Phylogeny and taxonomic revision of the Planistromellaceae

© 2012 Nationaal Herbarium Nederland & Centraalbureau voor Schimmelcultures

You are free to share - to copy, distribute and transmit the work, under the following conditions:Attribution: Youmustattributetheworkinthemannerspecifiedbytheauthororlicensor(butnotinanywaythatsuggeststhattheyendorseyouoryouruseofthework).Non-commercial: Youmaynotusethisworkforcommercialpurposes.Noderivativeworks: Youmaynotalter,transform,orbuilduponthiswork.Foranyreuseordistribution,youmustmakecleartoothersthelicensetermsofthiswork,whichcanbefoundathttp://creativecommons.org/licenses/by-nc-nd/3.0/legalcode.Anyoftheaboveconditionscanbewaivedifyougetpermissionfromthecopyrightholder.Nothinginthislicenseimpairsorrestrictstheauthor’smoralrights.

Persoonia 29, 2012: 11–28www.ingentaconnect.com/content/nhn/pimj http://dx.doi.org/10.3767/003158512X658766RESEARCH ARTICLE

INTRODUCTION

KellermaniawasestablishedbyEllis&Everhart(1885)toac-commodate an unusual coelomycete, K. yuccigena, with large, cylindrical, and septate conidia that were considered stipitate, and it occurred on dead leaves of Yucca(Asparagaceae, sub-family AgavoideaesensuAPGIII(2009)=Agavaceae in earlier classifications).ThestipesoftheconidiadescribedbyEllis&Everhart(1885)areinterpretedasapicalappendages(Sutton1968).Höhnel(1918a)adoptedthegenericnamePiptarthron that was provisionally suggested by Montagne for Septoria macrospora, a coelomycetous species without appendages that occurred on senescing leaves of Agave(Asparagaceae, subfamily Agavoideae=Agavaceaeinearlierclassifications).He provided a generic diagnosis and comparison to Keller-mania.Subramanian&Ramakrishnan(1954)establishedthecoelomycetous genus Alpakesa, which was characterised in part by conidia with multiple apical appendages, when their study revealed that Neottiospora yuccifolia, a species found on dead leaves of Yucca, was not congeneric with the type of Neottiospora.Several other studies have added new species to Alpakesa, Kellermania, and Piptarthron and/or provided revised generic

circumscriptions.Sutton(1968)restrictedKellermania to spe-cies with simple, blastic conidiophores; septate, hyaline, and appendage-bearing conidia; and sclerotioid, pycnidial conidio-mata, while discussing segregate genera and excluding many namesclassifiedinKellermania.Morgan-Jonesetal.(1972a)re-examined the type species of Alpakesa,modifiedthegenericcircumscription,andaddedtwospecies.Morgan-Jonesetal.(1972b)revisedtheconceptsofthetwoKellermania species thatwereacceptedbySutton(1968),addedtwonewspecies,and included taxa with or without a single apical conidial ap-pendage.Sutton (1977) synonymised theYucca-inhabiting genus Septoplaca with Piptarthron, but he was unable to de-termine which species was represented by the type, S. limbata.Sutton(1980)latertreatedS. limbata as Piptarthron limbatum and subsequently as P. yuccae(Sutton1983).Sutton(1980)maintained Alpakesa, Kellermania, and Piptarthron for genera havingmultiple,single,ornoconidialappendages,respectively.Sutton(1980)alsonotedthepossibilityofabroadlyexpandedKellermania for all species in the complex.NagRaj (1993)treated Alpakesa as a later synonym of Kellermania and listed theunexaminedorexcludedtaxa.Ramaley(1991,1992,1993,1995,1998)providedthenextmajor advances in the study of this group of fungi by discovering and describing the sexual states of Kellermania and Piptarthron, adding new species, and reviewing several coelomycetous species.ThegenusPlanistroma, the sexual state of Piptarthron, was characterised by subepidermal, ostiolate ascomata in multi-locular stromata with bitunicate asci and lackling paraphyses (Ramaley1991).Ramaley (1992)addedanotherspeciesofPlanistroma, P. obtusilunatum, with unilocular conidiomata more typical of Kellermania.Shenotedthe intermediatena-ture of the asexual state in that the lack of apical appendages

Phylogeny and taxonomic revision of the Planistromellaceae including its coelomycetous anamorphs: contributions towards a monograph of the genus KellermaniaA.M.Minnis1a,A.H.Kennedy2a,D.B.Grenier2,M.E.Palm3,A.Y.Rossman4

1 Center forForestMycologyResearch,NorthernResearchStation,USForestService,OneGiffordPinochetDr.,Madison,WI53726,USA;

correspondingauthore-mail:[email protected] NationalIdentificationServices,USDA-APHIS,B010A,10300BaltimoreAve.,Beltsville,MD20705,USA.

3 National IdentificationServices,USDA-APHIS/PPQ/PHP/RIPPS,4700RiverRoad,Riverdale,MD20737,USA.

4 SystematicMycology&Microbiology Laboratory,USDA-ARS,B010A,10300BaltimoreAve.,Beltsville,MD20705,USA.

a Bothauthorscontributedequallytothiswork.

Key words

AgavaceaeAscomycotaAsparagaceaeBotryosphaeriaceaeBotryosphaerialescoelomycetesDothideomycetesmolecular phylogenyPlanistromellaceaeSeptoplacataxonomy

Abstract ThecorespeciesofthefamilyPlanistromellaceae are included in the teleomorphic genera Planistroma and Planistromella and the connected anamorphic, coelomycetous genera Alpakesa, Kellermania, and Piptarthron.Thesegenerahavebeendefinedprimarilyonthebasisofascosporeseptationornumberofconidialappendages.Due to a lack of DNA sequence data, phylogenetic placement of these genera within the Dothideomycetes, evaluation ofmonophyly,andquestionsaboutgenericboundariescouldnotbeadequatelyaddressedinthepast.Isolatesofnearlyalloftheknownspeciesinthesegenerawerestudiedgeneticallyandmorphologically.DNAsequencedataweregeneratedforthenSSU,ITS,nLSU,andRPB1markersandanalysedphylogenetically.Theseresultsplacedthe Planistromellaceae, herein recognised as a distinct family, in an unresolved position relative to other genera within the order Botryosphaeriales.SpeciesrepresentingthecoregeneraofthePlanistromellaceae formed a clade andevaluationofitstopologyrevealedthatpreviousmorphology-baseddefinitionsofgeneraresultedinanartificialclassificationsystem.Alpakesa, Kellermania, Piptarthron, Planistroma, and Planistromella are herein recognised as belonging to the single genus Kellermania.Thefollowingnewcombinationsareproposed:Kellermania crassispora, K. dasylirionis, K. macrospora, K. plurilocularis, and K. unilocularis.Fivenewspeciesaredescribed,namelyK. confusa, K. dasylirionicola, K. micranthae, K. ramaleyae, and K. rostratae.Descriptionsofspeciesinvitroandakeytospeciesknownfromcultureareprovided.

Article info Received:18May2012;Accepted:6August2012;Published:25October2012.

12 Persoonia – Volume 29, 2012

GenBankaccesionno.2

Species Isolatecultureno.1 Host Locality SSU LSU ITS RPB1

Bagnisiella examinans CBS551.66 Lantana camara India GU296139 EU167562 GU301803 n/aBarriopsis fusca CBS174.26 Citrus sp. Cuba EU673182 EU673330 DQ377857 n/aBotryosphaeria corticis ATCC22927 Vaccinium sp. USA EU673176 DQ299247 EU673245 n/aBotryosphaeria dothidea CBS110302 Vitis vinifera Portugal EU673174 AY259092 EU673243 n/a‘Botryosphaeria’ tsugae CBS418.64 Tsuga heterophylla Canada EU673208 DQ458888 DQ377867 n/aDiplodia corticola CBS112546 Quercus ilex Spain EU673207 AY259090 EU673262 n/aDiplodia cupressi CBS168.87 Cupressus sempervirens Israel EU673209 DQ458893 EU673263 n/aDiplodia mutila CBS112553 Vitis vinifera Portugal EU673213 AY259093 AY928049 n/aDiplodia pinea A CBS109727 Pinus radiata SouthAfrica EU673220 DQ458897 EU673269 n/aDiplodia rosulata CBS116470 Prunus africana Ethiopia EU673211 EU430265 DQ377896 n/aDiplodia scrobiculata CBS109944 Pinus greggii Mexico EU673218 DQ458899 EU673268 n/aDiplodia seriata CBS119049 Vitis sp. Italy EU673216 DQ458889 EU673266 n/aDothiorella iberica CBS113188 Quercus suber Spain EU673156 AY573198 EU673230 n/aDothiorella sarmentorum CBS115038 Malus pumila Netherlands EU673159 AY573206 DQ377860 n/aGuignardia bidwellii CBS111645 Parthenocissus quinquefolia USA EU673223 FJ824766 DQ377876 n/aGuignardia citricarpa CBS102374 Citrus aurantium Brasil FJ824759 FJ824767 DQ377877 n/aGuignardia philoprina CBS447.68 Taxus baccata Netherlands FJ824760 FJ824768 DQ377878 n/aHelicomyces roseus CBS283.51 submergedbark,inbrook Switzerland AY856928 AY916464 AY856881 n/aKellermania anomala AR3471,CBS132218 Yucca brevifolia USA JX444899 JX444853 JX444869 JX444884Kellermania confusa AR3469,CBS131723 Yucca thornberi USA n/a JX444854 JX444870 JX444885Kellermania crassispora AR3463,CBS131714 Nolina micrantha USA JX444900 JX444855 JX444871 JX444886Kellermania dasylirionicola AR3465,CBS131720 Dasylirion leiophyllum USA JX444901 JX444856 JX444872 JX444887Kellermania dasylirionis AR3464,CBS131715 Dasylirion leiophyllum USA n/a JX444857 JX444873 JX444888Kellermania macrospora AR3468,CBS131716 Agavesp. USA JX444902 JX444858 JX444874 JX444889Kellermania micranthae AR3474,CBS131724 Nolina micrantha USA JX444903 JX444859 JX444875 JX444890Kellermania nolinae AR3475,CBS131717 Nolina erumpens USA JX444904 JX444860 JX444876 JX444891Kellermania nolinifoliorum AR3473,CBS131718 Nolina microcarpa USA JX444905 JX444861 JX444877 JX444892Kellermania plurilocularis AR3467,CBS131719 Yucca baccata USA n/a JX444862 JX444878 JX444893Kellermania ramaleyae MEP1260,CBS131722 Yucca sp. Mexico n/a JX444863 JX444879 JX444894Kellermania rostratae JB5.16.11-01,CBS131721 Yucca rostrata Mexico n/a JX444864 JX444880 JX444895Kellermania unilocularis AR3466(dead) Yucca baccata USA n/a JX444865 n/a n/aKellermania uniseptata AR3476,CBS131725 Yucca rupicola USA JX444906 JX444866 JX444881 JX444896Kellermania yuccifoliorum AR3472,CBS131726 Yucca brevifolia USA JX444907 JX444867 JX444882 JX444897Kellermania yuccigena AR3470,CBS131727 Yucca filamentosa? USA JX444908 JX444868 JX444883 JX444898Lasiodiplodia crassispora CBS110492 unknown unknown EU673189 EF622086 EU673251 n/aLasiodiplodia gonubiensis CBS115812 Syzygium cordatum SouthAfrica EU673193 DQ458892 DQ377902 n/aLasiodiplodia parva CBS356.59 Theobroma cacao SriLanka EU673200 EF622082 EU673257 n/aLasiodiplodia pseudotheobromae CBS116459 Gmelina arborea CostaRica EU673199 EF622077 EU673256 n/aLasiodiplodia rubropurpurea CBS118740 Eucalyptus grandis Australia EU673191 DQ103553 DQ377903 n/aLasiodiplodia theobromae CAA006 Vitis vinifera USA EU673197 DQ458891 EU673254 n/aLasiodiplodia venezuelensis CBS118739 Acacia mangium Venezuela EU673192 DQ103547 DQ377904 n/aMacrophomina phaseolina AFTOL1783,CBS227.33 Zea mays unknown DQ678037 n/a DQ377906 n/aMelanops sp. CBS118.39 Quercus borealis USA FJ824763 FJ824771 DQ377856 n/aMelanops tulasnei CBS116805 Quercus robur Germany FJ824761 FJ824769 FJ824764 n/aNeodeightonia phoenicum CBS122528 Phoenix dactylifera Spain EU673205 EU673340 EU673261 n/aNeodeightonia subglobosa CBS448.91 keratomycosisinhumaneye UnitedKingdom EU673202 EU673337 DQ377866 n/aNeofusicoccum luteum CBS110299 Vitis vinifera Portugal EU673148 AY259091 AY928043 n/aNeofusicoccum mangiferae CBS118531 Mangifera indica Australia EU673153 AY615185 DQ377920 n/aNeofusicoccum parvum CBS110301 Vitis vinifera Portugal EU673150 AY259098 AY928046 n/aPhaeobotryon mamane CPC12264 Sophora chrysophylla USA(Hawaii) EU673183 EU673331 DQ377898 n/aPhaeobotryosphaeria citrigena ICMP16812 Citrus sinensis NewZealand EU673180 EU673328 EU673246 n/aPhaeobotryosphaeria porosa CBS110496 Vitis vinifera SouthAfrica EU673179 AY343379 DQ377894 n/aPhaeobotryosphaeria visci CBS100163 Viscum album Luxembourg EU673177 EU673324 DQ377870 n/aPseudofusicoccum stromaticum CBS117448 Eucalyptus sp. Venezuela EU673146 AY693974 DQ377931 n/aSaccharata proteae CBS115206 Proteasp. Australia GU296194 n/a GU301869 n/aSpencermartinsia viticola CBS117006 Vitis vinifera Spain EU673166 AY905555 EU673236 n/a1 AFTOL:AssemblingtheFungalTreeofLife;AR:CulturecollectionofAmyRossman,housedatU.S.NationalFungusCollections(BPI),Beltsville,MD,USA;ATCC:AmericanTypeCultureCol-lection,Manassas,VA,USA;CBS:CBS-KNAWFungalBiodiversityCentre,Utrecht,TheNetherlands;CAA:ArturAlves,UniversidadedeAveiro,Aveiro,Portugal;ICMP:InternationalCollectionofMicroorganismsfromPlants,Auckland,NewZealand;JB:JosephBischoff,cultureshousedatBPI;MEP:MaryE.Palm,cultureshousedatBPI.

