Hikobial4:55-702003

MOlecularphylo窪enyOftheGrimmiales(Musci)basedOnchlOroplast mcLsequencesI

HIRoMITsuBoTA,YuMIAGENo,BELENEsTEBANEz,ＴＯＭＩoYAMAGucHIANDHIRoNoRIDEGucHI

TsuBoTA，Ｈ､，AGENo，Ｙ､，EsTEBANEz，Ｂ，YAMAGucHI，Ｔ＆DEGucHI，Ｈ、2003． MolecularphylogenyoftheGrimmiales(Musci)basedonchloroplastr6cLsequences・ Hikobial4:55-70．

FamilialcircumscriptionsoftheGrimmialesandtheirallies，includingthe Orimmiaceae，thePtychomitriaceaeandtheSeligeriaceae，havebeenrepeatedly discussedHerewepresentphylogeneticinferencesontheGrimmiaceaeandtheir ロ alliesbasedonmaximumlikelihoodanalysesofchloroplast76cLgenesequences・ Oursmdyshowsthat（１）theGrimmiaceae(s」at.)withtheinclusionofthegenera CtJ叩〕ﾉﾉosje/伽ａｎｄＰ〃c〃o〃か伽lismonophyletic;(2)Ｇ〃んo〃"伽hasaclose affinitytMrcjoaoftheDicranales;(3)theSeligeriaceaefblmadistinctcladesisterto theGrimmiaceae（s・lat)；（４）theGrimmiales（s」at.）withtheinclusionofthe SeligeriaceaearemonophyleticwhentheDrummondiaceae,theScouleriaceaeandthe Wardiaceaeareomitted;ａｎｄ(5)theDrummondiaceae-Scouleriaceaecladeappearsat thebasalpositionofthehaplolepidousmossgroupDicranidae，adjacenttothe Bryoxiphiaceae．SincethemonophylyoftheGrimmiales(s」at.)includingmembers oftheSeligeriaceaeiswellsupported,weproposeherethattheordercanbetreatedin awidesense,andincludesfamilieswhichhaveinｃｏｍｍｏｎｔｈｅＧ伽〃α/Mjgel･ja typeperistome、

H1、伽乃"Ｍα,Ｔｂ〃ｏ】'bwmg"cﾙﾉ＆ﾉﾌﾞﾉｧo"MDegz`chj,ＤＥｐｑ"ｍｅ"'ｑ/Ｄｊｏ/ogjcaノ 比je"Ｃｅ，Ｇｍａ"α/ｅ比/ioo／ｑ／Sbje"Ｃｅ,〃ｉｍｓｈｊ"ｚａＵ７ｍﾉeﾊﾉﾉﾘﾉ,ＫｔＪｇα"!(〕ﾉαｍα／-3-/， HjgZzsﾙﾉｰﾊjrosh〃α-sﾊﾉ,HIrosh〃α7398526,ノヒＷｎ ＹｊｌｍＭｇｅ"０，Depα"ｍｅ"／ｑ/Ｄｉｏ/OgjcaノＳｂ/e"Ｃｅ，ＦｔＪＭｌｌ〕ノｑ／Ｓｂｊｅ"Ｃｅ，脇、s/i〃α 肋加e'Fjlﾜﾉ,KtJgzJmJi〕′α"zα/-3-/,ノリｉｇａＷ－ｈ"ｏｓｈｊ"Ｔａ－ｓｈｊ,HIroSﾙ〃α739-8526,.ﾉﾋJpα〃 Be/6〃ＥＳ肋α"ez，Depα"α"Te"'０火Ｂｊｏ/oｇｊａ（Bor伽αZﾉ，Ｆａｃ"ﾉﾉα`ノ火Ｃｉｅ"ｃｊａｓ， 肋ⅣＣＭ/､１Ｍ伽"o"zα火MIC/rj仏280ｲ,α"/o6/α"ＣＯ,Mzc/ｒｊ仏Slpaj〃

