4:449.1895
 ≡Pseudaleuria fibrillosa (Massee)J.Moravec(‘Pseudoaleuria’),Acta
Mus.Morav.Sci.Biol.88:51.2003.
 Notes—AccordingtoMoravec(2005)thisisaspeciesof
Pseudaleuria, P. fibrillosa.
Otidea grandis (Pers.) Boud., Bull. Soc. Mycol. France 9:
10.1893
Basionym.Peziza grandisPers.,Ann.Bot.Usteri15:27.1795.
 ≡ Peziza abietinavar.grandis(Pers.)Pers.,Mycol.Eur.1:233.1822.
 ≡ Aleuria grandis(Pers.)Gillet,Champ.FranceDiscomyc.:42.1879.
 ≡ Scodellina grandis(Pers.)Seaver,N.Amer.Cup-fung.,Operc.:186.
1928.
 Notes—TheoriginalsenseofO. grandis corresponds most
likely to a species of the Peziza depressa-Ruhlandiella lineage
(Hansenetal.2005)duetothelackofasplit.Nooriginalmate-
rial of O. grandisseemstobekeptinPersoon´sherbarium(L).
ThenameO. grandis has been used for O. bufonia(Boudier
1905;specimenPC0093644isO. bufonia)orO. unicisa(Ka-
nouse1949,Liu&Zhuang2006).Weregardithereasanomen
dubium and confusuminagreementwithHarmaja(2009a).
Otidea grandisvar.scheremetjeffiiHenn.,Hedwigia42,3:
(116).1903
 Notes—Hennings(1903)describedthis taxon based on
specimens kept in formalin and the colours provided in the
protologueare probably imprecise.The spore size can fit
O. bufonia or O. onotica, but there is no type material extant in
BorS,andapreciseinterpretationcannotbeproposedhere.
Otidea harperianaRehm,Ann.Mycol.2:34.1904
Holotype.USA,Ohio,BlueMountains,onground,6June1903,Harper
333(S-F9961,exHerb.Rehm,‘Herb.R.A.andA.M.Harper333’)!
 Notes—Thisspeciesiscloselyrelatedorconspecificwith
Peziza phyllogenaCooke. The asci arestrongly amyloid in
MLZwithageneral bluingovertheapexandthesporesare
ornamented with irregular, low, separate warts that are higher
andmoredenselyplacedat the poles (forming ‘pole caps’).
Oursporemeasurementsfrom theholotype,18 –19.5×8–9
µm(Lm=18.6µm,Wm=8.5µm,from15spores) are larger
thanthosegivenintheprotologue,15–17×5–7µm.
Otidea hirneoloides(Berk.)Sacc.,Syll.Fung.8:96.1889
Basionym.Peziza hirneoloidesBerk.inBerkeley&Curtis,J.Linn.Soc.
Bot.10:365.1869.
 ≡Phillipsia hirneoloides(Berk.)Berk.,J.Linn.Soc.Bot.18:388.1881.
 Notes—Thewood-inhabiting,ear-shapedapothecia,and
especially, the cymbiform spores suggest this name is to be
referred to Phillipsia.Hansenetal.(1999)suggestedP. hirneo-
loides belongs to the Phillipsia domingensiscomplex.
Otidea lechria(Berk.&Broome)Sacc.,Syll.Fung.8:97.1889
Basionym.Peziza lechria Berk.&Broome, J.Linn. Soc.Bot.14: 103.
1875.
Holotype. Sri lanka,on rotten wood, Nov. 1867, G.H.K. Thwaites
(K(M)161847,exHerb.Berkeley).
 Notes—B.Spoonerannotatedtheholotypein2008,and
noted it has amyloid asci and belongs to Peziza.
Otidea lilacinaR.Heim&L.Remy,Bull.Soc.Mycol.France
48:65.1932
 Notes—Theornamented,multi-guttulatesporesandstraight
paraphyses suggest that this taxon does not belong to Otidea.
No original material could be traced in PC (B. Buyck, pers.
comm.).
Otidea lobataRodway,Pap.&Proc.Roy.Soc.Tasmania:116.
1925‘1924’
 Notes—Rifai(1968)statedthatthetypespecimenappears
to represent the inoperculate genus Discinella.

226 Persoonia–Volume35,2015
Otidea luculenta (Cooke)Massee,Brit.Fungus-Fl., 4: 450.
1895(‘leuculenta’)
Basionym.Peziza luculentaCooke,Mycographiapart3:121.1876.
 Notes—Theentire,orangeapotheciaandstraightparaphy-
ses with orange granules suggest this name does not belong
to Otidea.Nannfeldt(1966)notedthatO. luculenta has ‘other
affinities’thanOtidea,butdidnotprovideagenericplacement.