2 SSU:partial18SrRNAgene;LSU:partial28SrRNAgene;ITS:internaltranscribedspacerregions1&2including5.8SrRNAgene;RPB1:partialRNAPolymeraseIIsubunit1gene.

Table 1Isolatesandsequencesemployedinthisstudy.Taxonnamesinboldindicatenewlygeneratedstrainsanddata.

suggested placement in Piptarthron while the presence of unilocular conidiomata supported placement in Kellermania.Ramaley(1993)circumscribedPlanistromella, the sexual state of Kellermania, and noted that it was similar to Planistroma but distinct in having septate ascospores.Thegenus originallyincluded two new species that were correlated with one new and one existing species of Kellermania(Ramaley1993).Shedid not embrace synonymising the asexual and sexual states intoabroadlydefinedPlanistroma with Kellermania anamorphs. Ramaley(1995)addedfivenewspeciesbasedonbothasexualand sexual states and provided a key to the species of Keller-mania and Piptarthron.Mostnotablyamongthenewspecieswas Planistromella torsifolium, the first sexual state havingAlpakesa-type conidia, which provided further support for the

synonymy of Alpakesa and Kellermania(Ramaley1995).Lastly, Ramaley(1998)namedtwomoreteleomorphicspeciescon-nected with known species of Kellermania and illustrated another undescribed species with both Piptarthron and Plani-stromastates.Planistroma kellermaniae, which has aseptate ascospores, was connected to the anamorphic Kellermania nolinae, a species with Alpakesa-type conidia, providing further confusioninregardtogenericboundariesandRamaley(1998)suggestedthatare-evaluationwasnecessary.Barr(1996)observedthatseveralgeneraintheDothideales werenotclassifiedeasilyintoanyfamily,andsheestablishedthe family Planistromellaceae to accommodate “taxa having ascostromata,interthecialtissues,andschizogenouslyformed,periphysateostioles”.Planistromella and Planistroma were in-

13A.M.Minnisetal.:Planistromellaceae - genus Kellermania

cluded and three species were transferred to the former genus along with circumstantial links to anamorphs for two species (Barr1996).ThegeneraLoratospora, Eruptio, Microcyclus, and MycosphaerellopsiswerealsoclassifiedinPlanistromellaceae.DuetotheavailabilityofanumberofRamaley’sculturesthatrepresent nearly all of the known species of Alpakesa, Keller-mania, and Piptarthron and additional cultures obtained from plantdiseaseinterceptionsatU.S.portsofentry,asystematicstudywasmadetoaddressthefollowingquestions:1)Whatare the phylogenetic relationships between members of the Planistromellaceae and the Dothideomycetes?; 2) are thePlanistromellaceae and its generamonophyletic?; 3) doesmorphology of conidial appendages or ascospore septation correlatewith phylogeny?; and 4) are slightmorphologicaldifferences among otherwise similar isolates, which are often obtained from different hosts, indications of distinct phylogenetic species?Toanswer thesequestions,nuclearprotein-codingDNA and nuclear ribosomal DNA sequence data were gener-atedforseverallociandanalysedphylogenetically.Additionally,detailed studies were made of these species in culture, including micromorphological characters.Herbariumspecimenswereexaminedwheneverpossibleand/ornecessary.

MATERIALS AND METHODS

Morphology and herbarium materialDriedherbariummaterialwasrehydratedandviewedin3%KOH(Largentetal.1977),andmicroscopicobservationsofculturesweremadeofmaterialmountedin3%KOHorbuff-eredShear’smountingfluid(Graham1959).LengthtowidthratiosaregivenasQ.Meanvaluesforlength,widthandQareindicated by Lm,Wm,andQm, respectively,basedonn=30.HerbariumacronymsfollowThiers(2012).SeeFarr&Rossman(2012)foradditionalinformationaboutcollectionshousedattheU.S.NationalFungusCollections(BPI).NewspecimensweredepositedatBPI.TheRamaleycollectionsfromwhichmanyof the cultures utilized in this studywere obtained, ifextant,aremissing.

CulturesIsolates were grown in plastic Petri plates on Difco potato-dextroseagar (PDA),whichwaspreparedaccording to themanufacturers’ instructions.Growth conditionswere 24°Cwitha12hlight/darkregimen.Culturesweremeasuredandphotographed after 2wk.Notes on colour and general ap-pearanceweremadeafter 2–3wk.Terminology for colourincludes general terms as well as standard terminology with thesamplereferencecode inparenthesesfromKornerup&Wanscher(1967).Culturesthathadnotsporulatedafter3wkwere continuously incubated under the same conditions for up to severalmonths andperiodically re-examined.Referencecultures were deposited at the Centraalbureau voor Schimmel-cultures(CBS-KNAWFungalBiodiversityCentre,Utrecht,TheNetherlands;Table1).

DNA extraction, PCR amplification, and sequencingDNAwas extracted from freshmycelium usingQiagen’sDNeasyPlantMiniKit (Germantown,MD).RibosomalDNAfromthenuclearsmallsubunit(SSU),theinternaltranscribedspacerregion(ITS;ITS1,5.8S,ITS2),andthenuclearlargesubunit(LSU)werePCRamplifiedusingtheprimerpairsNS1andNS4(Whiteetal.1990),ITS5andITS4(Whiteetal.1990),LROR (Moncalvo et al. 2000) and LR7or LR5 (Vilgalys&Hester1990),respectively.Additionally,aportionofthelargestsubunitoftheRNApolymeraseII(RPB1)wasamplifiedusingtheprimerpairRPB1-AcandRPB1-Cr(Mathenyetal.2002).EachregionwasamplifiedusingGoTaq(Promega,Madison,

WI)andassociatedstandardreagentsfollowingthemanufac-turer’srecommendationsincluding2.0mMMgCl2and1.5µMofeachprimer.ThermalcyclingconditionsforRPB1andLSUwereaccordingtoMalkusetal.(2006)andReebetal.(2004),respectively.ThermalcyclingconditionsfortheSSUandITSwere:95°Cfor60s;35cyclesat95°Cfor15s,50°C(55°CforITS)for20s,and72°Cfor60s;and72°Cfor180s.Cyclesequencing and fluorescent labelling was conducted with the samecorrespondingPCRprimersthatwereusedforeachlocusandtheBigDyeTerminatorv.3.1CycleSequencingkit(Ap-pliedBiosystems,FosterCity,CA).ThepurifiedproductswerethensequencedonanABI3730automatedDNAsequencer.GeneiousProv.5(Drummondetal.2010)wasusedtoeditelectropherograms and to build consensus sequences that were submittedtoGenBank(http://www.ncbi.nlm.nih.gov;Table1).

Data matrix and phylogenetic analysisFor the purpose of determining the phylogenetic position of Planistromellaceae among the Dothideomycetes, sequences of theSSUandLSU from the type speciesof Kellermania, Piptarthron, Planistroma, and Planistromella were manually incorporated into thealignment ofSchochet al. (2009) us-ing theprogramGeneiousProv.5 (Drummondetal.2010;i.e.,Dothideomycetes alignment). Identical sequenceswereremoved from this alignment.AMaximumLikelihood (ML)analysiswasconductedinRAxMLv.7.3.0(Stamatakis2006)usingthe‘RAxML-HPC2onXSEDE’toolviatheCIPRESSci-enceGateway(Milleretal.2010),selectingtheGTRGAMMAnucleotide substitution model and Opegrapha dolomitica and Schismatomma decolorans asoutgroups(Schochetal.2009).Aseparaterapidbootstrap(bs)analysisof1000iterationswasconductedwithidenticalsettings.Multiple-sequence alignments were conducted for each of the datasets from the PlanistromellaceaeinGeneiousProv.5(Drum- mondetal.2010)usingMUSCLEv.3.6(Edgar2004),adjustedmanually,andthenconcatenated(i.e.,Kellermaniaalignment).Congruenceamongthesefourdatapartitions(SSU,ITS,LSU,RPB1)wasevaluatedbycomparingtheirtopologiesinsearchofwell-supported clades (posterior probability >0.85)withconflicting compositions, and separately with the Incongruence LengthDifferencetest(ILD;Farrisetal.1994).TheindividualtopologieswereconstructedwithBayesian inference (BI) inMrBayesv.3.1.2(Huelsenbeck&Ronquist2001,Ronquist&Huelsenbeck2003)wherebest-fittingmodelsforeachdatasetweredeterminedinMrModeltestv.2.2(Nylander2004)bytheAkaikeInformationCriterion(AIC;Posada&Buckley2004).Allotherparameterswereleftasdefault.Theposteriorprobability(pp)distributionoftreeswasestimatedfromthosecollecteduntil the standard deviation of split frequencies reached less than0.01 (1million generations)minus the burn-in (10%),whichwasdeterminedintheprogramTracerv.1.5(Rambaut&Drummond2007).TheILDtestwasimplementedinPAUPv.4.0b10(Swofford2003)asthePartitionHomogeneityTestwith1000homogeneity replicates.Each replicatesearchedtreespacewith100randomadditionsequences(RAS)sav-ing10treesperRASwhileignoringuninformativecharacters.ThemonophylyofcorePlanistromellaceae was tested by con-ductingaphylogeneticanalysisofamatrixcomprisedofSSU,ITS,andLSUfromawiderangeofBotryosphaeriaceae se-lectedfromtheanalysesofCrousetal.(2006),Phillips&Alves(2009),andSchochetal.(2009);plustypespeciesofgeneraof Kellermania, Piptarthron, Planistroma, and Planistromella as wellasatleastonerepresentativeofeachmajorcladeofcorePlanistromellaceae; and Helicomyces roseus(CBS283.51)asoutgroup(i.e.,Botryosphaeriaceaealignment).ThismatrixwasalignedwithMUSCLEv.3.6(Edgar2004)andanalysedwithBI,asdescribedabove.


Dothideomycetes alignment Botryosphaeriaceae alignment Kellermania alignment

N Length No.V.C. No.I.C. DNAModel N Length No.V.C. No.I.C. DNAModel N Length No.V.C. No.I.C. DNAModel

SSU 283 1027 473(46%) 295(29%) N.A. 44 1133 94(8%) 42(4%) GTR+I+G 10 1025 9(1%) 5(0.5%) HKY+I

ITS N.A. N.A. N.A. N.A. N.A. 42 377 130(34%) 92(24%) SYM+I+G 15 351 45(13%) 27(8%) K80+I+G

LSU 327 1853 689(37%) 542(29%) N.A. 45 584 134(23%) 82(14%) GTR+I+G 15 883 29(3%) 18(2%) GTR+I+G

RPB1 N.A. N.A. N.A. N.A. N.A. N.A. N.A. N.A. N.A. N.A. 15 795 221(28%) 160(20%) GTR+G

Combined 342 2880 1162(40%) 837(29%) GTRGAMMA 45 2091 302(14%) 208(10%) Mixed 15 3054 304(10%) 210(7%) Mixed

Table 2Asummaryofmatrixpartitionstatistics(ingrouponly)foreachalignmentwhichwasanalyzedphylogeneticallyinthepresentstudy.N=thenumberoftaxonomicunitsinthealignment,Length=thenumberofnucleotidecharactersinthealignment,No.V.C.=thenumberofvariablecharacters,No.I.C.=thenumberofphylogeneticallyinformativecharacters,DNAmodel=themodelofnucleotidesubstitionusedinananalysisforthecorrespondingpartition.