TheorderGrimmialesisoneofthebasalmem- Orimmiaceae（asGrinunieae）andPtychomitria- bersofthehaplolepidousmosses，ａcconmodat- ceae（asPtychomitrieae）intheGrimmiales（as ingthetwofamiliesGrimmiaceaeandPtychomi- )inthefirsteditionof"Dienatiirli- triaceae・TheorderSeligeriales，accommodating chenPHanzenfamilien，，、Subsequently,Brotherｕｓ 品 thefamilySeligeriaceae，isalsooneofthebasal (1924-1925）transferredthePtychomitriaceaeto orders・Circumscriptionandtaxonomicposition thelsobryales、Nyholm（1954-1960)placedthe ofthefamiliesGrimmiaceae，， PtychomitriaceaeintheEubryales，inwhichare Seligeriaceaeandtheirallieshavebeenrepeat- includedthefamiliesprimarilywithadiplolepi- edlydiscussedAfterthefirstdescriptionofthe dousperistome:Bryaceae,Mniaceae,Aulacomnia- GrimmiaceaeasanindependentfamilybyArnott ceae，Meesiaceae，Timniaceae，Catoscopiaceae， (1825)，manyworkersvariouslyproposedtheir Bartramiaceae，Orthotrichaceae，andHedwigia- systemsontheGrimmiaceaeandtheirallies ceaeNoguchi（1954)discussedtherelationship (Tableｌ）Brotheruｓ（1901-1902）placedthe betweenPltW/io〃/伽加，Ｇ７加伽ａａｎｄＲａｃｏ〃ノー rj"mbasedonleafstrucmres，especiallyoma- lContributionfmmtheLaboratoryofPlantTaxonomy mentationofleafcells,ａｎｄｐｌａｃｅｄＰひc/io〃〃i"m andEcology,HiroshimaUniversity､ＮＳｅｒ・No.５２６． intheGrimmiaceae・Noguchi（1974)suggested 5６ ＨｉｋｏｂｉａＶｏｌｌ４,No.1，２００３ thatRaco伽かｊ"misanintennediategenusbe- PCRamplificationsofsegmentsfbr『bcLgene， tweｅｎＰｔＷｈｏｍかj"ｍａｎｄＧｒ加伽ａｏｎｔｈｅｂａｓｉｓ anddirectsequenceanalyseswerecarriedoutus- ｏｆｔｈｅｌｅａｆｃｅｌｌｓａｎｄｅｘothecialcelldevelopment、 ingstandardconditionsaccordingtothemanufac- Deguchi(1979)treatedP〃c〃o加伽"ｍａｓａｍｅｍ‐ turer'sinstructionsReactionswereperfbnnedin beroftheGrimmiaceaeemphasizingthemorpho- athennalcycler（ThermalCycler9600,Perkin logicalandanatomicalaffinityofcalyptra，leaf Elmer)withPCRamplificationkitsandsynthetic cellsandsetatwistingVitt(1984),andBuckand primers(TsubotaeM/200lb)．DirectDNAse‐ Goffinet（2000）treatedthePtychomitriaceaeas quenceanalysisofthePCRproductswasper- anindependentfamilyoftheGrimmiaceaelnre- lbnnedusingABIorAmershamkitsaccordingto centyears，severalmolecularphylogeneticworks themanufacturer1sinstructions・Thesequences relevanttothephylogenyofhaplolepidousmoss- wereelectrophoresedonautomatedsequencers eshavebeencarriedout(LaFarge＆coworkers (ABI373AandABIPmSM310),andanalyzed 2000,2002;GofTmete/α/､2001),showingthe onthedata-analysisSystems、Thesequencesob- monophylyoftheGrimmialesandSeligeriales， tainedinthepresentstudyweresubmittedtothe ａｌｔｈｏｕｇｈｔｈｅｎｕｍｂｅｒｓｏｆＯＴＵｓｆｂｒｔｈeseorders ＤＤＢＪ/ＥＭＢＬ/GenBanklnternationalNucleotide seemtobenotenoughtoclarifyanddiscusstheir SequenceDatabaseCollaborationDetailsand phylogeneticpositionswithinthesystem・ fUrtherinfbnnationfbrDNAextractingprotocols， Theohjectivesofthisstudywere:（１）tocon‐ amplificationprimers，PCRprotocolsandthose structaphylogenetictreefbrtheGrimmiaceae， ofsequencingreactionsaredescribedpreviously theSeligeriaceaeandtheiralliesbasedonmaxi- inTsubotaandcoWorkers(1999,2000,2001a,ｂ） mumlikelihoodcriteria；（２）ｔｏｔｅｓｔｔｈｅｍｏｎｏ‐ Ｓｂ"'cesα"`Ｍ"αかze`、"α phylyofthefamiliesandclarifytheftlmilialand Dataset--9076cLgenesequencesfbr86moss genericrelationshipsWepresentberephylog- specieswereexaminedfbrtheanalyses(Table２， eneticinferencesfbrthefamiliespreviouslytreat- withＤＤＢＪ/EMBL/GenBankAccessionNos.） edintheGrimmialesandSeligeriales，basedon The76cLgenesequencesofthegrinmialeanand themaximum-likelihood(ＭＬ)analysesof90se- seligerialeanmossesweremanuallyalignedwith quencesofthechloroplastribulｏｓｅｌ,5-bisphos- ７１mosssequencesregisteredintheDNAdata- phatecarboxylase/oxygenaselargesubunit(『bcL)， base、Outgrouptaxawereselectedbasedonthe togetherwithrecentinfbnnationonthemolecular phylogenetichypothesispropｏｓｅｄｂｙＬａＦａｒｇｅｅ／ phylogenyofgrimmialeanmosses，anddiscuss α/､(2000)andTsubotaetal.(ｉｎｍｓ.,andtheab‐ theimplicationsofourfindingsontheGrimmia- stractoftheMOSymposium2003） ceaeandtheirrelatedfamilies． 』ﾉﾉg""?e"j-Manualalignmentwascarriedout ibrthedatasetagainstacompleteγbcLsequence

Materialsandlnethods ofLe"co6M/"zsca6r"ｍ(ABO29388)withanedi- torprogramandouroriginalprogram，Sclean Phylogeneticanalysesbasedonther6cLgene 287（Tsubotal998-2003，inourhttpsites,see sequenceswereperfbnnedfbllowingthemethods AppendixB)．Undetenninedsites,gaps,andre‐ ofTsubotaandcoworkers（1999,2001a,2002)． gionsnotclearlyalignablefiDrallsequenceswere ② Themethodconsistsoftwosteps:（１）detennin- excludedfiomtheanalyses・Amongsequences ingsequencedata(DNAextractionPCRamplifl- aligned，identicalsequencesarerepresentedbya cationandDNAsequencing，orDNAdatabase singlesequence,andanunalignablesequencewas homologysearch);and(2)dataanalyses， omittedftomthedatasets．

ＤＭ欧伽ajo",ＰＣＲ。〃/抗cα/ｊｏ〃α"`/Ｄ/V,４ A"αｌｂＭＭｍ"oqb Se9"e"cmg Treeswereconstructedbythefbllowingthree TotalDNAiswereextractedfi･omfiPeshsamples methods:theneighborjoining(NJ;Saitou＆Nei ordriedherbariumspecimensbymodificationsof l987)，themaximum-parsimony（ＭＰ；Fitch thｅＣＴＡＢｍethod(Murray＆Thompsonl98u l971),andthemaximum-likelihood(ＭＬ;Felsen- Arikawa＆Higuchil999;TsubotaeM2000)or steinl981)．Thetreesobtainedbythethree phenol-chlorofbrmmethod(TsubotaeM/1999)． methodswereappraisedbythelog-likelihood HTsuBoTA,Ｙ・ＡGENｏ,BEsTEBANEz,ＴＹＡＭＡＧｕｃＨＩＡＮＤＨ・DEGucHI 5７ measurewithNucMLinMOLPHY23b3(Ada- MJx〃"川ﾉﾉﾋcﾉﾉﾉjooaα"α加応一MLtreeswere chi＆Hasegawal996,seeAppendixB） constructedwithNucMＬｉｎＭＯＬＰＨＹａｎｄｆａｓｔ－ Ｄｊｓｊα"Ｃｅα"α11〕入,jJ-ANJtreewasconstructed ＤＮＡｍｌ1.2.2(Olsenetα/､1994)．InMLanaly- withNuｃＭＬａｎｄＮＪｄｉｓｔｉｎＭＯＬＰＨＹｕｓｉｎｇｔｈｅ ses，thebest-confbnnedtransition/transversion ＨＫY85model（Hasegawae/αＬ1985）fbrthe (Ts/ＴＶ）parameterwasestimatedbyNucML distanceestimations accordingtocalculationswiththeHKY85model PZJ芯加o'1Ｗ"αOA,js-MPtreeswereconstructed employingthe“-topt，,optionparameteras2､093 byPAUPRat(Sikes＆Lewis2001),whichisa (as4185intheNucMLnotationalsystem)．ML tooltoimplementtheParsimonyRatchetsearches treeswereconstructedwiththelocalrearrange- (Nixonl999)withPAUP*,overPAUP40blO mentsearchffomtｈｅＮＪｔｒｅｅｂｙＮｕｃＭＬａｎｄ (Swoffbrd2002)．hanalysesbyPAUPRat,MP ＮＪｄｉｓｔＩｎａｎａｌｙｓｅｓｂｙｆｔｌｓｔＤＮＡｍｌ，MLtrees treesweresearchedusingtheParsimonyRatchet weresearchedwiththeglobalrearrangements searchstrategyintwenty200iterationruns． optionsTheTs/Tｖｐａｒａｍｅｔｅｒｗａｓｔｈｅｎｕｓｅｄａｓ