Otidea luteonitens(Berk.&Broome)Massee,Brit.Fungus-
Fl.4:449.1895
Basionym.Peziza luteonitens Berk.&Broome,Ann.Mag.Nat.Hist.,ser.
II,7:180.1851.
 Notes—ThisnameiscurrentlyplacedinAleuria as Aleuria
luteonitens(Berk.&Broome)Gillet.
Otidea micropus(Pers.)Sacc.,Syll.Fung.8:98.1889
Basionym.Peziza micropus Pers., Icon.Desc.Fung. 2:30.1800: Fr.,
Syst.Mycol.2:54.1822.
 Notes—ThisnameisasynonymofPeziza varia(Hedw.:
Fr.)Fr.sensuHansenetal.(2002).
Otidea microspora (Kanouse)Harmaja,Karstenia15:32.1976
Basionym.Otidea alutacea var.microspora Kanouse,Mycologia41:668.
1949.
 Notes—Kanouse(1949)describedthistaxonasavariety
of O. alutacea.Sheindicatedtwodifferentcollectionsasthe
type;A.H. Smith9351afterthediagnosisandA.H. Smith17699
inthe materialexamined.WehavestudiedA.H. Smith9351
(MICH14406,dupl.UPSF-629985!)andithasfarlargerspores
thanstated intheprotologue (13 –15.5×7– 8 µmvs9–10 ×
5.5–6.5µm).Theoblongspores,anectalexcipulumoftextura
angularis,andtheabsenceofyellowpigmentinKOHindicate
it belongs to the O. alutaceacomplex.AsforA.H. Smith17699
(UPSF-629996!),thesporesmatchtheoriginal description.
TheapothecialshapeissimilartothatofO. rainierensis.AGen-
BankITSsequenceofaparatypeofO. microspora(A.H. Smith
30502)differsonly1bpfromtheITSsequenceoftheholotype
of O. rainierensis(Fig.3),butweprefernottoselectalectotype
untilalltheoriginalmaterialhasbeenexamined.Wetherefore
treatthenameasdoubtfulforthetimebeing.
Otidea neglectaMassee,Grevillea22:66.1894
 Notes—ThisnamewaserectedasanewnameforO. auri-
culainthesenseofRehm(1883),Saccardo(1889)andBresa-
dola(1884,asPeziza).Thespeciesthatthesethreeauthors
treated under the epithet auricula is Wynnella silvicola(Beck)
Nannf.initscurrentsense.
Otidea obtecta (Schwein.)Sacc., Syll.Fung.8:98.1889
Basionym.Peziza obtectaSchwein.,Trans.Amer.Philos.Soc.ser.2,4:
170.1832‘1834’.
 Notes—Theoriginaldescriptiongivesstipitate,1–1.5cm
wide, cinnamon-coloured apothecia with a split, growing among
leaves.Seaver(1928)statedthattheidentityofO. obtecta is
uncertain.NooriginalmaterialcouldbelocatedinPHandthe
identificationofthisspeciescannotbeinferred.
Otidea olivaceaBucholtz,Bull.Soc.Imp.NaturalistsMoscou
2:325.1897
 Notes—Thecurved,uniguttulatesporesandtheoccurrence
on a rotten trunk suggest that this taxon does not refer to a
species of Otidea.TheBucholtzherbariumwasboughtbythe
FH,butnoauthenticmaterialofO. olivacea could be located
there.
Otidea onoticavar.ochracea(Fr.)Sacc.,Syll.Fung.8:95.
 1889
Basionym.Peziza onoticavar.ochraceaFr.,Syst.Mycol.2:48.1822:Fr.
loc.cit.(‘ß ochracea’).
 ≡Peziza ochracea(Fr.)P.Karst.,Not.Sallsk.FaunaFl.Fenn.Forh.10:
110.1869.
 ≡Otidea ochracea(Fr.)Seaver,Bull.Lab.Nat.Hist.IowaStateUniv.5:
45.1904.
 Notes—Fries(1822) stated thatthis variety is close to
O. onotica.ItwaselevatedtospeciesrankbyKarsten(1869),who
later considered it a synonym of O. onotica(Karsten1871).As
no original material exists and the original description is meagre,
wearenotabletoprovideagoodinterpretationofthistaxon.
Otidea pleurota(W.Phillips)Sacc.,Syll.Fung.8:97.1889
Basionym.Peziza pleurota W.Phillips in Cooke, Mycographia part 5:
208.1878.
 ≡Iotidea pleurota(W.Phillips)Clem.,Gen.Fungi:175.1909.
Holotype.England,1877,W. Phillips(K(M)29973).
 Notes—Eckblad(1968)observedstronglyamyloidasciin
the type and concluded it belongs to Peziza.B.Spooneran-
notated the type as ‘Peziza badiofusca ?’.