0.03

Pseudofusicoccum stromaticum CBS 117448

Diplodia cupressi CBS 168.87

Kellermania nolinae AR 3475

Neofusicoccum mangiferae CBS 118531

Diplodia corticola CBS 112546

Diplodia pinea CBS 109727

Barriopsis fusca CBS 174.26

Spencermartinsia viticola CBS 117006

Lasiodiplodia crassispora CBS 110492

Kellermania plurilocularis AR 3467

Macrophomina phaseolina AFTOL 1783

Guignardia bidwellii CBS 111645

Phaeobotryosphaeria porosa CBS 110496

Dothiorella iberica CBS 113188Dothiorella sarmentorum CBS 115038

Guignardia philoprina CBS 447.68

Botryosphaeria tsugae CBS 418.64

Neofusicoccum parvum CBS 110301

Kellermania macrospora AR 3468

Diplodia scrobiculata CBS 109944

Helicomyces roseus CBS 283.51

Diplodia mutila CBS 112553

Lasiodiplodia pseudotheobromae CBS 116459

Phaeobotryosphaeria citrigena ICMP 16812

Diplodia seriata CBS 119049

Diplodia rosulata CBS 116470

Kellermania yuccigena AR 3470

Botryosphaeria dothidea CBS 110302

Lasiodiplodia rubropurpurea CBS 118740

Guignardia citricarpa CBS 102374

Neodeightonia phoenicum CBS 122528Neodeightonia subglobosa CBS 448.91

Lasiodiplodia venezuelensis CBS 118739

Melanops tulasnei CBS 116805

Neofusicoccum luteum CBS 110299

Lasiodiplodia gonubiensis CBS 115812

Kellermania anomala AR 3471

Phaeobotryosphaeria visci CBS 100163

Melanops sp. CBS 118.39

Lasiodiplodia theobromae CAA 006

Botryosphaeria corticis ATCC 22927

Lasiodiplodia parva CBS 356.59

Phaeobotryon mamane CPC 12264

Kellermania yuccifoliorum AR 3472

Saccharata proteae CBS 115206Bagnisiella examinans CBS 551.66

0.75

1

1

1

1

0.59

0.98

0.68

0.99

1

1

0.96

0.80

0.86

0.90

1

0.93

0.92

10.950.96

0.75

1

1

1

1

Fig. 1Bayesianinference50%majority-rulephylogramoftheBotryosphaeriaceaealignmentbasedonanalysisofthecombinedSSU,ITS,andLSUdata.ThistreerevealsamonophyleticKellermania (corePlanistromellaceae)inanunresolvedpositionamongfourothermajoringrouplineagesofBotryosphaeriales.

ThemajorcladesofthecorePlanistromellaceaewereidentifiedfromtheresultsofamixed-modelBIanalysisoftheKellermania alignment plus Helicomyces roseus(CBS283.51)inMrBayesv. 3.1.2 using themethodsoutlined above.This phylogenywasalsousedtoi)testthemonophylyofgenericconceptsinPlanistromellaceae;ii)determinerelationshipsamongitsspe-ciesincludingnoveltaxa;andiii)evaluatetheevolutionofkeymorphologicalcharactersandhostrelationships.AllalignmentsandresultingtreesweredepositedintoTreeBASE(S13234),andnomenclaturalnoveltiesinMycoBank(Crousetal.2004).

RESULTS

Data matrix and phylogenetic analysesSequencing was successful for all core Planistromellaceae taxawith living cultures except for theSSUofKellermania

confusaAR3469,K. dasylirionisAR3464,K. plurilocularisAR3467,K. ramaleyaeMEP1260,andK. rostratae JB5.16.11-01,whichweretreatedasmissingdatainallalignments.TheDothideomycetesalignmentcontained342ingrouptaxawithatotallengthof2880characters,837(29%)ofwhichwerephylogeneticallyinformative(Table2).TheBotryosphaeriaceae alignmentincluded45ingrouptaxa(Table1)withatotallengthof2091characters,208(10%)ofwhichwerephylogeneticallyinformative(Table2).TheKellermaniaalignmentcontained15ingrouptaxawithatotallengthof3054characters,210(7%)ofwhichwerephylogeneticallyinformative(Table2).

Phylogenetic relationships of the PlanistromellaceaeMaximum Likelihood analysis of Dothideomycetes revealed highsupport(bs=91%)foracladecomprisedstrictlyofBotryo-sphaeriaceaethatwascongruentwithSchochetal.(2009)with


theexceptionofawell-supportedclade(bs=99%)containingthe type species of the genera Kellermania, Piptarthron, Plani-stroma, and Planistromella of Planistromellaceae(phylogramnot shown).Bayesian inference of theBotryosphaeriaceae alignment suggested that Planistromellaceae is one of fivemonophyletic members of Botryosphaeriaceae s.l. (seeDis-cussion):i)Bagnisella examinans;ii)Saccharata proteae;iii)Melanops(pp=1.0);iv)Planistromellaceae(pp=1.0);andv)the remaining members of the family including the core mem-bers of Botryosphaeriaceae(pp=1.0;Fig.1).

Phylogenetic diversity and relationships within KellermaniaInspection of the individual phylogenies resulting fromBIanalysesofSSU,ITS,LSU,andRPB1revealedasinglewell-supported incongruence.This incongruencewas betweentheSSUandRPB1treeswhereKellermania macrosporaAR3468was sisterwith a clade containingK.crassispora AR3463,K. dasylirionicolaAR3465,K. micranthaeAR3474,andK. nolinaeAR3475intheSSUanalysis(pp=0.94);whereasK. macrosporawaspartofapolytomy(pp=1.0)intheRPB1analysis containing two other clades, one comprised of K.uni-septataAR3476,K.yuccifoliorumAR3472,andK.yuccigena AR3470,andanothercontainingK.anomalaAR3471andK.nolinifoliorumAR3473.Resultsof the ILD test (Farrisetal.1994)suggestedthattheSSU,ITS,LSU,andRPB1datafrom Planistromellaceaewerecongruent(P=0.230)andthussuitableforcombinedanalysis.OurBI analysis of theKellermania alignment resulted in a consensusphylogram (Fig. 2) comprisedof fivemajor line-ages: i)Kellermania macrosporaAR3468,K.confusaAR3469,K.ramaleyaeMEP1260,andK. rostrataeJB5.16.11-01(pp=0.89);ii)K. yuccigenaAR3470,K.uniseptataAR3476,and K.yuccifoliorumAR3472(pp=0.95);iii)K.anomalaAR

0.3

K. yuccifoliorum AR 3472, Yucca /s/s

K. macrospora AR 3468, Agave /n/-

K. anomala AR 3471, Yucca /s/s/?

K. micranthae AR 3474, Nolina /n/a/?

K. nolinifoliorum AR 3473, Nolina /s/s


K. dasylirionis AR 3464, Dasylirion /n/-

K. dasylirionicola AR 3465, Dasylirion /n/-

K. yuccigena AR 3470, Yucca/s/s

K. crassispora AR 3463, Nolina /n/a

K. nolinae AR 3475, Nolina /m/a

K. plurilocularis AR 3467, Yucca /n/a

K. ramaleyae MEP1260, Yucca /n/s/?

K. confusa AR 3469, Yucca/n/-

K. rostratae JB-5.16.11-01, Yucca /n/-

K. uniseptata AR 3476, Yucca /m/s

0.74

0.95

0.93

0.70

0.53

0.89

1

0.98

0.54

0.96

0.69

0.60

lineage-i

lineage-ii

lineage-iii

lineage-iv

lineage-v

Fig. 2Bayesianinference50%majority-rulephylogramofKellermania (corePlanistromellaceae)basedonanalysisofthecombinedSSU,ITS,LSU,andRPB1alignment.Hatchmarksindicatethatthebranchlengthwasshortenedby50%forpresentationpurposes.Thistreerevealsfivelineages,i–v.Hostgenus,numbersofconidialappendages,andascosporeseptationaregivennexttotaxonnamesinthatorder.Forconidialappendages,n=none,s=single,m=multiple.Forascosporeseptation,a=aseptate,s=septate,-=unknown.Aquestionmarkindicatesuncertaintyinthelinkofasexualandsexualstates.

3471,K.nolinifoliorumAR3473(pp=1.0);iv)K.micranthaeAR3474,K.dasylirionisAR3464,K.crassisporaAR3463,K.da-sylirionicolaAR3465,andK.pluilocularis AR3467(pp=0.74);andv)K.nolinaeAR3475(Fig.2).Lineagev,whichisbasedupon a single isolate, is herein labelled as distinct because its position among the other lineages is unresolved and varied in differentanalyses.Therelationshipsamongthesefivelineagesarenotwellresolvedthoughsomemajorgroupingsareweaklysupported and a sister relationship of lineages ii– iii receives somesupport(pp=0.70).Membersoflineagesi– iiiwereallisolated from either Agave or Yucca(Asparagaceae, subfamily Agavoideae),withtheexceptionofK.nolinifoliorumAR3473on Nolina(Asparagaceae, subfamily Nolinoideae).Membersof lineages iv–v were isolated from members of Nolina or Da-sylirionis(subfamilyNolinoideae)withtheexceptionofK.plurilocularis AR3467onYucca (Fig.2).Lineages i– iiipossessseptate ascospores and lineages iv–v possess aseptate as-cospores, but see K. unilocularisbelowandinFig.3.Lineagei possesses conidia without conidial appendages, lineage ii pos-sesses conidia with single or multiple appendages, lineage iii possesses conidia with single appendages, lineage iv pos-sesses conidia without appendages, and lineage v possesses conidiawithmultipleappendages.

Colony characteristics and micromorphology of KellermaniaColony colour typically ranged from greenish to greyish tones, but K. crassisporauniquely remainedpink.Differenceswereobserved in growth rates, but in general slight differences in growth rates and overall colony appearances were observed fromsubculturetosubculture.Thus,tendenciestowardslowervsfastergrowthratesaremoreusefulthanexactmeasurements.Conidiomata were produced in culture by all species except K. crassispora.Theyarecharacterisedas:superficialorimmersed,


0.06

K. nolinifoliorum AR 3473

K. micranthae AR 3474


K. dasylirionicola AR 3465

K. yuccifoliorum AR 3472

K. confusa AR 3469

K. plurilocularis AR 3467

K. macrospora AR 3468

K. nolinae AR 3475

K. yuccigena AR 3470

K. uniseptata AR 3476

K. dasylirionis AR 3464

K. ramaleyae MEP 1260

K. crassispora AR 3463

K. unilocularis AR 3466

K. anomala AR 3471

K. rostratae JB-5.16.11-01

0.99

0.58

1

0.95

0.89

0.82

0.74

0.72

0.99

0.99

Fig. 3Bayesian inference50%majority-rule phylogramofKellermania basedonanalysisoftheITS,includingK. unilocularis.Hatchmarksindicatethatthebranchwasshortenedby50%forpresentationpurposes.

scattered to densely gregarious, at times confluent becoming fused, discrete or associated with stromatal growths including irregularly column-like and rarely mat-like structures, frequently more or less round in shape, with or without necks, black, at times coveredwithwhitish,greenish,orgreengreyhyphae.Conidiog-enous cells lining the inner conidiomatal walls were holoblastic, determinate, discrete or integrated on short single-celled conidi-ophores,doliiformtocylindrical,hyaline,andsmooth.Conidia werevariableinsizeandshape,aseptate,singletomultiseptate,andbear0–multipleapicalappendages.Alessusefulcharacterincludedthepresenceofafrillatthebaseofconidia.Conidia,which developed within weeks or sometimes only after months, wereexudedfromconidiomatainwhitish,mucilaginousmasses.Conidia often germinate quickly and, in some several month old cultures, nearly all conidia in the mucilaginous masses had eithergerminatedordesiccated.Inthedescriptionsofcultures,colony appearance, growth rate, and conidia are the most valu-ablecharactersfordistinguishingspecies.

TAxONOMy

PlanistromellaceaeM.E.Barr,Mycotaxon60:433.1996. Type genus.PlanistromellaA.W.Ramaley.

Kellermania Ellis&Everh.,J.Mycol.1:153.1885.

Type species.Kellermania yuccigenaEllis&Everh. =Piptarthron Mont.exHöhn.,Hedwigia:60:203.1918. Type species.Piptarthron macrosporum(Durieu&Mont.)Höhn. =Alpakesa Subram.&K.Ramakr.,J.IndianBot.Soc.33:204.1954. Type species.Alpakesa yuccifolia(J.G.Hall)Subram.&K.Ramakr.as‘yuccaefolia’. ?=SeptoplacaPetr.,Sydowia17:271.1964‘1963’. Type species.Septoplaca limbataPetr. =Planistroma A.W.Ramaley,Mycotaxon42:69.1991. Type species.Planistroma yuccigenumA.W.Ramaleyas‘yuccigena’. =Planistromella A.W.Ramaley,Mycotaxon47:260.1993. Type species.Planistromella yuccifoliorumA.W.Ramaley.

Kellermania anomala(Cooke)Höhn.,Sitzungsber.Kaiserl. Akad.Wiss.,Math.-Naturwiss.Cl.,Abt.1,124:84.1915.—

Fig.4,5

Basionym.Discella anomala Cooke,Grevillea7:11.1878.

≡Discula anomala (Cooke)Sacc.,Syll.Fung.3:677.1884. ?=Wettsteinina yuccigena M.E.Barr,Contr.Univ.MichiganHerb.9:547.1972as‘yuccaegena’. ≡Planistromella yuccigena (M.E.Barr)M.E.Barr,Mycotaxon60:434.1996.