Table１．ＣｌassificationsoftheOrimmiaceae,PtychomitriaceaeandSeligeriaceae,orpartsoftheftlmily， andpossiblyrelatedgenera・Thetableshowsthedistributionoffamiliesandgenerathathavebeencon- sideredtobelongingtotheGrimmialesandSeligerialesatonetimeoranotherandincludedinthepresent analysis． Ｂｍｃｈ＆Schimper Schimper(1876）Limpricht(1885-1889）Brothems('901-1902）Fleischer(1915-1923） (1837-1846） Seligeriaceae Seligeriaceae 、icranales 6MgUJ7a Seligerieae Stﾀﾉﾙg［?rm Seligerieae Seligeriaceae Bﾉﾉ"c/ｍ Sｾﾉ'9℃rm Bﾉﾉ"ぬａ 比｣I/gcr/α StWgw、 ＣｂｍＰﾘﾉﾉbsje"zｲ耐 Blindieae β〃7c/7ａ βﾉﾉ"cﾒﾉα B"、ﾉｶﾞﾋJ Campylosteliaceae OIthotrichaceae Brachyodonteae Bmcﾉ(〕ﾉﾋﾞﾉb""""１ Grimmiaceae P〃Ｃ〃ｏｍｊｗ""l Cmγ収ｿﾉbsだ"""７ ｃｍ"ﾉﾌﾘﾉﾉＯＳ雌ﾉﾉ""l Ptychomitrieae Grimmiaceae Gblp"o"""７２ｨ"’ G卯/､'''/かｊ"ｍ Sbo"ん'･ｍ ＣＤＳＣｊ"odb〃 Grimmiaceae Grimmiaceae ＰｌＤﾉcho'"ｊ"ｍｍ Ｃｂｓｃｊ"ocjb〃 Grimmieae Cinclidonteae ＥＵ《gOlMo""""'"７ Ｓと〃jSj減""’ Grimmiaceae Gr/"ｍｍＪ C/"cﾉﾉﾋﾉＷ"ｓ ｃｔJ"ﾘﾘﾉﾉnFje"""１ ０r/〃〃ね Sb〃ｊｓｊ城""ｊ Ｒａｃｏｍｊかｊ"m （Sbo"ん'ね） Scoulerieae Ｒ力αＣＯ"""７""Z Ｇ'･〃”ｊａ Ptychomitrieae Ｇｒｉｍｍｉｅａｅ SboMIerm Ｒａｃｏ"""７"'〃 CoFcj"ocﾉb〃 Ｓｂ"1F'jdh/"l Grimmieae Isobryales Ｐ〃c〃ｏｍｊ"7"m CQFc"Todb〃 Cosc/"ＯＬ/b〃 GOlpﾉｍｍｊ"７""’ Ｇﾉﾌﾞ"1"ｊｍ Ｇ'．ｊ"ｌ"Jliq EJipoa7""' Ｒａｃｏ〃"'７""l Grj"'""α 随"/"rje"b「 Ptychomitrieae Ｓｃ"ｌＷｊ的Ⅲ"ノ Orthotrichaceae Bmc伽花ﾙﾉｨ"’ Ｒｈａｃｏ"ﾉﾉ〃/""Ｉ G〃んo"'ｊ"７""， （PI(〕ﾉc力o""か/""7） Ｃ、"Pﾘﾉﾉbsr/e"Ⅲ"】 （GOlp/io"""72J"'） （Ch1”γﾉbs'e""腕）

Dixon(1924） Brothems(1924-1925）Jones(1933)inGrout，NyhoIm('954-1960）Churchill(1981） ＆GrouMl936） Dicranaceae Dicranales Grimmiaceae Seligerieae Seligeriaceae Seligeriaceae Seligeriaceae Grimmioideae Sb/jgU7m「 Sc/jg〃ね Scﾉigcrm Bﾉﾉ'7., GJ7"】"】ね「 Dicraneae β肋めα B""｡、 sｾﾉﾉgヅノα Ｓどんな'枕""７ B""ぬａ Ｂ'pcﾉiyzjb""z'"’ Coscinodontoideae Grimmiales Grimmiaceae Cbsc/"ＯＬﾉb〃 Grimmiaceae Grimmiaceae Grimmieae Grimmiales Ptychomitrioideae G7”ｍｍ Scoulerioideae QIWm"Ｊｊかj""I Gnmmiaceae R“ｏｍｊ〃、"Ｊ ＲﾉﾉαＣＯ""か'"〃 Sbo"た〃α Grjmmm Sとﾉｸﾊﾟﾉ減"ｍ Ｃｍ７ＶリノnFze"""ｊ CQFcmodb〃 Grimmioideae Scoulerieae Ｃｂｓｃｊ"oab〃 ＰＯﾉｃｈｏｍｊかｊ""l ＰＯﾉｃｈｏｍｊｗ"ｍ CnSlc/"oclb〃 Sbo"んrm ｏｒ/加加/α Ｇ卯"o〃ｊ"'""】 Ｇ""ｍｍ Ptychomitrieae Ｒ"αＣＯ〃ｊＷ１〃 CtJ"v2ymFje""、 ＳｂｶﾊﾞF"cﾉﾉﾉｨｍ Ch｢"リリノoJMe"ｌｌｍ Sbo"ん〃ａ Ｒａｃｏｍｊ〃ｊ"” Ｐ〃cﾉ7o"'/ｗ""ｊ Eubryales Ｒｈａｃｏ"""'ｗ、 Ptychomitriaceae lsoblyales Q1Mpho'ｗ７"ｍ Ptychomitiaceae Ｐｌ０/c"o""か/"〃 CtJ”jMbsjeﾉｶﾞ"､ ＧＩ(HP力o"'"'/''''， Ｐﾊﾉcho”/か畝、 5８ HikobiaVoLl4,No.1,2003