Otidea radiculata(Sowerby)Bres.,FungiTrident.ser.2,fasc.
11–13:72.1898
Basionym.Peziza radiculata Sowerby,Col.Fig.Engl.Fung. 1:46(un-
numberedpage),t.114.1797:Fr.,Syst.Mycol.2:81.1822.
 Notes—ThisnameisplacedinSowerbyella, as S. radicu-
lata(Sowerby) Nannf. Yao&Spooner(2006)examined the
typeatKandconfirmeditsplacement.
Otidea reisneriVelen.,ČeskéHouby4– 5:872.1922
 Notes—Svrček(1976)studiedthetypematerialandcon-
cluded it is a synonym of Sowerbyella radiculata(Sowerby:Fr.)
Nannf.
Otidea schulzeriQuél.inSchulzer,Hedwigia24,4:150.1885
 Notes—TheapotheciaofO. schulzeri were described as
elongated on one side, split, pale yellow-grey and pseudo-
stipitate.However,theverythickflesh(3–4mm),thestraight
paraphysesandthesporesize(20–28µm)suggestthistaxon
does not represent Otidea.AlsoNannfeldt(1966)statedthat
O. schulzeri could hardly belong to Otidea.
Otidea silvicolaBeckinSacc.,Syll.Fung.8:97.1889
 Notes—ThisisWynnella silvicola.Thenamewascreated
for Peziza atrofusca Beck, a later homonym for P. atrofusca
Berk.&M.A.Curtis.
Otidea sparassisQuél.,Rev.Mycol.(Toulouse)54:65.1892
 Notes—TheuniguttulatesporesdonotsuggestanOtidea.
ThistaxonmightrefertoasparassoidHelvellaspecies.
Otidea subonoticaHenn.,Hedwigia36:232.1897
 Notes—Theoriginaldescriptiondoesnotprovideanydis-
cordant feature for Otidea,exceptitwasreportedfromBrazil.

227
I.Olariagaetal.:AmonographofOtidea
No type material of O. subonoticaexistsinHenningsherbarium
inB(Carbone2009)orinS,anditscorrectplacementcannot
beinferredwithcertaintyhere.
Otidea succosa(Berk.)Thüm.,Mycoth.Univ.15:no.1411.
 1879
Basionym.Peziza succosa Berk.,Ann.Mag.Nat. Hist., ser.I, 6: 358.
1841.
 Notes—This is currently considered to be a species of
Pezizas.l.(Hansenetal.2005).
Otidea tasmanicaRodway,Pap.&Proc.Roy.Soc.Tasmania:
116.1925‘1924’
 Notes—Rifai(1968)studiedthetypespecimenofO. tas-
manica in Kew and synonymised it with Peziza praetervisa
Bres.(sensu Dennis, asRifai depicted ornamentedspores
fromthetypespecimen).Thus,O. tasmanica might be close
to, or a synonym of, Peziza subviolaceaSvrček.
Otidea violaceaA.L.Sm.&Ramsb.,Trans.Brit.Mycol.Soc.
 5:237.1916
Holotype.England,Warwickshire,W.B. Grove1915(K(M)30407,exHerb.
W.B.Grove).
 Notes—Parslow&Spooner(2013)examinedtheholotype
and concluded it is a species of Peziza(cf.azureoidesDona-
dini).
AcknowledgementsThismonographbecamemorecomprehensivethanks
to the great efforts of numerous mycologists who provided us with interesting
material.Allour colleagueswhocontributedcollections,discussed issues
orhelpedusinotherwaysarewarmlythanked.EspeciallywethankNancy
S.Weber,RosanneHealy,MatsKarströmandTapioKekkiforputtingatour
disposalvaluablematerial accompanied by colour photographs.Wealso
thank the curators and staff in the herbaria mentioned in the taxonomic
part,H.J.M.Sipman(B),M.HerreraandI.Salcedo(BIO),H.Knudsen(C),
P.G.Jamoni and D.Bolognini(GMFN), D.H.Pfisterand G. Lewis-Gentry
(FH),P.SaloandS.Stenroos(H),E.D.Liu(HKAS),Hong-MeiLu(HMAS),
B.Aguirre-Hudson, B. Dentinger and B.Spooner(K),R. Rabeler and
P.Rogers(MICH),D.McLaughlin(MIN),A.B.MujicandJ.Spatafora(OSC),
B.Buyck (PC),A. Freire-Fierro(PH),J. Holec (PRM),S.Huhtinen (TUR,
TUR-A)andR.Berndt(ZH),forsearchingmaterialandarrangingloans.We
appreciateKerstinGillen´sassistanceduringfieldworkin2010.JuanSantos
andXiang-Hua Wangcollectedpart ofthematerial duringtheirparticipa-
tionintheSwedishPezizomycetesproject.WeareindebtedtoXiang-Hua
WangforsequencingafewimportantOtidea collections, for discussions and
helpineditingthemanuscript,andSeppo Huhtinenfor instructivediscus-
sionsonresinousexudates.WethankD.H.HawksworthandL.A.Parrafor
nomenclaturaladvice.DonaldH.PfisterandTrondSchumacherreviewed
the manuscript and we are grateful for their valuable comments and correc-
tions.FundingforthisresearchwasprovidedbyagrantfromtheSwedish
TaxonomyInitiativetoK.H.(grantno.143/ 071.4).