Culturecharacteristics—ColoniesonPDA43–44mmaf-ter2wkat24°Cwitha12hlight/darkregimen;surfaceneardullgreen(30E4),coveredwithlow,cobwebbytovelutinous,greenish hyphae; margin uneven, whitish; reverse black with margin uncoloured.Conidia 40–69× 5–8µm,Q=5–14.3(Lm=52.3µm,Wm=7.3µm,Qm=7.4),cylindricaltofusiform,apices tapering to a relatively acute point, tapering towards and typically truncate at bases that may bear an indistinct frill,1–2(–3)-septate;wallssmooth,thinandhyaline,andnotsignificantlyconstrictedatsepta;contentshyaline;appendagesabsentorpresent,attimesscarcelyvisible,upto11µmlong,apical,single,appearingasashort,filiform,hyalinemucro. Habitat&Distribution—DeadleavesofYuccaspp.(Morgan-Jonesetal.1972b,NagRaj1993,Farr&Rossman2012).Thetype was found on Y. draconis(Cooke1878),probablycorrectlynamed Y. brevifolia.ThisfungalspeciesiswidelydistributedinthewesternhalfoftheUSA(Morgan-Jonesetal.1972b,Farr&Rossman2012).

Specimen examined.USA,Arizona,MohaveCo.,0.3milesfromGemAcresRd.,exitmile20fromU.S.Hwy.40,ondeadleavesofYucca brevifolia, 3June1992,coll.A.W. Ramaley,AR3471(CBS132218)isolatedbyA.W.RamaleyfromAWR9228(9229ontube),driedcultureonPDA(BPI882814).

Notes—Ramaley(1993)notedthattheappendagemaybehard to observe or absent and that the middle cell in conidia with twoseptaisregularlyshorter.Weobservedthattheappendage(whenpresentinculture)wasshorterthaninpreviousreportsfrommaterialinnature(Morgan-Jonesetal.1972b,NagRaj1993).Shoemaker&Babcock(1987)redescribedP. yuccigena on Yucca glauca.Barr (1996)provisionally linkedPlanistro-mella yuccigena to K. anomala based on the circumstantial occurrenceofbothonthesameleaves.Thislinkisindirectlysupported by the morphological similarity of the sexual state to that of K. nolinifoliorum,asisterspecies.

Kellermania confusaMinnis&A.H.Kenn.,sp. nov.—Myco-BankMB801095;Fig.4,5

Etymology.Thenamereferstoconfusioninregardtotheidentityofthisfungus.

Culturecharacteristics—ColoniesonPDA35–40mmafter 2wk at 24°Cwith a 12h light/dark regimen; surface nearpalegrey(30B1),coveredwithlow,whitecobwebbytoveluti-nous hyphae; margin uneven, whitish; reverse grey with black conidiomatavisible.Conidia38.5–64×6.5–9.5µm,Q=4.8–8(Lm=49.9µm,Wm=8.3 µm,Qm=6.0), obclavate, at timescurved, apices tapering towards and slightly acute, typically truncateatbases,(1–)2(–3)-septate;wallssmooth,thin,hya-line, and not constricted at septa; contents hyaline; append-agesabsent. Habitat&Distribution—DeadleavesofYucca thornberi.ItisknownonlyfromthetypelocalityintheUSA:AZ.

Specimen examined.USA,Arizona,SantaCruzCo.,Interstate10,mile283.6,northsideofroad,ondeadleavesofYucca thornberi,13Apr.1992,coll.A.W. Ramaley,AR3469(CBS131723)isolatedbyA.W.RamaleyfromAWR9212,driedcultureonPDA(holotype,BPI882824).

Notes—Thisspeciesisdistinguishedfromtheothersthatoccur on Yuccabyits(1–)2(–3)-septateconidiathatlackap-


Fig. 4 Cultures of Kellermaniaspecies.a–e.Cultures(surfaceandreverse)onPDAat2wkafterincubationat24°Cwitha12hlight/darkregime;a.K. anomala(AR3471);b.K. confusa(AR3469);c.K. crassispora(AR3463);d.K. dasylirionicola(AR3465);e.K. dasylirionis(AR3464).

c

b

d

a

e

c

b

d

a

e


pendages.Ramaley(inlitt.)notedthattwoisolatesidentifiedas Piptarthron macrosporum, this one from Yucca and another from Agave,hadconsistentdifferencesinconidialmorphology.Morphological and DNA sequence data support the separation of this species from Kellermania macrospora on Agave.

Kellermania crassispora(A.W.Ramaley)Minnis&A.H.Kenn.,comb. nov.—MycoBankMB801096;Fig.4

Basionym.Piptarthron crassisporum A.W.Ramaley,Mycotaxon55:261.1995. =Planistroma nolinaeA.W.Ramaley,Mycotaxon55:258.1995.

Culturecharacteristics— ColoniesonPDA30–36mmafter 2wkat24°Cwitha12hlight/darkregimen;surfacenearpink-ishwhite(10A2)tocottoncandy-pink,coveredwithlow,veluti-nous hyphae; margin uneven, whitish to pale pink; reverse near palered(7A3).Noconidiomataobserved.Noconidiaobserved.

Habitat&Distribution—DeadleavesofNolina micrantha, Nolina sp. (Ramaley1995, thisstudy). It isknown from theUSA:NM,TX(Ramaley1995,thisstudy).

Specimen examined.USA,Texas,CulbersonCo.,GuadalupeMtns.NationalPark,5.6milesfromhighwayalongWilliamsRanchRd.,ondeadleaves of Nolina micrantha,23Oct.1995,coll.A.W. Ramaley,AR3463(CBS131714)isolatedbyA.W.RamaleyfromAWR9536,driedcultureonPDA(BPI882815).

Notes—TheculturethatwestudieddidnotsporulatethoughRamaley(1995)observedconidiainculture.Ramaley(1995)statedthatconidiaonthehostwere56.8–78.4×12.8–14.4µm,cylindrical with rounded apices, aseptate, and without append-ages.Thelinkbetweentheasexualandsexualstatesisbasedon cultural similarities thatwerenotedpreviously (Ramaley1995).

Fig. 5a.ConidiaofKellermania anomalafromculture(AR3471);b.conidiaofK. confusafromculture(AR3469);c.conidiaofK. dasylirionicola from culture (AR3465);d.conidiaofK. dasylirionisfromculture(AR3464);e.conidiaofK. macrosporafromculture(AR3468);f.conidiaofK. micranthae from culture (AR3474);g.conidiaofK. nolinaefromculture(AR3475);h.conidiaofK. nolinifoliorumfromculture(AR3473);i.conidiaofK. plurilocularisfromculture(AR3467).—Scalebars=30µmforall.

cb

d

a

g h i

e f


Kellermania dasylirionicolaMinnis&A.H.Kenn.,sp. nov. — MycoBankMB801097; Fig.4,5

Etymology.ThisspeciesisnamedforitsoccurrenceonDasylirion leio-phyllum.

Culturecharacteristics—ColoniesonPDA22–25mmafter2wkat24°Cwitha12hlight/darkregimen;surfacewhitishto near pinkishwhite (10A2), coveredwith low, tomentoseto velutinous hyphae; margin uneven, whitish; reverse near orangewhite(5A2).Conidia32–45×3–5µm,Q=6.7–12.5(Lm=39.4µm,Wm=4.5µm,Qm=9.0),moreorlesscylindri-cal with irregularly curved shape, flexuous, apices tapering to a somewhat acute point, typically truncate at bases, aseptate; walls smooth, thin, and hyaline; contents hyaline; appendages absent. Habitat&Distribution—DeadleavesofDasylirion leiophyl-lum.ItisknownonlyfromthetypelocalityintheUSA:TX.

Specimen examined.USA,Texas,PecosCo.,1.2milesnorthofT2400onU.S.Hwy.285,roadsideplant,ondeadleavesofDasylirion leiophyllum, 7May1994,coll.A.W. Ramaley,AR3465(CBS131720)isolatedbyA.W.RamaleyfromAWR9413,driedcultureonPDA(holotype,BPI882821).

Notes—Althoughsimilar toKellermania dasylirionis, this second species on Dasylirion is distinguished by its aseptate conidia with irregular, curved shapes and slower growth on PDA.Ramaley(inlitt.)notedaseptate,curlyconidiainmaterialfrom the host in nature and speculated that the collection may representanundescribedspecies.DNAsequencedataandassociatedanalysesconfirmthisasanewspecies.

Kellermania dasylirionis(A.W.Ramaley)Minnis&A.H.Kenn.,comb. nov.—MycoBankMB801098;Fig.4,5

Basionym.Piptarthron dasylirionisA.W.Ramaley,Mycotaxon55:263.1995.

Culturecharacteristics—ColoniesonPDA45–52mmaf-ter2wkat24°Cwitha12hlight/darkregimen;surfaceneargreenish grey (30C2), coveredwith low, velutinoushyphaethat often covers conidiomata; margin uneven, whitish; reverse near brownish grey (7E2) to greyish orange (5B3).Conidia 45–64×5–9.5µm,Q=6.3–12.3(Lm=57.1µm,Wm=7.2µm,Qm=8.1),cylindricaltonarrowlyfusiformwithsomedegreeofcurvature, apices tapering to a relatively acute point, tapering towards and typically truncate at bases, 0–1-septate, septa approx.medianwhenpresent;wallssmooth,thin,hyaline,andnot constricted at septa; contents hyaline, at times granular; appendagesabsent. Habitat&Distribution—DeadleavesofDasylirion leiophyl-lum, D. wheeleri, Dasylirionsp.(Ramaley1995,thisstudy).ThisspeciesisknownfromMexicoandtheUSA:AZ,TX(Ramaley1995,thisstudy).

Specimen examined.USA,Texas,BrewsterCo., BigBendNationalPark, Sotol Vista, on dead leaves of Dasylirion leiophyllum,25Oct.1994,coll.A.W. Ramaley,AR3464(CBS131715)isolatedbyA.W.RamaleyfromAWR9441,driedcultureonPDA(BPI882816).

Notes—Ramaley(1995)notedthatconidiainculturewereshorterandnarrowerthanthoseonthehostinnature.

Kellermania macrospora(Durieu&Mont.)Minnis&A.H.Kenn., comb. nov.—MycoBankMB801099;Fig.5,6

Basionym.Septoria macrosporaDurieu&Mont.,Expl.Sci.Algerie1:589.1849. ≡Hendersonia montagneiCooke,NuovoGiorn.Bot.Ital.10:19.1878.Note:Thisnom.nov.wasestablishedsincetheepithet ‘macrospora’wasoccupied by H. macrospora Berk.&Broome1850. ≡Hendersonia piptarthraSacc.,Michelia2:111.1880.Note:Thisnom.nov.wasestablishedsincetheepithet‘macrospora’isoccupiedbyH. mac-rospora Berk.&Broome1850.Itisanom.illeg.viasuperfluous,ICBNArt.52

(McNeilletal.2006),sinceH. montagnei was already published as a re-placementname. ≡Stagonospora macrospora(Durieu&Mont.)Sacc.,Syll.Fung.3:450.1883. ≡Piptarthron macrosporum(Durieu&Mont.)Höhn.,Hedwigia60:203.1918.

Culture characteristics—Colonies onPDA52mmafter2wkat24°Cwitha12hlight/darkregimen;surfacenearlightgrey(30C1),withscatteredlow,hyphaebetweenconidiomata;margin uneven, whitish; reverse uncoloured, black, to near grey-ishbrown(6D3).Conidia54.5–93×6.5–11µm,Q=4.9–13(Lm=72.2µm,Wm=9.0µm,Qm=8.3),cylindricaltoobclavate,at times slightly curved, apices tapering towards and obtuse to slightly acute, typically truncate at bases that frequently bear a marginalfrill,3–5(–7)-septate;wallssmooth,thin,hyaline,andnotconstrictedatsepta;contentshyaline;appendagesabsent. Habitat&Distribution—DeadleavesofAgave americana, Agave sp. (Sutton1980,Farr&Rossman2012).Basedonscatteredreports,thisspeciesisknownfromAfrica(Algeria),Europe,andNorthAmerica(USA)(Höhnel1918a,Sutton1980,Farr&Rossman2012).Thisspecieshasalsobeenreportedfrom species of Yucca fromvarious locations (Sutton1980,Farr&Rossman2012),butwehavefoundnospecimensonYucca.

Specimen examined.USA,Arizona,CochiseCo.,northsideofroad,I10mile322.5,ondeadleavesofAgave sp.,13Apr.1992,coll.A.W. Ramaley, AR3468(CBS131716)isolatedbyA.W.RamaleyfromAWR9205,driedcultureonPDA(BPI882817).

Notes—ThisspecieswasdescribedoriginallyfromAgave.Collections on Yucca, including the type of Kellermania multi-septata,havebeentreatedasconspecific(e.g.Sutton1980),but it appears that K. macrospora is limited to Agave.Plani-stromella parryi, described originally from Agave shawii(Cooke1885),Plowrightia agaves, described from Agavesp.(Maublanc1903),andPlowrightia williamsoniana, described from Agave americana (Kellerman1906), are potential synonyms (Barr1996)aswellaspossiblythesexualstateofK. macrospora, buttheseteleomorphsarepoorlyknown.

Kellermania micranthae Minnis&A.H.Kenn.,sp. nov.— MycoBankMB801100;Fig.5,6

Etymology.ThenameisderivedfromNolina micrantha,thehost.