TableLContinued

Vitt(1984） Noguchietal Iwatsuki(1991） Buck＆Goffinet(2000） (1987-1988 Iwatsukietal.（2００１ OIthotrichineae Seligeriaceae Dicranales Grimmiales Erpodiaceae B"ｍｍ Seligeriaceae Grimmiaceae A"ノロcqpノルm ＳＭｇＵ７ｍ B""c加 Aﾉig7mT腕m Ezpoczj"加 Ｂｍｃﾉ！〕ﾉﾋﾞﾉb"""腕 比ﾉｵg［ｗａ Cbscmocjb〃 陀"/"rje肋 Ｂｍｃ/i〕ﾉzjb"""〃 COFcmodb"ｊＭｂ 陥ﾉﾋﾉ、 Grimmiaceae ＤＪ,ｌｐｍｄｍ Cbscmodb〃 Grimmiales Ｇ７ｊ'""/α Leucodontineae Ｇｒｍｍｍ Grimmiaceae 勿伽sjeﾉﾉﾛ「 Wardiaceae Ｒａｃｏ〃ｊか/"〃 GJ7"ｗｊａ ノヒ1ｶﾞi`e"o6ひ"ｍ リグbｍ７ａ Ｐ〃cAomjかｊｚｨｍ Ｓｂｈｉｙ/城"ｍ Ｌｅ"cclpe'℃力αe""〃 ＣｔＪ"ql2〕ノルs花"""’ Ｃｏｓｃ/"oclb〃 Ｒａｃｏｍｊか加加 Seligeriineae Ｒａｃｏｍ/か/""Ｔ ＳｂｈｊＳｊｊめ"m Seligeriaceae Erpodiaceae ＰＩ(ｗｈｏ"/かｊ"” Ptychomitriaceae B肋ｄｍ JC"〃r/e肋 ＣｔＪ７"ＩＤﾉﾉｍｊｅ""" CtJ"１回〕ﾉﾉｍｊｅ""腕 Ｂｍｃﾉ！〕ub"""" Ａ"ﾉﾋJcclpﾉﾉﾉｨ” GOlPho""か/"〃 Ｈｗ"e"o/omqpsJs Ｇ卯ｈｏｍか/"" Erpodiaceae Ｐリc力ｏｍ/かjqpS/Ｓ MigE7m Gblph｡"''"j""ノ Ｐ〃c"０ｍ"ｊ"m Sbﾉ伽岬"" 比"/"γｊｅ化 Scouleriaceae ７>Dcho6ひ"加 Ａ"ノロcqPﾉﾊｨ" Sbo"ん7ｍ 陀〃"c肱"３ 〃ｊｃﾉb""""〃 Drunmondiaceae Grimmiineae DJwmmo"dia Ptychomitriaceae Oz"qPﾘﾉｏｓｊＵｇ７"〃 Seligeriales Gblphomjかﾉﾂﾞ"' Seligeriaceae ＰＩＯﾉchomjZr/qPsjs B"ｍｍ PlOﾉｃｈｏｍｊか/"m Ｂｍｃ/1〕ﾉﾋﾞﾉb"""" Grimmiaceae たりﾉｍｅ"ｏｂ腕ｑｐｓｊｓ A"g7jmmja ＭｉｇＵ７ｍ CbScmodb〃 Ｚ>'ocho6〃"m ＣＯｓｃ/"oc/b"だ肋 Wardiaceae Ｄｒ〕lpmdb〃 リリb"〃ａ Ｇ７/”ｍｊａ ＬＯｎ>Ｄ８ｍｍＴｍ 〃ぬｨsjC肋 JIq版`e"０６び"m Ae"｡bﾉ1yqpﾊﾉﾉｂ ＲａＣｏｍｊかｍｍ Ｓｂ力むｍＺｊ"ｍ Ｓｂｏ"んrね

２．０９３fbrtheanalysesbyfastDNAmL cedures：theapproximatelyunbiased（AU）ｔｅｓｔ T7eeＣＯ"qpa7zso〃ｗﾉﾉﾉｉノog-ﾉﾉﾉMjhoocノα"‘ｓｅｖ－ (Shimodaira2000，2002）usingthemultiscale ｅｍＩ/tesZF-TreecompansonwiththeMLcriteria bootstraptechniquqandtheBayesianposterior wascalTiedoutbyNucMLtoevaluatetheresult- probability(PP)calculatedbytheBayesianinfbr- ingtreesSeveralmeasureswereusedestimating mationcriterion（BIC）approximation（Schwarz thelog-likelihoodofdiffierenttreetopologies,the l978;Hasegawa＆Kishinol989)． standarderrors(SES)ofthediffercnceinlog-like- BooZiW叩卿ｊ－ＵｓｉｎｇＮｕｃＭＬ,abootstrapping lihoo｡(Kishino＆Hasegawal989),andtheboot- testwithlocalbootstrapprobabilities(Adachi＆ strapprobabilityestimatedbytheRELL(resam- Hasegawal996;LBPsiｎ％)wasappliedfbrthe plingofestimatedlog-likelihoodofsites)method bestMLtopology・LBPisarelativebootstrap (KishinoetaLl990;Hasegawa＆Kishinol994)． ffequencyobtainedfiomatopologDノsearchby TheprogrampackageCONSELOlf(Shimodaira localrealTangementsofNucMLThevalueis ＆Hasegawa200Dwasalsousedtocalculatethe comparablewiththatofFelsenstein,ｓ（1985） p-valuesoftheconfidencefbrthebifilrcating bootstrapprobability,anditisalittlelargerthan candidatetopologiesusingseveraltestingpro－ thestandardbootstrapprobability． HTsuBoTA,Ｙ・ＡGENo,BEsTEBANEz,TYAMAGucHIANDH・DEGucHI 5９