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INDEX to species, varieties and forms
Accepted names treated in the monograph are in bolditalic.
abietina 166,167,193,195,197,224
abietinavar.grandis 225
abietinaf./var.nigra195
alba185,224
alutacea 175–177,179,181,182,224,225
alutacea var.microspora 226
angusta176,198,200
apophysata 167,168,178,179,180–182
atrofusca 226
aurantia223,224
aurantia var. atromarginata224
aurantia var. stipitata 224
auricula167,185,187,226
auriculariiformis224
azureoides227
badiofusca226
bicolor 178,211
borealis168,171,174,176–178,212,213
boudieri180
brevispora174,179, 205
brunneoparva168,179,187, 191
bufonia175,176,179,185,192,205,206,209,
 211,213,219,224,225
caeruleopruinosa168,174,177,179,213,
215,219
caligata 189,193,195,197
cantharella167,178,189,191,192,193,197,
 216,219
cantharellavar. minor188,189,219,220
cinerascens185,224
cochleata166,167,180,183–185,195,206,
 207,223,224
cochleata var.alutacea182
cochleatavar.umbrina206,175
concinna167,168,174,175,177,178,189,
 210,213,216,219,223
crassa174,185,187
daliensis167,168,178–180,181,192,197
darjeelensis224,225
dochmia 224
domingensis 225
doratophora 225
euplecta 225
felina179,180,185,187,225
fibrillosa 225
flavidobrunneola177,179,213,216,217
formicarum168,171,174,176,177,179,189,
197, 200, 201
fuckelii185,187
fusconigra 192,205
grandis205,207,208,209,225
grandisvar.scheremetjeffii 225
harperiana 225
hirneoloides 225
indivisa195–197
integra168,174,178,223
kauffmanii171,175,213,214,223
kaushalii174,176,177,178,202, 205
kunmingensis178,184,185
lactea174,178,179,213, 219
lechria 225
leporina 166,167,171,174,175,177,179,185,
 188,189,192,209– 211
leporina f. major185
leporinaf./var.minor185,195
leporina var.onotica 209
leporinavar.rubescens185
lilacina 225
lobata 225
lohjaënsis 200
luculenta226
luteonitens226
marsupium var.pyxidata216
micropus226
microspora 226
minor174,177,178,189,213,217,219, 222
mirabilis171,174–176,179,192,205,207,
209,211,213
myosotis185,187
nannfeldtii168,175–177,179,185,189,191,
 197,198, 201, 202
neglecta226
obtecta226
ochracea226
olivacea192,226
olivaceobrunnea 178,192
onotica166,167,174,177–179,205,209,
 225,226
onoticavar. brevispora174,205,206
onoticavar.ochracea 226
oregonensis168,174,177,178,213,217,
 219,220,223
papillata175,179,185,189,191,198,200
papillataf.pallidefurfuracea189
pedunculata206,207,209
phlebophora167,168,174,178,213,217,219,
220, 222, 225
phyllogena 225
platyspora178,179,180,182
pleurota 226
praetervisa227
propinquata167,177,178,191–193,195,224
pseudobadia 206,209
pseudoleporina168,174,176,177,179,185,
188,197,200,201
purpurea179,205,211
pusilla192
pyxidata 216
radiculata226
rainierensis174,175,179,213,214,217,222,
223,226
reisneri 226
rhenana 224
rosea209
schulzeri 226
scutellata 216
shimizuensis 167
silvicola 167,185,187,226
sinensis 174,178,213
smithii168,171,176,179,192,205,209,211
sparassis226
subconcinna219
subformicarum168,171,174,179,197,198,
200
subonotica 226
subpurpurea178,211
subterranea 167,178,179,181
subviolacea 227
succosa227
tasmanica 227
tianshuiensis 178,213
tuomikoskii 171,177,179,185,189, 200, 202
umbrina 184,187,206– 208
unicisa 167,168,174,177,178,202,205,
211, 225
varia226
violacea227
yunnanensis 167,174,178,179,202,204,205