Culturecharacteristics—ColoniesonPDA38–48mmafter2wkat24°Cwitha12hlight/darkregimen;surfaceneargrey-ishgreentogreenishgrey(30B6–30B2)togreyishgreentodullgreen(30E5–30E4),attimesportionscoveredwithlow,white,cottony hyphae; margin more or less even, whitish; reverse neargreyishgreen togreenishgrey (30C3–30C2).Conidia 43–61× 6.5–9.5µm,Q=4.5–7.8 (Lm=49.2µm,Wm=8.6µm,Qm=5.8),cylindricaltoobclavate,attimescurved,apicestapering towards and obtuse, rounded or truncate at bases, aseptate; walls smooth, thin, hyaline; contents hyaline, at times granular;appendagesabsent. Habitat&Distribution—DeadleavesofNolina micrantha.ThisspeciesisknownonlyfromthetypelocalityintheUSA:TX.

Specimen examined.USA,Texas,CulbersonCo.,GuadalupeMtns.NationalPark,5.6milesfromhighwayalongWilliamsRanchRd.,ondeadleaves of Nolina micrantha,23Oct.1995,coll.A.W. Ramaley,AR3474(CBS131724)isolatedbyA.W.RamaleyfromAWR9536,driedcultureonPDA(holotype,BPI882825).

Notes—Thisspeciesisdistinguishedfromtheothersknownfrom Nolina by the combination of its aseptate conidia that lack appendagesanditsgreenishcolourationinculture.Ramaley(inlitt.)statedthatthiswasanundescribedspeciesandgaveittheprovisional name, Piptarthron sotoli.Anundescribedspecies


Fig. 6 Cultures of Kellermaniaspecies.a–e.Cultures(surfaceandreverse)onPDAat2wkafterincubationat24°Cwitha12hlight/darkregime;a.K. macrospora(AR3468);b.K. micranthae(AR3474);c.K. nolinae(AR3475);d.K. nolinifoliroum(AR3473);e.K. plurilocularis(AR3467).

c

b

d

a

e

c

b

d

a

e


(Ramaley1998)withaseptate,Planistroma-type ascospores, which was found on the type specimen of P. kellermaniae on Nolina erumpens inTexas,hasananamorphthatissimilartoK. micranthaeandthetwospeciesmaybeconspecific.

Kellermania nolinae(Pollack)NagRaj,inNagRaj,Coelomy-cetousanamorphswithappendage-bearingconidia:442.1993.—Fig.5,6

Basionym.Bartalinia nolinae Pollack,Mycologia39:620.1947.

≡Alpakesa nolinae (Pollack)Morgan-Jones,NagRaj&W.B.Kendr.,Canad.J.Bot.50:879.1972. =Planistroma kellermaniaeA.W.Ramaley,Mycotaxon66:510.1998.

Culturecharacteristics—ColoniesonPDA9–20mmafter2wk at 24°Cwith a 12hlight/dark regimen; surface neargreenishgrey(25C2),formingtiersofraisedmounds,smoothtograiny, at times portions covered with scattered white, velutinous, aerial hyphae; margin uneven, whitish; reverse near greyish white(25B1).Conidia35–48×8–11µm,Q=3.7–6(Lm=40.5µm,Wm=8.8µm,Qm=4.7), fusiform toobclavate, at timesslightly curved, tapering or not towards generally obtuse apices, tapering towards and typically truncate at bases that frequently bearafrill,2–3-septate;wallssmooth,thinandhyaline,andnotconstricted at septa; contents hyaline; appendages present, 8–24µmlong,apical,3–5,filiform,unbranched,hyaline. Habitat&Distribution—DeadleavesofNolina erumpens, N. microcarpa(Pollack1947,Ramaley1998,thisstudy).ItisknownfromtheUSA:AZ,TX(Pollack1947,Ramaley1998,thisstudy).

Specimen examined.USA,Texas,BrewsterCo.,BigBendNationalPark,3.75milesfromU.S.Hwy.385onroadtoTheBasin,ondeadleavesofNolina erumpens,9May1994,coll.A.W. Ramaley,AR3475(CBS131717)isolatedbyA.W.RamaleyfromAWR9408,driedcultureonPDA(BPI882818).

Notes—Thelinkbetweentheasexualandsexualstatesis based on the production of characteristic conidia in cultures derivedfromasci(Ramaley1998).

Kellermania nolinifoliorumA.W.Ramaley,Mycotaxon55: 255.1995.—Fig.5,6

=Planistromella nolinifoliorumA.W.Ramaley,Mycotaxon66:509.1998.

Culturecharacteristics—ColoniesonPDA23–26mmafter2wkat24°Cwitha12hlight/darkregimen;surfaceneardarkgrey(28F1),at timesportionscoveredwithscatteredwhite,cobwebby, aerial hyphae; margin uneven, whitish, somewhat slimyinappearance;reversenearlightgreytodarkgrey(28D1–28E1)with offwhitemargin.Conidia 37–57.5× 5–9.5µm,Q=5.6–9.3(Lm=49.1µm,Wm=7.1µm,Qm=7.0),fusiform,straight, slightly curved to somewhat sigmoid, apices tapering to a relatively acute point, tapering towards and typically truncate atbasesthatmaybearmucilaginousmaterial,approx.medianly1-septate;wallssmooth,thinandhyaline,andnotsignificantlyconstricted at septa; contents hyaline; appendages present or scarcelyvisible,perhapsabsent,upto5µmlong,apical,single,appearingasablunt,mucilaginousmucro. Habitat&Distribution—DeadleavesofNolina microcarpa, N. micrantha, Nolinasp.(Ramaley1995,1998,thisstudy).ThisfungusisknownfromtheUSA:AZ,NM,TX(Ramaley1995,1998,thisstudy).

Specimen examined.USA,Arizona,YarapaiCo.,0.1milenorthofBigCreek,westsideofHwy.17atmile262.1,ondeadleavesofNolina micro-carpa,19July1996,coll.A.W. Ramaley,AR3473(CBS131718)isolatedbyA.W.RamaleyfromAWR9610,driedcultureonPDA(BPI882819).

Notes—Theconnectionbetweentheasexualandsexualstate is based on the production of characteristic conidia in culturesobtainedfromasci(Ramaley1998).

Kellermania plurilocularis(A.W.Ramaley)Minnis&A.H. Kenn.,comb. nov.—MycoBankMB801101;Fig.5,6

Basionym.Piptarthron pluriloculare A.W.Ramaley,Mycotaxon42:63.1991.

=Planistroma yuccigenumA.W.Ramaley,Mycotaxon42:69.1991as‘yuccigena’.

Culturecharacteristics—ColoniesonPDA20–29mmaf-ter2wkat24°Cwitha12hlight/darkregimen;surfaceneargreenishgrey (30D2),at timesportionscoveredwithwhite,cottony hyphae; margin uneven, whitish; reverse near light grey (18D1).Conidia30.5–56×5–8µm,Q=4.2–11.7(Lm=41.5µm,Wm=6.0µm,Qm=7.2),slightlycurved,falcate,tostronglycurvednearingU-shapes,apicestaperingtoarelativelyacutepoint, typically truncate at bases, aseptate; walls smooth, thin andhyaline;contentshyaline;appendagesabsent. Habitat &Distribution—Dead leaves ofYucca baccata (Ramaley1991, thisstudy).This fungus isknownonly fromtheUSA:CO(Ramaley1991,thisstudy).

Specimen examined.USA,Colorado,LaPlataCo.,Durango,alongbiketrailbetweenChapmanHillskislopeandLion’sDen,ondeadleavesofYucca baccata,8June2000,coll.A.W. Ramaley,AR3467(CBS131719)isolatedbyA.W.RamaleyfromAWR2001b,driedcultureonPDA(BPI882820).

Notes—Thesizerangeof theconidiaof theholotypeofPiptarthron plurilocularewasreportedas(48–)59–76(–98)× (4–)5.5–7(–8)µm(Ramaley1991).Theconidiallengthofthepresentisolate(AR3467)inculturewasmuchshorter,buttheoverall appearance of the conidia was basically the same as thoseoftheholotype.Thelinkbetweentheasexualandsexualstatesisbasedonculturalsimilarities(Ramaley1991).

Kellermania ramaleyaeMinnis,M.E.Palm&Rossman, sp. nov.—MycoBankMB801102;Fig.7,8

Etymology.ThisspeciesisnamedinhonourofAnnetteW.Ramaleyforher outstanding contributions towards the study of Planistromellaceae and coelomycetousanamorphs.

Onhost(holotype):Foliicolous,stromataapprox.0.5–0.75mmdiam, scattered to gregarious, round in top view, subepidermal, immersed becoming erumpent after peeling back of disc-shapedepidermal tissue, black,multilocular.Conidiophores absent.Conidiogenous cells lining the basal and lateral, locu-larwalls,moreorlessdoliiformtoslightlycylindrical.Conidia 48–70.5×11–16µm,Q=4.3–5.3(Lm=58.8µm,Wm=12.6µm,Qm=4.7),cylindricaltoslightlyclavateorrarelycurved,apicesobtuse, typically truncateatbases,2–4(–5)-septate;wallssmooth,thin,hyaline,andnotsignificantlyconstrictedatsepta;contentshyaline,attimesgranular;appendagesabsent. Culturecharacteristics—ColoniesonPDA7–21mmafter2wkat24°Cwitha12hlight/darkregimen;surfaceneardullgreen(26D3,29E4,30E4,)todarkgreen(29F4),coveredwithlow, cottony to velutinous hyphae; margin uneven, whitish; re-verse multi-coloured: uncoloured, black, near greyish turquoise (24D5),anddullgreen(27E3).Conidia35–51×9.5–16µm,Q=2.6–4.6(Lm=42.7µm,Wm=12.3µm,Qm=3.5),cylindri-cal to slightly clavate or rarely curved, apices obtuse, typically truncateatbases,2–4-septate;wallssmooth,thin,hyaline,andnot constricted at septa; contents hyaline, at times granular; appendagesabsent. Habitat&Distribution—DeadleavesofYucca sp.Itisknownonly from the material from Mexico intercepted at the Laredo PlantInspectionStation.

Specimen examined. Mexico,InterceptedatLaredo,Texas,USAondeadleaves of Yucca sp.,2Dec.1985,coll.S. Vesper,MEP1260(CBS131722)isolatedbyM.E.PalmfromBPI525045(holotype,as ‘P. macrosporum’),driedcultureonPDA(BPI882823).


Fig. 7 Cultures of Kellermaniaspecies.a–e.Cultures(surfaceandreverse)onPDAat2wkafterincubationat24°Cwitha12hlight/darkregime;a.K. ramaleyae(MEP1260);b.K. rostratae(JB5.16.11-01);c.K. uniseptata(AR3476);d.K. yuccifoliorum(AR3472);e.K. yuccigena(AR3470).

c

b

d

a

e

c

b

d

a

e


Notes—Thisspeciesisdistinguishedfromtheothersthatoccur on Yuccabyitsconidiainculturewith2–4septa,Q=2.6–4.6,andappendageslacking.Somedifferenceswereobservedindimensionsofconidiaproducedonthehostvsinculture.Aloculoascomycete producing tardily septate ascospores that wasfoundonthehostmaybethesexualstateofthisspecies.

Kellermania rostrataeMinnis,A.H.Kenn.&J.F.Bisch., sp. nov.—MycoBankMB801103;Fig.7,8

Etymology.ThenameisderivedfromYucca rostrata,thehost.

Onhost(holotype):Foliicolous,stromataapprox.0.5–1mmdiam, scattered to gregarious, round to ellipsoid in top view, subepidermal, immersed becoming erumpent after peeling back of disc-shaped epidermal tissue, black,multilocular.Conidiophores absent.Conidiogenous cells lining the basal and lateral, locular walls, more or less doliiform to slightly cy-

lindrical.Conidia30.5–48×5–6.5µm,Q=6.3–9.3(Lm=41.3µm,Wm=5.3µm,Qm=7.8),cylindricaltoobclavate,attimescurved, apices tapering towards and obtuse to slightly acute, typically truncate at bases, aseptate; walls smooth, thin, and hy-aline;contentshyaline,attimesgranular;appendagesabsent. Culturecharacteristics—ColoniesonPDA52mmafter2wkat24°Cwitha12hlight/darkregimen;surfacewhitishtoneargreenishgrey(30C2),attimesportionscoveredwithscatteredlow, wispy, white hyphae; margin uneven, whitish; reverse uncol-ouredwithconidiomatavisible.Conidia33.5–62.5×5–9.5µm,Q=4.2–9(Lm=44.1µm,Wm=7.1µm,Qm=6.3),cylindricaltoobclavate, at times curved, apices tapering towards and obtuse toslightlyacute,typicallytruncateatbases,0–1(–3)-septate;walls smooth, thin, hyaline, and not constricted at septa; con-tentshyaline,attimesgranular;appendagesabsent. Habitat&Distribution—DeadleavesofYucca rostrata.ThisspeciesisknownonlyfromaninterceptionofMexicanmaterial.