Table2Listofspeciesandfamiliesinvestigatedfbrr6cLgenewiththeaccessionnumberandvoucheror referenceofthesequences、TreatmentoffamiliesfbllowsBuckandGoffinet(2000）Fultherinfbnnation fbrthesequencesobtainedinthepresentstudyisshowninAppendixA

Class/Order/Familv/SDecies AccessionNo References/Ori2in Grimmiales Grimmiaceae(sstr） COscmo`ｌb"c"６，s"s(Hedw）Ｓｐｍｃｅ ＡＢ１２５５７５ Thissmdy Ｇｒｍ"ｊａｑＰｊｃⅢﾉmZJHornsch ＡＢ１２５５７６ Thisstudy GrjmmmeﾉmgmZJKaulf ＡＢｌ２５５７７ Thissmdy GrmmmﾉﾋWgzJm(Brid)Ｂｒｉｄ ＡＦ２３１０８１ MishlcretaL(Database） Ｇｍ'mljZJノロcvjgmZJ(Brid)Ｂｒｉｄ ＡＦ４７８２３０ Magombo(2003） Ｇｒｊｍｍｊａｏｒ６ｊｃｚＭａｒｊｓＢｍｃｈ ＡＢｌ２５５７８ Thissmdy Ｇ７/mmmIpﾉﾉi/bmRBeauv ＡＢｌ２５５７９ Thissmdy Grjmmmrp"んｊ"αね(Hedw）Ｓｍ ＡＢ１２５５８０ Thissmdy Ｒａｃｏ"j"ｊ"、/bscjcuノロ”(Hedw8)Brid ＡＦ２３１０８２ Mishleretal.(Database） Racomj"ｊ"'MIscjc"ノロ”varn”Wrj士Ｃａｒｄｏｔ ＡＢl25581 Thisstudy Ｒａｃｏｍｊ"ｊ"加八ＭＤＯ"ｊｃ"mDozy＆Ｍｏｌｋ AB125582 T1nsstudy Ｓｂﾉ７１F"成"mqpoc"p""(Hedw)Ｂｍｃｈ＆Schimp AF231065[aｓＧ７ｊｍｍｍｌ Mishleretal.(Database） qpoca7pα］ SbhjF'域"”sかｊｃ/"、(Tilmer)LoeskeexMartensson ＡＢ１２５５８３ Thisstudy Ptychomitriaceae Cm'ﾘﾘﾉﾉｂｓ'e""msZzxjcoﾉﾋJ(RWeber＆DMohr)Bruch＆ ＡＢｌ２５５８４ Thissmdy Ｓｃｈｉｍｐ Ｇ卯ﾙo"ﾂﾉかｊ"、ﾉｸ""'ｊ"ｊ"，""'(Mitt）Cardot ＡＢｌ２５５８５ Thisstudy G〃んo"ｊかｊ""ｗｊ""ｊｍＦｍ""(SOkamura)Ｂｒｏｔｈ ＡＢｌ２５５８６ Thissmdy Ｐ０ﾉcho"Ｔｊかｊ"伽｡〉ｗｍｍｏ"ぬｊ(Wilson)Sull＆Lesq ＡＦ２２６８２１ Wheeleretal.(Database） Pl1ycﾉiomjW"ｍｇｍｚＺ"erjLesq ＡＦＯO５５４９ GoHInetetaL（1998） POﾉｃｈｏｍｊかj"〃ｇ[JmherjLesq ＡＦ２３１３１３ LaFalgeetal(2000） P(yc/ｉｏｍｊかj"加咋"､'"腕(Mitt）AJaeger ＡＢｌ２５５８７ Thisstudy POﾉｃｈｏｍｊかｊ"ｍｗｊＬＦｏ"ｊｊＳｕｌｌ.＆Lesq ＡＢｌ２５５８８ Thisstudy Scouleriaceae Sbo"ノセグロα9"αjjcaHook AF226822 WheclcretaL(Database） Drummondiaceae D'ｗｍｍｏ"dmo6/"s(/bﾉｍＭＵｌｌＨａｌ AF232697 CoxetaL(２０００） Ｄｒ"mmo"c"p"'２ＰＣ"s(Hedw)EBritton AFOO5542 GoHInetetal(1998） Seligeriales Seligeriacｅａｅ Ｂ""dmiac"、(Hedw)Bruch＆Schimp AF226817 WheeleretaL(Database） B""c加加α9℃"ｔＪ"ｊｃａＳｃｈｉｍｐ AF478232 Magombo(2003） Ｂ""cZm/qpoJ7jcaBroth ABl25590 Tｈｉｓｓｍｄｙ MJgFrmα"sかjacaTSchauer AB12558９ Ｔｈｉｓｓｍｄｖ Wardiaceae ﾉﾘｂｍｍﾉｾXg7Dme"jaJHarv.＆Hook ＡＪ２７５１７０ Coxetal.(2000） Dicranales BIyoxiphiaceae B'〕ﾉoxjP"ｊ"ml7orvegjc"伽(Brid)Mitt ＡＦ２３１２９４ LaFaIgeetaL(2000） Fj”jc！blnsdb《6/"sRBeauv ＡＦ２３１３０３ LaFalgcetal.(2000） Dicranaceae A”/oα/bﾉbajα(Hedw)Loeske ＡＦ２３１２９２ LaFalgeetal.(２０００） d”､αたんe〃(Dicks）Ｂｍｃｈ＆Schimp ＡＦ２３１２９３ LaFalgeetal(2000） B'ＤＫ〃e”んα"α(SuU)MUllＨａｌ ＡＢｌ２２０３３ Ogurictal.(2003） Ｃα''１のノロP"sα'苫O’'ＤＣα'(ﾉbl7(MUllHal)Broth ＡＦ２３１３１５ LaFalgectaL(2000） CWlodb""""ﾉﾉﾋﾞ""erj(Schimp)Ｓｔｉｒｔ ＡＦ２３１３１８ LaFalgeetaL(2000） Ｄｊｃｍ"e/ﾉWjeje'Ｄ"'αﾉﾙ[(Hedw）Ｓｃｈｉｍｐ ＡＦ２３１２９６ LaFalgeetal(2000） Ｄｊｃｍ"ocjb"""”cノセ""c/b/"、(Brid)EBritton ABl22034 Ogurictal(2003） Djclm7owejsjacmmZJ(Hedw)Ｌｉｎｄｂ AF231297 LaFalgeetal(2000） Ｄ/c、""mscqpcW"ｍＨｅｄｗ・ AF478226 Magombo(2003） Kmerm6bﾉﾉﾉﾉﾉ(Bruch＆Schimp)Broth AFZ3l３０６ LaFalgeetal(2000） Le"ＣＯﾉbmar"/e"Ｍｇ７ｊ(MUllHal)ＣＨWright ＡＦ２３１３０８ LaFalgeetaL(2000） Oc/０６ﾉﾋp/、'w、α必ｊｄｉＪｍＨｅｄｗ ＡＦ２２６７９５ WheeleretaL(Database） Ｏ"CODﾉiolwswcZﾉiノセ"6e頑ｊｊＢｒｉｄ ＡＦ２３１３１０ LaFalBeetal(2000） 6０ ＨｉｋｏｂｉａＶｏＬｌ４,No.1,2003

Table2continued．

Class/Ordel/Familv/species AccessionNo Refbrences/Origin Dicranaceae O7的o成c、""、/iｲﾉw〃(Hook)GRoth ＡＦＺ３１３１１ LaFalgeetal(2000） Dicnemonaceae E"cα"!〃/o“〃ｍ１ｲeﾉﾙｸﾞﾉＨａｍｐｅ＆MtjllHal ＡＦ２３１３１９ LaFalgectal(2000） LeucobIyaceae Le"co6r〕ﾉ"ｍｓｃａ６'wmSandeLac ＡＢＯ２９３８８ T1subotaetal（1999） Djかｊｃ/ilmpa雌力〃(Hedw)Ｈａｍｐｅ ＡＦ２３１３０２ LaFalgeetaL(2000） Ｐノセ"r/d7"ｍａｃ"、ｊ"α"〃Lindb・ ＡＦ２３１３１２ LaFalgeetal.(2000） Rha6dbwｅｍｍｃ"""ん、(Mitt）HJameson AFOO5544 Go伍netetal（1998） Erpodiaceae A"ﾉbicqpj"腕力oqIg1Ini"so"me(Hampe＆MUllHal)Broth ＡＦＯＯ５５４５ GoHinetctaL(1998） 形"/"rjeﾉﾉbsj"e"sjs(Vent）ＭUllHal ＡＢｌ２５５９１ Thisstudy Bar6"ﾉﾋJCC"voﾊｲｍＨｅｄｗ、 AF478225 Magombo(2003） ＰＣ"jZJ伽je7med7α(TiJrner)FUrnr・ AB125592 Thisstudy PSC"`/bﾘｍ６ﾉＢｐﾉﾘCWS“"ｊ”erjα"α(Paris)ＨＡＣｒｕｍ AF226805 Whcelerctal(Database） 乃r/"ノロo6"（s“ｊ､α(MtjllHal)Mitt AFZ26823 WheeleretaL(Database） Zbr/"ﾉﾋﾝﾉwrα"s(Hedw)RGaertn,ＢＭｅｙ＆Scherb AJ275169 Coxetal.(2000） Qmbﾙﾉ"PC花W2e"ｊＳｗＢ AF226789 WhcelcrctaL(Database） ＬａｲcqPhα"esa必csce"sMUll､Hal、 AF226798 Wheclerctal.(Database） Ｍ７ｊ〃'”"〃０６/"s（/b""〃(Lindh)ＨＲｏｈ AF226777 Wheeleretal(Database） Ｓｂﾉ灯力qpodb"〃γjez"jjMont AF22678３ WheeleretaL(Database） Orthotrichales Orthotrichaceae 紐飯細紐躯眠紐征嘘師飯 ０００００００００００ ５５５５５５５５５５５ ５５５５５５５５５５５ ４４２２２２３２４４３ ３１３７４８７２７０２ A"!〃ん幼"〃ﾉﾋV1po"ｊｃ"〃(Hedw）Schimp ０００００００００００ Go伍netetal.（1998） B'〕ﾉodiXo"/ｑｐｅｒｊｃﾉｍｅｍﾉﾉSSainsbury GoBHnCtetal.（1998） Cmz/b"e化9"ｊ"9"ﾋﾞﾉﾋ、α(Hornsch)Ｖｉｔｔ GoHinetetaL(1998） Ｇ'Ｄ"/ｊｅﾉﾉﾋJqpjaｲﾉbjcJ(Hook)ＨＡＣｍｍ＆Steere Go伍netetal(1998） ノMYC'Ｄｃｏｍα/e""ｊｓ(Hook＆Grev)Ｖｉｔｔ GoHinetctaL(1998） ＭｂｌｃＪＤ"ﾉﾉ"ｊ""'ｊ"α"Ｗbﾉﾉ""！(Hook＆Grev）Schwiigm GoBHnetetal(1998） Orjhoかｊｃ/，"'''０６'"s(/bﾉﾉ2J”Ｂｒｉｄ GoBHnetetal(1998） Ｓｂﾙﾉ、ﾉFej"ｊｍ６１ｍＩﾉ"〃Schwiigr． GofIinetctal（1998） 〔ﾉﾉﾋﾞａｓ"w"lPaﾉ>"/ＣＯノロ(MnllHal)RHZander Go伍netetal(1998） 〔化、ルノeα(Hookf＆Wilson)Ｍｉｔｔ Go価netetal.（1998） 〃gDcﾉb"ｊ"lIer"eclli"sBruch＆Schimp・ GoHinetctal（1998） Hedwigiales Hedwigiaceae HedWjgmcﾉﾉﾉα、(Hedw)PBeauv・ AF478234 Magombo(２００３） ＨｍＷﾉｇｊａｃﾉﾉﾉα/α(Hedw)EBeauv AFOO5517 GoHinetetal（1998） BIyales Bartramiaceae BcJmacJ7mα/iα此ｒｍ"αＨｅｄｗ・ ＡＦ２３１０９０ DeLunaetal.(2000） Bryaceae Br〕ﾉ"ｍｃｑｐﾉﾉﾉﾛﾉ℃Ｈｅｄｗ、 ＡＹ１６３０２７ Pederscn＆Hcdcniis(Dababase） Mniaceae PﾉﾋJgjo"'"ｊ""'ﾉqpo"ｊｃ"'〃(Lindb)TJKop ＡＢＯ５０９９２ Tmbotaetal(ZOOla） Rhizogoniales Hypnodendraceae fblp"ｏｚｊｂ"ぬり"ｍｅ"ｚｊｅｓｊｊ(Hook)Paris ＡＦ２３ｌＯ９３ Mishleretal(Database） Hookerimles Daltoniaceae Be"ｊｍｍ"ｊａｅﾉj"76αねＨ､Akiy.,Ｔ・Ｙａｍａｇ＆Suleiman ＡＢｌＯ３３５１ Akiyamaetal.(2003） HypnaleS Ibpnaceae ノヲ)lp"""，Ｐｍ"'αq/bmleWilson ＡＢＯ29384 TmbotaetaL（1999） Leucodontaceae DozMJﾉqpo"ｊｃａＳａｎｄｅＬａｃ● ＡＢｌ２５５９３ Thisstudy Ｈ・TSUBOTA,Ｙ,AGENO,Ｂ,ESTEBANEZ,ＴＹＡＭＡＧＵＣＨＩＡＮＤＨ・DEGUCHI 6１