Fig. 8a.StromataofKellermania ramaleyae(holotype,BPI525045)onleafofYuccasp.;b.conidiaofK. ramaleyae(holotype,BPI525045)onleafofYucca sp.;c.stromataofK. rostratae(holotype,BPI884092)onleafofY. rostrata;d.conidiaofK. rostratae(holotype,BPI884092)onleafofY. rostrata;e.conidiaofK. ramaleyaefromculture(MEP1260);f.conidiaofK. rostrataefromculture(JB5.16.11-01);g.conidiaofK. unilocularis(isotypeofPiptarthron uniloculare,BPI1110167)onleafofY. baccata;h.conidiaofK. uniseptatafromculture(AR3476);i.conidiaofK. yuccifoliorumfromculture(AR3472);j.conidiaofK. yucci- genafromculture(AR3470).—Scalebars=1mmforstromata,=30µmforconidia.

cb da

g

h i j

e f


Specimen examined.Mexico,InterceptedatLosIndios,Texas,USAondead leaves of Yucca rostrata,12May2011,coll.A. Garza,JB5.16.11-01(CBS131721)isolatedbyJ.F.BischofffromBPI884092(holotype),driedcultureonPDA(BPI882822).

Notes—Thisspeciesisdistinguishedfromtheothersthatoc-cur on Yuccabyitstypicallycylindricaltoobclavate,0–1(–3)-sep-tateconidiainculturethatlackappendages.Conidiaproducedonthehostwereaseptate.Inculture,septawereobservedonasmallpercentageofconidia.Somedifferencesinsizeofconidiawereobservedbetweenmaterialfromhostandinculture.

Kellermania unilocularis(A.W.Ramaley)Minnis&A.H.Kenn.,comb. nov.—MycoBankMB801104;Fig.8

Basionym.Piptarthron uniloculareA.W.Ramaley,Mycotaxon45:451.1992.

=Planistroma obtusilunatumA.W.Ramaley,Mycotaxon45:450.1992.

Onhost(isotypeofPiptarthron uniloculare):Conidia48–72× 8–9.5µm,Q=5–7.6(Lm=56.4µm,Wm=9.0µm,Qm=6.3),cylindrical, at times curved, apices straight, obtuse, less com-monly tapering towards, typically rounded to inconspicuously truncateatbases,2–4(mostly3)-septate;wallssmooth,thin,hyaline, andnot significantly constricted at septa; contentshyaline;appendagesabsent. Culture characteristics—Only sterile, irregular stromatalgrowthsobservedinthedriedculture. Habitat&Distribution—DeadleavesofseveralYuccaspp.(Ramaley1992).The typesof both theasexual andsexualstates were found on Yucca baccata (Ramaley 1992).ThisspecieshasbeenreportedfromtheUSA:CO,NV(Ramaley1992,thisstudy).

Specimens examined.USA,Colorado, LaPlataCo.,Durango, alongbiketrailbetweenChapmanHillskislopeandLion’sDen,ondeadleavesofYucca baccata,8June2000,coll.A.W. Ramaley,AR3466isolatedbyA.W.RamaleyfromAWR2001a,driedculture(BPI883224);RidgeS.ofSmelterMountain, Cactus ridge, on dead leaves of Yucca baccata,10Feb.1992,coll.A.W. Ramaley,AWR9018(isotypeofPiptarthron uniloculare,BPI1110167).

Notes—Theconidiaof the isotypewereobserved tobeslightlyshorterthanreportedbyRamaley(1992),andnoconidiawereobservedwith5septaalthoughRamaley(1992)reportedthatsomehad5septa.Thecultureofthisspeciesisdeadandonlyadriedcultureofthisisolateisextant,butRamaley(1992)reportedthattypicalconidiawereformedinculture.Thelinkbetween the asexual and sexual states is based on the produc-tionoftypicalconidiainculturesobtainedfromasci(Ramaley1992).An additional analysis was performed as the Kellermania analy-sisabovebutwiththeadditionofanexistingITSsequencefromK. unilocularisAR3466.Theresultssuggestedasisterrela-tionship between K. unilocularisandlineagei(pp=0.82).Thisexpanded clade with lineage i was member of a larger clade (pp=0.89)alsocontaining lineage ii (Fig.3).Theunilocularstromata of K. unilocularis is shared by members of lineage ii havingconidiawithappendages.Itslackofconidialappendagesissharedbymembersoflineageihavingplurilocularstromata.Itsaseptateascosporesdistinguishitfrombothlineages.Thisistheonlyspeciesinorcloselyrelatedtolineagesi–iii(Fig.2)withaseptateascospores.Itwasexcludedfromotherphylogeneticanalysesduetomissingdata.

Kellermania uniseptata(Morgan-Jones,NagRaj&W.B. Kendr.)NagRaj,inNagRaj,Coelomycetousanamorphs withappendage-bearingconidia:443.1993.—Fig.7,8

Basionym.Alpakesa uniseptata Morgan-Jones,NagRaj&W.B.Kendr.,Canad.J.Bot.50:879.1972.

=Planistromella torsifoliorumA.W.Ramaley,Mycotaxon55:265.1995.

Culture characteristics—Colonies on PDA 10–13mmafter 2wk at 24°Cwith a 12h light/dark regimen; surfacenearorangewhite(6A2),slightlyslimyinappearance,attimesportions covered with low, cottony hyphae; margin uneven, whitish; reverse off white to nearly concolourous with upper surface.Conidia27–45×8–17.5µm,Q=2–4.2 (Lm=35.8µm,Wm=12.4µm,Qm=3.0),cylindrical,fusiform,obclavate,to ellipsoid, apices tapering or not, obtuse, tapering or not towards and typically truncate at bases bearing indistinct frills, approx.medianly1-septate;wallssmooth, thinandhyaline,andnotsignificantlyconstrictedatsepta;contentshyaline,attimesgranular;appendagespresent,13–41.5µmlong,apical,3–5(–8),filiform,unbranched,flexuous,hyaline. Habitat &Distribution—Dead leaves ofYucca rupicola (Morgan-Jonesetal.1972a,Ramaley1995,thisstudy).ThisspeciesisknownonlyfromtheUSA:TX(Morgan-Jonesetal.1972a,Ramaley1995,thisstudy).

Specimen examined.USA,Texas,KimbleCo.,junctionofI10andU.S.Hwy.290,ondeadleavesofYucca rupicola,24Oct.1994,coll.A.W. Ra-maley,AR3476(CBS131725)isolatedbyA.W.RamaleyfromAWR9432,driedcultureonPDA(BPI882826).

Notes—Theconnectionoftheasexualandsexualstatesiscircumstantial(Ramaley1995).

Kellermania yuccifolia(J.G.Hall)NagRaj,inNagRaj,Coe-lomycetous anamorphs with appendage-bearing conidia: 443.1993.

Basionym.Neottiospora yuccifoliaJ.G.Hall,Phytopathology5:57.1915as ‘yuccaeafolia’. ≡Alpakesa yuccifolia(J.G.Hall)Subram.&K.Ramakr.,J.IndianBot.Soc.33:205.1954as‘yuccaefolia’.

Culturecharacteristics—Thisspecies isnotknown fromculture.Itischaracterisedbyitsaseptateconidiawithmultipleappendages(Hall1915,Morgan-Jonesetal.1972a,NagRaj1993). Habitat&Distribution—DeadleavesofYucca filamentosa, Y. gloriosa, and Yuccasp.(Morgan-Jonesetal.1972a,Sutton1980).ThisspeciesiswidespreadintheUSAbasedonscat-teredreports(Morgan-Jonesetal.1972a,Sutton1980,Farr&Rossman2012).ThetypewascollectedonYuccasp.intheUSA:WA(Hall1915).

Specimen examined —None.

Notes—NoDNA sequence data exist for this species,which is the type of the genus Alpakesa(Subramanian&Ram-akrishnan1954).Consideringthatthedatapresentedhereforother species of Kellermania indicate that conidial septation and appendage number are not important for distinguishing genera, this species belongs in the genus Kellermania.Theoccurrenceof this species on Yucca and its multiple conidial appendages suggestaprobablephylogeneticplacementinlineageii.

Kellermania yuccifoliorumA.W.Ramaley,Mycotaxon47: 262.1993.—Fig.7,8

=Planistromella yuccifoliorumA.W.Ramaley,Mycotaxon47:261.1993.

Culturecharacteristics—ColoniesonPDA27–40mmafter2wkat24°Cwitha12hlight/darkregimen;surfaceneardarkgreen(30F5–30F3),coveredwith low,cottony tovelutinoushyphae;marginuneven,whitish;reversegreytoblack.Conidia 40–65.5×8–11µm,Q=3.9–7.4(Lm=52.6µm,Wm=9.8µm,Qm=5.4),cylindricaltoobclavate,apicestaperingtowardsandobtuse, typically truncate to less commonly somewhat rounded atbases,2–3-septate;wallssmooth,thinandhyaline,andnotsignificantlyconstrictedatsepta;contentshyaline;appendages


present,attimesscarcelyvisible,possiblyabsent,upto1.5µmlong,apical,single,appearingasashort,dome-likemound. Habitat&Distribution—Dead leavesofYucca baccata, Y. brevifolia(holotypesofbothnames),Y. schidigera, Y. thorn-beri(Ramaley1993,thisstudy).ThisspecieshasbeenreportedfromtheUSA:AZ,CA,UT(Ramaley1993,thisstudy).

Specimen examined.USA,Arizona,MohaveCo.,0.3milesfromGemAcresRd.,exitmile20fromU.S.Hwy.40,ondeadleavesofYucca brevifolia, 3June1992,coll.A.W. Ramaley,AR3472(CBS131726)isolatedbyA.W.RamaleyfromAWR9229,driedcultureonPDA(BPI882827).

Notes—Ramaley(1993)reportedvariationinconidialsizesbetween specimens found on different hosts and between fresh anddriedconidia.Giventhediversityfoundinthisstudy,itisdifficulttostatewithcertaintythatallspecimensfromallhostsareconspecific.Theconidiafromthecultureexaminedhereweresmallerthanthosereportedfromthehostinnature(Ra-maley1993).Theconnectionoftheasexualandsexualstatesiscircumstantial(Ramaley1993).

Kellermania yuccigenaEllis&Everh.,J.Mycol.1:154.1885as ‘yuccaegena’.—Fig.7,8

=Planistromella uniseptataA.W.Ramaley,Mycotaxon47:267.1993

Culturecharacteristics—ColoniesonPDA19–25mmafter2wkat24°Cwitha12hlight/darkregimen;surfacewhitishtonearpalegrey(30B1),coveredwithcottonyhyphae;marginuneven, whitish; reverse greyish with black conidiomata vis-ible.Conidia37–56×8–13µm,Q=3.3–6.2(Lm=48.2µm,Wm=10.1µm,Qm=4.8),cylindricaltosomewhatobclavate,apices tapering towards or not and typically obtuse but some-times slightly acute, tapering or not, typically truncate at bases, approx.medianly1(–2)-septate;wallssmooth,thinandhyaline,and not or slightly constricted at septa; contents hyaline; ap-pendagespresent,5–35µm(Lm=19.6µm)long,apical,single,filiform,unbranchedtorarelybranchedwithasinglebifurcation,flexuous,hyaline. Habitat&Distribution—DeadleavesofseveralspeciesofYucca(Morgan-Jonesetal.1972b,NagRaj1993,Ramaley1993,Farr&Rossman2012).Thetypeoftheasexualstatewasfound on Y. glauca(asY. angustifolia)(Ellis&Everhart1885)and the type of the sexual state was found on Y. elata(Ramaley1993).ThisspeciesiswidelydistributedinthewesternhalfoftheUSA(Morgan-Jonesetal.1972b,Ramaley1993,Farr&Rossman2012).

Specimen examined.USA,NewMexico,ChavesCo.,mile302.05onU.S.Hwy.380,ondeadleavesofYucca filamentosa?,24Oct.1993,coll.A.W. Ramaley,AR3470(CBS131727)isolatedbyA.W.RamaleyfromAWR9325,driedcultureonPDA(BPI882828).

Notes—Theasexualandsexualstateswerelinkedthroughthe observation of typical conidia in cultures derived from asci (Ramaley1993).

ExCLUDED, POORLy KNOwN AND UNCERTAIN TAxA

Diatrype acervataEllis&Everh.,J.Mycol.4:75.1888. ≡ Planistromella acervata (Ellis&Everh.)M.E.Barr,Mycotaxon60:434.1996.

Notes—ThisspeciesdescribedfromYucca filamentosa in NewJersey(Ellis&Everhart1888)couldbethesexualstateof a Kellermania, but confusion around a species complex as well as a Stigmina anamorph have hindered the development of a proper species concept (Barr 1996).Other homotypicsynonymslistedinMycoBankarenotlistedhere.

Endothia parryiFarl.exCooke,Grevillea13:102.1885. ≡ Planistromella parryi (Farl.exCooke)M.E.Barr,Mycotaxon60:435.1996.

Notes—ThisspeciesfoundonAgave shawii(Cooke1885)hasplurilocularstromatawithseptateascospores(Barr1996).It may represent a distinct species of Kellermania or the sexual state of K. macrospora.OtherhomotypicsynonymslistedinMycoBankarenotlistedhere.

Hypocrea agavesMaubl.,Bull.Soc.Mycol.France19:292.1903.

≡ Plowrightia agaves (Maubl.)Maubl.,Bull.Soc.Mycol.France23:143.1907.

Notes—Thisspeciesbasedonmaterial fromAgave sp.inMexico(Maublanc1903)maybethesexualstateofaKel-lermania, possibly K. macrospora. It hasbeen treatedasasynonym of Planistromella parryi(Barr1996).

Kellermania attenuataMorgan-Jones,NagRaj&W.B.Kendr.,Canad.J.Bot.50:1643.1972.

Notes—Intheprotologue,thisspecieswasreportedtohaveconidia70–85×3–5µm,cylindrical,aseptate,appendageslacking (Morgan-Jonesetal.1972b).The type fromMexicooccurs on Yucca filifera, but other collections from widespread North American localities occur on other Yuccaspp.(Morgan-Jonesetal.1972b).Basedonthefigureoftheconidiaintheprotologue(Morgan-Jonesetal.1972b), the length towidthratio is12–15.ThoughtreatedasasynonymofPiptarthron limbatumbySutton(1980),wholatertreatedbothoftheseassynonyms of Piptarthron yuccae(Sutton1983),thisseemstobe a distinct species on Yucca.

Kellermania majorDearn.&Barthol.,Mycologia16:163.1924.

Notes—Intheprotologue,thisspecieswasreportedtohaveconidia55–75×11–14µm,2-septatewiththemiddlecellhalfthesizeoftheendcellswith1–2appendages,15–18×3–4µm(Dearness1924).AccordingtoDearness(1924),thetypefromtheUSA:CAoccursonHesperoyucca whipplei(asYucca whipplei;subfam.Agavoideae).Thereareafewotherreportsof this species on Yuccaspp.fromthewesternUSA(Farr&Rossman2012).ThoughtreatedasasynonymofKellermania anomalabyMorgan-Jonesetal.(1972b),Sutton(1980),andNagRaj(1993),thisisalmostcertainlyadistinctspeciesgiventhe importance of host associations since it is the only species described originally from the host genus Hesperoyucca.

Kellermania multiseptataMorgan-Jones,NagRaj&W.B. Kendr.,Canad.J.Bot.50:1644.1972.

Notes—In the protologue, this plurilocular specieswasreportedtohaveconidia50–68×6–7.5µm,cylindricaltoob-clavate,aseptate,3–4-septate,appendageslacking(Morgan-Jonesetal. 1972b).The type from theUSA:TXoccursonYucca macrocarpa and an additional collection was reported from theUSA:AZonYucca brevifolia (Morgan-Jonesetal.1972b).Basedonthefigureoftheconidiaintheprotologue(Morgan-Jonesetal.1972b),thelengthtowidthratiois8.2–12.ThoughtreatedasasynonymofPiptarthron macrosporum by Sutton(1980),thisseemstobeadistinctspeciesonYucca.

Phyllachora yuccaeEllis&Everh.,Bull.TorreyBot.Club22:440.1895.

≡ Piptarthron yuccae(Ellis&Everh.)B.Sutton,Trans.Brit.Mycol.Soc.81:407.1983.


Notes—In the protologue, this specieswas reported tobe an immature ascomycete and a question mark was placed nexttothegenustoindicateuncertaintyaboutitsclassification(Ellis&Everhart1895).According to theirobservations, theimmatureasci, if interpretedasconidia,aregivenas50–60×7–8µm,cylindrical,taperedtowardsapices,aseptate.Thetype from Mexico occurs on Yucca glauca (asY. angustifolia)(Ellis&Everhart1895).BasedoncorrespondencewithM.E.Barr,Sutton(1983)determinedthatthisspeciesrepresenteda Piptarthron and that P. limbatum and K. attenuata were latersynonyms.Ifthetypematerialisacoelomycetebearingaseptate conidia that lack appendages, this would likely be a distinctspeciescorrectlyclassifiedinKellermania.However,M.E.Barr’sannotationofthetypespecimenfrom1970(NY)states that the specimen is an immature K. anomala.

Plowrightia williamsonianaKellerm.,J.Mycol.12:186.1906.

Notes—Kellerman(1906)describedthisspeciesfromGua-temalan material found on Agave americana.Thecombinationof host, plurilocular stromata, and septate ascospores suggests a possible link with Kellermania macrospora.Ithasbeentreatedas a synonym of Planistromella parryi and Plowrightia agaves (Barr1996).

Septoplaca limbataPetr.,Sydowia17:272.1964‘1963’. ≡ Piptarthron limbatum(Petr.)B.Sutton,inSutton,Thecoelomycetes:54.1980.

Notes—Intheprotologue,thisspecieswasdesignatedasthe type of the genus Septoplaca, and it was reported to have indistinctly pseudoseptate conidiameasuring 35–60(–78)× 3–3.5 (Petrak 1964).The type from theUSA:AZoccurson Yucca macrocarpa (Petrak1964).Sutton (1977) treatedSeptoplaca as a synonym of Piptarthron, but he was unable to placethetypespecies.Assumingtheconidiamayactuallybeaseptate and after considering conidial measurements, Sutton (1977)suggestedthatS. limbata was an earlier synonym for Kellermania attenuata.Later,Sutton(1980)formallytransferredS. limbata to Piptarthron, treated K. attenuata as a synonym, and considered the species to be aseptate. Sutton (1983)treated these names as later synonyms of P. yuccae.AlthoughPetrak(1964)mayhaveconfusedpseudoseptateconidiawiththe euseptate conidia in Piptarthron, it seems unlikely that he wouldconfuseaseptatewithseptateconidia.

KEy TO KELLERMANIA SPECIES IN CULTURE

1.Colonydistinctlypink . . . . . . . . . . . . . . . . K. crassispora1.Colonynotpink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22.Conidiawithmorethanoneapicalappendage. . . . . . . . 32.Conidiawith0–1apicalappendages. . . . . . . . . . . . . . . 53.Conidiaaseptate . . . . . . . . . . . . . . . . . . . . . . K. yuccifolia*

3.Conidiaseptate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44.Conidia1-septate. . . . . . . . . . . . . . . . . . . . K. uniseptata4.Conidia2–3-septate . . . . . . . . . . . . . . . . . . . . . K. nolinae5.Someconidiawith1apicalappendage . . . . . . . . . . . . . 65.Allconidiawithnoapicalappendages . . . . . . . . . . . . . . 96.Manyapicalappendagesgreaterthan12µmlong. . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. yuccigena6.Apicalappendagesnotorrarelygreaterthan12µmlong 77.Allconidia1-septate. . . . . . . . . . . . . . . . K. nolinifoliorum7.Someconidiamorethan1-septate . . . . . . . . . . . . . . . . . 88.Conidia1–2(–3)-septate,Q=5–14.3,middlecellshorter

than end cells . . . . . . . . . . . . . . . . . . . . . . . . . K. anomala

8. Conidia2–3-septate,Q=3.9–7.4,middlecelllongerthanend cells ........................ K. yuccifoliorum

9. Conidiaaseptate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109. Someconidiaseptate . . . . . . . . . . . . . . . . . . . . . . . . . 1210. Conidiatypicallycylindricaltoobclavate. . K. micranthae10. Conidiatypicallycurved . . . . . . . . . . . . . . . . . . . . . . . 1111. Conidiaslightlycurved,falcate,tostronglycurved,nearly

U-shaped . . . . . . . . . . . . . . . . . . . . . . . . K. plurilocularis11. Conidiamore or less cylindricalwith irregularly curved

shapes, flexuous . . . . . . . . . . . . . . . . . K. dasylirionicola12. Conidiamostly0–1-septate. . . . . . . . . . . . . . . . . . . . 1312. Conidiamostlywith2ormoresepta. . . . . . . . . . . . . . 1413. Conidiatypicallycylindricaltoobclavate. . . K. rostratae13. Conidiatypicallywithsomedegreeofcurvature . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. dasylirionis14. Conidia1–3(mostly2)-septate . . . . . . . . . . K. confusa14. Someconidiawith4ormoresepta . . . . . . . . . . . . . . 1515. Someconidiawithmorethan5septa,Q=4.9–13 . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. macrospora15. Conidiawith5orlesssepta,maximumQ=±7.6 . . . . 1616. Conidia2–4-septate,Q=2.6–4.6 . . . . . . K. ramaleyae16. Conidia2–4-septate,Q=5–7.6. . . . . . . K. unilocularis**

*Not known from culture **Basedonstudyofisotypeonhost(BPI1110167)

DISCUSSION

Phylogenetic analyses support the placement and monophyly of the core Planistromellaceae in the order Botryosphaeriales of class Dothideomycetes where the Planistromellaceae rep-resent a well-supported clade comprised of the coelomycetous genera Alpakesa, Kellermania, and Piptarthron with associated teleomorphic genera Planistroma and Planistromella.WithinBotryosphaeriales, it is positioned in a polytomy with four other lineages of Botryosphaeriaceaes.l.,threeofwhichoccupybasalpositionsinpreviousstudies(Philips&Alves2009,Schochetal.2009)relativetothefourththatincludesthecorediversityof Botryosphaeriaceae(Fig.1).ThiscoreBotryosphaeriaceae lineage(pp=1.0)iscomprisedoftwomainclades,onewithBotryosphaeria(pp=1.0)andanother(pp=0.75)withPseudo-fusicoccum stromaticum and Guignardia(pp=1.0).Theposi-tion of Guignardiaissignificantduetoitsunclearpositionbasedonpreviousstudies(Crousetal.2006,Philips&Alves2009).ThoughthefamilyBotryosphaeriaceae has been recognised withabroadcircumscription, i.e.sensulato(Philips&Alves2009),andPlanistromellaceae could well be treated as a later synonym, we prefer to maintain the family Planistromellaceae as distinct from Botryosphaeriaceae s.str. and suggest thatfurthersamplingisneededtoclarifyfamilialclassificationwithinthe Botryosphaeriales.ThePlanistromellaceaeissignificantlyand phylogenetically distinct from the Botryosphaeriaceae.All genera of core Planistromellaceae are here considered to constitute one genus, namely Kellermania.Fourothergeneraclassified inPlanistromellaceae (Barr 1996) are redisposedas follows: Eruptio has been shown to be a member of Myco-sphaerellaceae (Verkleyetal.2004);Loratospora has been shown to be a member of the Phaeosphaeriaceae (Schochetal.2009);andnoneofthespeciesofMycosphaerellopsis including the type species, M. myricariae(Höhnel1918b),havebeen placed phylogenetically; Microcyclus,typifiedbyMicro-cyclus angolensis(Sydow&Sydow1904),isonlyrepresentedby DNA sequence data of Microcyclus ulei, and based on BLASTsearchesofGenBankusingavailableITSsequences,it does not belong in the order Botryosphaeriales.TheclosesthitsresultingfromtheBLASTsearchesaremembersoftheMycosphaerellaceae,butitsfamilialclassificationisuncertain.


Among the Planistromellaceae(Kellermania)arefivelineagesthat,withtheexceptionoflineageiv,areeachwellsupported.Lineagevistreatedseparatelyasitspositionisunresolved.These lineageswere recovered inall analysesof individualgeneswiththeexceptionoftheSSU.ThelackofresolutionintheSSUtreewasduetoalowlevelofsequencedivergence,and thusa lackof informativecharacters (Table2).Overall,the relationshipsamong the five lineageswerenot stronglysupported.In general, the phylogenetic groupings within the Planistromel-laceae (Kellermania) do not correspondwith the presenceor absence and numbers of conidial appendages nor with conidialseptation(e.g.multipleappendagesinlineagesiiandv;appendagesabsentinlineagesiandiv).Previousgenericcircumscriptions of these coelomycetes based on conidial ap-pendagesalonearenotwellsupported,afindingwhichwasalso recently observed among other genera of appendaged coelomycetes (Barber et al. 2011,Crous et al. 2012).Thetype species of Kellermania, K. yuccigena, and Piptarthron, P. macrosporum(≡K. macrospora),belongtoseparatelineagesiiandi,respectively,thatnotablyformaclade(pp=0.89)withK. unilocularisintheITSanalysis(Fig.3).Lineageii,whichin-cludes K. yuccigena, also includes three well-known species of Planistromella for which DNA sequence data exist including the type species, Planistromella yuccifoliorum(=K. yuccifoliorum).Thoughlineagesi–iiiinFig.2containonlyspecieswithseptateascospores, Planistroma obtusilunatum, linked to P. uniloculare (≡K. unilocularis), asmentioned above appears to belongamong these lineages (Fig. 3). It has aseptate ascospores(Ramaley1992).Otherspecieslackingascosporeseptation,namely Planistroma yuccigenum, the type of the genus and linked to P. pluriloculare(≡K. plurilocularis)andPlanistroma nolinae, linked to P. crassisporum(≡K. crassispora),belongtolineage iv and Planistroma kellermaniae, linked to Kellermania nolinae,representslineagev.Thus,Planistroma and Planis-tromella do not form well-supported clades if circumscribed byascosporeseptationalone.Consideringthebiologicalandphylogeneticsimilaritiesofthespeciesinthefivemajorline-ages and the lack of resolution among the lineages, there is little pragmatic value in recognising any subclades or lineages asdistinctgenera.Thus,theentirecladeishereinrecognisedas a broadly definedgenusKellermania, the generic name having priority. In accordancewith the changes enacted intheMelbourneCodeinregardtoArt.59(Norvell2011),wedonotrecogniseseparateanamorphicandteleomorphicnames.Morphological variation of conidia among isolates from different hosts or even the same host, especially in regard to append-ages, septation, and shape of conidia, is frequently an indicator ofthepresenceofdistincttaxa.However,slightdifferencesinsize, septation, andappendagesof conidia amongmaterialfrom fresh collections, herbarium collections, and culture were frequentlyobservedforagivenspecies(Ramaley1993,1995,thisstudy).Inspiteofthesedifferences,theoverallappearanceof conidia on the host and in culture is similar for a given species and species differences tend to be obvious when considering within species andamong speciesmorphological variation.DNA sequence data are required to distinguish species and to confirmspeciesrecognitioninambiguouscases.From an ecological and evolutionary point of view, it is interest-ing to note that several lineages including lineages i, sisters ii– iii,andiv(Fig.2)arelargelycorrelatedwithplanthostsfromdifferent host clades in the Asparagaceae, subfamilies Agavoid-eae and NolinoideaesensuAPGIII(2009).Thissuggeststhathost phylogeny is correlated with the phylogeny of these fungi tosomedegree.However,samplingwithinthesefungiremainslimitedanditistooearlytoformulatefinalconclusionsgiventhelackofresolutionamongthelineages.Althoughanumber

of Kellermania species have been recognised historically as oc-curring on numerous hosts, we expect that there are few, if any, plurivorousspeciesgiventheevidenceofhostspecificityandnumbersofcryptictaxafoundinthisstudy.However,multipleisolates were not available for species of Kellermania reported from numerous hosts, thus, this hypothesis needs further study as many members of the Botryosphaeriales are known to inhabit multiplehosts(vanNiekerketal.2004,Glienkeetal.2011).