Table2continued

Class/Order/Familv/SDecies AccessionNo References/Ｏｒｉ２ｉｎ Outgrouptaxa FunariaIes Funariaceae FiJ"”ねﾉﾘ9ｍ”eかjcaHedw． AFOO5513 GoEHnetetal(1998） Ｐ伽c”ｊ"℃ﾉﾉｂＩｐａだ"s(Hedw)Ｂｍｃｈ＆Schimp ABO66207 Sugita＆Sugiura(Databasc） Encmlyptales Encalyptaceaｅ 勘cablPmm7堰lpmcaZpaHedw AF478239 Magombo(2003） Timmimleg Timmiaceae 刀加ｍｊαα"sかｍｃａＨｅｄｗ・ AJ275185 CoxetaL(2000） 刀加加mmegrMPo"ね"αＨｅｄｗ、 AF478242 Magombo(2003） Diphysciales Diphysciaceae DlPﾉｶ〕川Scｊ""んん加""ｍＭｉｔｔ AF478222 Magombo(2003） Ｄｉｐ伽c､”ﾉb｢加""〃(Cardot)Magombo,Zacharia AF478213[as】'舵rjoZjα Magombo(2003） Lekodi 〃71/b/､］ PoIytrichopsida Tbtraphidales Buxbaumiaceae B"x6α""jQpipe7jBest AF478211 Magombo(2003） Polytrichales Polytrichaceae PoOﾉ"/c"cmb妙"ｓａ７な､'"s(Hampe)Ｍｉｔｔ AF478207 Magombo(2003）

ＣＯ"se"s"ｓかee-50％majority-ruleconsensus Ｔｈｅdatamatrixwaspreparedfbrphylogenetic treesfbrthetopologieswithhigh-rankinglog- analysesbymanualalignmentofthesequences likelihoodvaluesandpassiｎｇｔｈｅＡＵｏｒＰＰｔｅｓｔ Fortheanalyses,atotalofl,l69sites(81.9％ｉｎ ｗｅｒｅａｌｓｏｃｏｍｐｕｔｅｄｂｙＰＡＵＰ蝋． 1,428bpoftheentiresequence)of90r6cLse- quenceswereusedNucleotidefTequenciesfiom

Results thedaｔａｓｅｔｗｅｒｅＡ＝２９４％，Ｃ＝１６．９％，Ｇ＝ 21.0％,ａｎｄＴ＝32.7％、Inthesitesofthealigned Ｏ６ｍｊ"e‘seQwe"Ｃｅ血ra r6cLsequences，３９５sites（＝33.8％inL169 New76cLseqUencesweredetenninedfbrl9 sites)werevariablc species：＊COscj"oc/O〃ノカ"〃/応，＊Ｇ７〃macJpjc"‐ P/Moge"erjcα"αlbAses /伽,で'.ｅ/o"g川,で7.ｏ'6ｊｃ"/α加,*Ｇｒ.ｐﾉﾉ娩、， Thedatamatrixwaspreparedfbranalysesof Ｇ'．ｐ"Ｍ"川，書Raco〃t7jz"〃んscjc"〃ｅｖａｒ． phylogeneticrelationshipsofthegrimmialeanand 〃ｏＶｊ７ｊｃ/e,ＹＲ,ノ叩o"zcz""，*ＳｂｈＭ血""0ｓかｊｃｊ""０， seligerialeanmosses，includingotherhaplolepi- ０J'，Ｗ/oste/伽Saxjco/LJ,で卯/to〃/伽加ﾉj幽伽ノー dousmossesAtotalof692topologies,ofwhich 伽"川，で/、〃""ｒｊＭｍ"腕，やlﾜﾉcﾙo〃かｊ"ｍｃ/b〃 onlylltreetopologieswerebifilrcate，ｗｅｒｅob- 〃""ＭＤｊ・WjLSO"ﾉﾉ,$Ｍｊｇｅ７ｊαα"Sr7jaCa,Ｂ/ｊＭａ tainedfiomthreeanalyses：１ＭＬａｎｄｌＮＪｔｏ－ ﾉ叩o"jca,*厄"〃ｊｅﾉﾉasj"elM,ＰＣ伽〃e""e伽α， ｐｏｌｏｇｉｅｓｂｙＮｕｃＭＬ；１ＭＬｂｙｆａｓｔＤＮＡｍｌ；ａｎｄ andDoZ〕′αノ叩o"jca,ｆｂｒｗｈｉｃｈｌ４ｔｈｅｓｐｅｃｉｅｓ ６８９ＭＰｂｙＰＡＵＰＲａｔｏｖｅｒＰＡＵＰ*４Ｍorede- ｗｉｔｈｔｈｅａｓｔｅｒｉｓｋｗｅｒｅｓｈｏｗｎａｓａｎｅｎｔｉｒｅｓｅ－ tailedtopologiesweresearchedthroughtheob- ｑｕencel,428bplongTheobtainedsequences tainedtreesusinglog-likelihoodmeasureand weresubmittedtotheDNAdatabaseunderthe severaltests(Table３)．HKY85modelwasused serialDNADatabaseAccessionNos・ａｓｓｈｏｗｎｉｎ astheestimatemodelwiththetransition/transver- Ｔａｂｌｅ２ａｎｄＡｐｐｅｎｄｉｘＡ,thelattercontaining sionparameterestimatedfiDmthedatasetfbrthe voucherinfbnnationThevoucherspecimensof MLtreeas4287．Thebest-supportedMLtree thesequencedmaterialinthepresentstudyare withthehighestlikelihoodvalueisshowninFig depositedinHIRQ LThelog-likelihoodvaluefbrtheMLtreewas Se9"e"Ceαﾉjg7zme"／ -l2066.0±５５４４．Ｔｗｏ５０％majority-rulecon－ 6２ HikobiaVoLl4,No.1,2003