AcknowledgementsThe authors extend heartfelt gratitude toAnnetteW.Ramaleyfordetailedobservationsoftheseunusualfungiaswellashercontribution of the cultures used in this study and her notes without which this studywouldnothavebeenpossible.TheauthorsthankJohnH.Wiersemafor providing the nomenclaturally correct names for some species of Yucca thatwere identified by others. JosephF.Bischoff kindly provided fungalmaterialsandothersupportfromtheinitiationofthisstudytoitslaterstages.LisaA.CastleburyiswarmlyacknowledgedforprovidinganITSsequenceof Kellermania unilocularis.PedroW.Crousgenerouslyassistedwiththepublicationofthismanuscriptandadvisedonmattersoffamilialtaxonomy.

REFERENCES

APGIII.2009.AnupdateoftheAngiospermPhylogenyGroupclassificationfortheordersandfamiliesoffloweringplants:APGIII.BotanicalJournaloftheLinneanSociety161:105–121.

BarberPA,CrousPW,GroenewaldJZ,PascoeIG,KeaneP.2011.Reas-sessingVermisporium(Amphisphaeriaceae),agenusoffoliarpathogensofeucalypts.Persoonia27:90–118.

BarrME.1996.Planistromellaceae,anewfamilyintheDothideales.Myco-taxon60:433–442.

CookeMC.1885.Synopsispyrenomycetorum.Grevillea13:100–109.CrousPW,GamsW,StalpersJA,RobertV,StegehuisG.2004.MycoBank:anonline initiative to launchmycology into the21stcentury.Studies inMycology50:19–22.

CrousPW,SlippersB,WingfieldWJ,Rheeder J,MarasasWFO, et al.2006.PhylogeneticlineagesinBotryosphaeriaceae.StudiesinMycology55:235–253.

CrousPW,VerkleyGJM,ChristensenM,Castañeda-RuizRF,GroenewaldJZ.2012.Howimportantareconidialappendages?Persoonia28:126–137.

DearnessJ.1924.Newandnoteworthyfungi.III.Mycologia16:143–176.DrummondAJ,AshtonB,BuxtonS,CheungM,CooperA,etal.2010.Ge-neiousv5.1.Availablefromhttp://www.geneious.com.

EdgarRC.2004.MUSCLE:multiplesequencealignmentwithhighaccuracyandhighthroughput.NucleicAcidsResearch32:1792–1797.

EllisJB,EverhartBM.1885.Newfungi.JournalofMycology1:148–154.EllisJB,EverhartBM.1888.Newspeciesof fungi fromvarious localities.JournalofMycology4:73–82.

EllisJB,EverhartBM.1895.New species of fungi.Bulletin of theTorreyBotanicalClub22:434–440.

FarrDF,RossmanAY.2012.Fungaldatabases,systematicmycologyandmicrobiologylaboratory,ARS,USDA.RetrievedMarch30,2012fromhttp://nt.ars-grin.gov/fungaldatabases/.

FarrisJS,KällersjöM,KlugeAG,BultC.1994.Testingsignificanceofincon-gruence.Cladistics10:315–319.

GlienkeC,PereiraOL,StringariD,FabrisJ,Kava-CordeiroV,etal.2011.Endophytic and pathogenic Phyllosticta species, with reference to those associatedwithCitrusBlackSpot.Persoonia:47–56.

GrahamSO.1959.Theeffectsofvariousreagents,mountingmedia,anddyesontheteliosporewallsofTilletiacontroversaKühn.Mycologia51:477–491.

HallJG.1915.NotesuponWashingtonfungi.Phytopathology5:55–58.HöhnelFvon.1918a.Fungi imperfecti. Beiträge zurKenntnis derselben.Hedwigia60:129–209.

HöhnelFvon.1918b.MycologischeFragmente.AnnalesMycologici 16:35–174.

HuelsenbeckJP,RonquistF.2001.MrBayes:Bayesianinferenceofphylo-geny.Bioinformatics17:754–755.

KellermanWA.1906.AnewPlowrightiafromGuatemala.JournalofMycol-ogy12:185–187.

KornerupA,Wanscher JH. 1967.Methuenhandbookof colour, 2nded.Methuen&CoLtd,London,UK.

LargentD,JohnsonD,WatlingR.1977.HowtoidentifymushroomstogenusIII.Microscopicfeatures.MadRiverPress,Eureka,CA.

MalkusA,ChangP-FL,SabinaMZ,ChungK,ShaoJ,etal.2006.RNApolymeraseIIgene(RPB2)encodingthesecondlargestproteinsubunitinPhaeosphaerianodorumandP.avenaria.MycologicalResearch110:1152–1164.


MathenyPB,LiuYJ,AmmiratiJF,HallBD.2002.UsingRPB1sequencestoimprovephylogeneticinferenceamongmushrooms(Inocybe,Agaricales).AmericanJournalofBotany89:688–698.

MaublancA.1903.Surquelquesespécesnouvellesdechampignonsinfé-rieurs.BulletindelaSociétéMycologiquedeFrance19:291–296.

McNeillJ,BarrieFR,BurdetHM,DemoulinV,HawksworthDL,etal.(eds).2006. InternationalCode of Botanical Nomenclature (ViennaCode):Adoptedby theSeventeenth InternationalBotanicalCongress,Vienna,Austria,July2005.ARGGantner,Ruggell,Liechtenstein.

MillerMA,PfeifferW,SchwartzT. 2010.Creating theCIPRESScienceGatewayforinferenceoflargephylogenetictrees.In:ProceedingsoftheGatewayComputingEnvironmentsWorkshop(GCE),14Nov.:1–8.NewOrleans,LA.

MoncalvoJM,LutzoniFM,RehnerSA,JohnsonJ,VilgalysR.2000.Phylo-genetic relationships of agaric fungi based on nuclear large subunit ribo-somalDNAsequences.SystematicBiology49:278–305.

Morgan-JonesG,NagRajTR,KendrickB.1972a.Generacoelomycetarum.V.AlpakesaandBartalinia.CanadianJournalofBotany50:877–882.

Morgan-JonesG,NagRajTR,KendrickB.1972b.Generacoelomycetarum.VI.Kellermania.CanadianJournalofBotany50:1641–1648.

NagRajTR. 1993.Coelomycetous anamorphswith appendage-bearingconidia.MycologuePublications,Waterloo,Ontario,Canada.

NiekerkJMvan,CrousPW,GroenewaldJZ,FouriePH,HalleenF.2004.DNAphylogeny,morphologyandpathogenicityofBotryosphaeriaspeciesongrapevines.Mycologia96:781–798.

NorvellL.2011.Fungalnomenclature.MelbourneapprovesanewCode.Mycotaxon116:481–490.

NylanderJAA.2004.MrModeltest,v.2.Programdistributedbytheauthor.EvolutionaryBiologyCentre,UppsalaUniversity.

PetrakF.1964,‘1963’.Septoplacanov.gen.,eineneuegattungderscole-cosporenSphaeropsideen.Sydowia17:271–274.

PhillipsAJL,AlvesA. 2009.Taxonomy, phylogeny, andepitypification ofMelanops tulasnei, the type speciesofMelanops.FungalDiversity 38:155–166.

PollackFG. 1947.Two additions to the fungi imperfecti.Mycologia 39:617–621.

PosadaD,BuckleyTR. 2004.Model selection andmodel averaging inphylogenetics:advantagesofakaike informationcriterionandBayesianapproachesoverlikelihoodratiotests.SystematicBiology53:793–808.

RamaleyA.1991.FungiofYuccabaccata1.Piptarthronpluriloculareanditsteleomorph,Planistromayuccigena.Mycotaxon42:63–75.

RamaleyAW.1992.FungifromYuccabaccata.2.Planistromaobtusilunatumsp.nov.,and itsanamorph,Piptarthronunilocularesp.nov.Mycotaxon45:449–460.

RamaleyAW.1993.New fungi fromYucca:Planistromella yuccifoliorum,gen.etsp.nov., itsanamorph,Kellermaniayuccifoliorum,sp.nov.,andPlanistromellauniseptata,sp.nov., the teleomorphofKellermaniayuc-cigena.Mycotaxon47:259–274.

RamaleyAW.1995.NewspeciesofKellermania,Piptarthron,Planistroma,andPlanistromella frommembers of theAgavaceae.Mycotaxon 55:255–268.

RamaleyAW.1998.NewteleomorphsoftheanamorphicgenusKellermania.Mycotaxon66:509–514.

RambautA,DrummondAJ.2007.Tracerv1.5.Computerprogramanddocu-mentationdistributedbytheauthorsathttp://beast.bio.ed.ac.uk/Tracer.

ReebV,LutzoniF,RouxC.2004.Multilocusphylogeneticcircumscriptionofthe lichen-forming fungi family Acarosporaceae and its position within the Ascomycota.MolecularPhylogenyandEvolution32:1036–1060.

RonquistF,HuelsenbeckJP.2003.MrBayes3:Bayesianphylogeneticinfer-enceundermixedmodels.Bioinformatics19:1572–1574.

Schoch CL, Crous PW,Groenewald JZ,Boehm EWA,Burgess TI,etal.2009.A class-wide phylogenetic assessment ofDothideomycetes.Studies inMycology64:1–15.

ShoemakerRA,BabcockCE. 1987.Wettsteinina.Canadian Journal ofBotany65:373–405.

StamatakisA. 2006.RAxML-VI-HPC:maximum likelihood-basedphylo-geneticanalyseswiththousandsoftaxaandmixedmodels.Bioinformatics22:2688–2690.

SubramanianCV,RamakrishnanK.1954.Alpakesa,anewgenusof theSphaeropsidales.JournaloftheIndianBotanicalSociety33:203–205.

SuttonBC.1968.Kellermania and its segregates.Canadian Journal ofBotany46:181–196.

SuttonBC.1977.CoelomycetesVI.Nomenclatureforgenericnamespro-posedforCoelomycetes.MycologicalPapers141:1–253.

SuttonBC.1980.Thecoelomycetes.Fungiimperfectiwithpycnidia,acervuli,andstromata.CommonwealthMycologicalInstitute,Kew,Surrey,England.

SuttonBC.1983.NomenclaturalnotesonDeuteromycetes.TransactionsoftheBritishMycologicalSociety81:407.

SwoffordDL.2003.PAUP*.PhylogeneticAnalysisUsingParsimony(*andothermethods),Version4.SinauerAssociates,Sunderland,Massachusetts.

SydowH,SydowP.1904.Novae fungorumspecies.AnnalesMycologici2:162–174.

ThiersB.2012[continuouslyupdated].IndexHerbariorum:Aglobaldirectoryofpublicherbariaandassociatedstaff.NewYorkBotanicalGarden’sVirtualHerbarium.http://sweetgum.nybg.org/ih/.

VerkleyGJM,Starink-WillemseM,IperenAvan,AbelnECA.2004.Phylo-geneticanalysesofSeptoriaspeciesbasedontheITSandLSU-D2regionsofnuclearribosomalDNA.Mycologia96:558–571.

VilgalysR,HesterM.1990.Rapidgeneticidentificationandmappingofen-zymaticallyamplifiedribosomalDNAfromseveralCryptococcusspecies.JournalofBacteriology172:4238–4246.

WhiteTJ,BrunsT,LeeS,TaylorJW.1990.Amplificationanddirectsequen-cingoffungalribosomalRNAgenesforphylogenetics.In:InnisMA,GelfandDH,SninskyJJ,WhiteTJ(eds),PCRProtocols:Aguidetomethodsandapplications.AcademicPress,Inc.,NewYork:315–322.

Documents

Phylogeny and taxonomic revision of the Planistromellaceae