Table３．Comparisonoflog-likelihoodscoresamongthe692resultingtopologiｅｓｆｂｒ/bcLsequencesinthegrim- mialeanmosseswiththeHKY85model(Hasegawaαα/1985）Thelog-likelihoodvalueswerecalculatedbyNuc‐ ＭＬｉｎＭＯＬＰＨＹ２３ｂ３(Adachi＆Hasegawal996),andallthep-valuesbyCONSELO1f(Shimodaira＆Hase‐ gawa2001）ffomlO,O00repetitions，Thelog-likelihoodvaluesofthehighestlikelihoodtreesaregiveninangle brackets,andthedifferencesinlog-likelihoodofaltemativetreeshDmthatoftｈｅＭＬｔｒｅｅａｒｅｓｈｏｗｎ±theirstandard errors(SE)．Thep-valuesthatarenotsignifiCantata＝O05areemphasizedinboldtypaTopologieswithasterisks intheAUandPPcolumnswereusedfbrtheconsensustrees(valuesaresupenmposedontheMLtreeinFig」)．●

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－４．４±４０．０ ＷＷＷＷＷ －４．４±２９３ －４．５±３８２ －４．５士４０１ －４５±４０６ 六六★★★ 、叩〉（叩ＵｎｍｕｎｍＵ⑪叩〉 『■且（叩Ｕ■■二へｍＵｎｍＵ 〆、〉つ□》つり》の〃▲（ｕ〃 ロ『）（ｚＵヘクムグハＵ句，ｊ ５５６７８ ６６７７０ ６９０７５ ４４４４３ ８７２２２ ０４９２９ ＭＭＭＭＮ ＰＰＰＰＪ 士士士士士 ’｜’｜’ 688 ●●●●● 0.000 0.002 PAUPRat 689 0.000 0.002 PAUPRat 690 0.000 0.001 PAUPRat 691 0.000 0.001 PAUPRat

692 0.000 0.０００ NuＣＭＬ＋NJdist Nooftopologiesfbr 665 ７ consensustree Note-AU:approximateunbiasedtest（Shimodaira2000,2002）usingthemultiscalebootstraptechnique,ＰＰ： BayesianposteriorprobabilitycalculatedbytheBayesianinfbnnationcriterion(BIC）approximation（Schwarz l978;Hasegawa＆Kishinol989),ＢＰ:bootstrapselectionprobabilityofFelsenstein(1985)estimatedbytheRELL resamplingmethod(KishinoetaL1990;Hasegawa＆Kishinol994）

sensustreeswerealsoobtainedfbrthe7and656 LBPvalueinMLtree，１００％supportinthecon- topologieswithhigh-rankinglog-likelihoodvalues sensustreefbrthetopologiespassingtheAUtest， thａｔｐａｓｓｅｄｔｈｅＡＵｏｒＰＰｔｅｓｔ,respectively；and 100％supportintheconsensustreefiortheto- thevaluesfbrpercentageofsupportedtopologies pologiespassingthePPtest]，100/100/100, fbreachbranchweresupenmposedinFigl 99/100/100,respectively)．WithintheDicranidae ThreelargecladescorrespondingtotheDicra- clade，ninesubcladeswereresolved：theGrim- nidae，Bryidae，andFunariaceae-Encalyptaceae， mialescladecolTespondingtotheGrimniales(s・ membersoftheFunariidae，wereresolvedwith lat.）withmclusionoftheSeligeriaceae（100／ highsupport（LBP/AU/ＰＰ＝９９/100/100［99％ 100/100);thePottiaceae-Erpodiaceaeclade(96／ HTsuBoTA,Ｙ・ＡGENｏ,BEsTEBANEz,ＴＹＡＭＡＧｕｃＨＩＡＮＤＨ・DEGucHI 6３

JOQUJJ BtjdjUﾉＴ 難癖艘鮒脇瘤 ７ｊＳｌｊｄＬ 88ノ ｎｊａｌａｅ ｍｊａ伯， 7ｍｍ'ａ ＭＬｔｒｅｅ 鑿 ｉａＱ'bｊ ７ｍｌａｐ ｍｌａＰ（

ＪｕｂＪﾛﾛﾌﾉ７，，ﾉ1ｍ １%;IiH； ＬＢＰ/Ａｕ/ＰＰ ＯｍｉＩｍｌ iales Ｒａｃｏｈ =|GⅢ ７ ｑｙｌｏｓｈ

－ ｃフ １００/７０ａ －ＰＩ /chom １ＵＷＵ ychom ｕｏノBljndia ﾌα1００Ｒｌｉｎ付 】佇 Ｔ i南!荒＝雛ｉｉ窺講鱗 B５ 96ﾉﾌﾞ００/9５ 型ili:ｉＨｉｉ=;簔壹i叢|蝋''1':`……… ＩＨＩ郷haPesaL ｃｅｎｓ

94ノ 【〕のｍｂｌｅＤｈ后 /dUm ＤＵｌＣ７ａｎｅｌｌａｈｅｌｅｍ ＤＤｊＣ７ａｎumscOpa， =＝Ｏ'thod/C'anun ヨグぴj白ｈｖＤｍｍｅｆｒ７ｂｆ 叩一一ＤＢ山’、

６ 池卯佗 蕊ililli繍

3,,鵬6Kﾉﾂ<ＲｌＩｌ;yiil鰯ョ 「一 l1Wmum .○ＷｕｍｌｍＵｍｌ７ mutissjmum ﾐJJEiJlI ●C ｗｎｈｌｐｎｆ elg〃 6３ 9０

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100ﾉﾌﾞ００/1００

■■ ０.O1substitutions/site

FigLThebest-supportedMLtreewiththehighestlikelihoodvaluefbrthealignedl,１６９bpofthe90r6cLgene sequences(HKY85model[Hasegawaerα/､1985];２α/β＝4.287;lnL＝-12066.0±S544byNucML);andthe cladeswithintheGrimmiales(slat.[＝sensupresent])clade、Thehorizontallengthofeachbranchisproportional totheestimatednumberofnucleotidesubstitutions・TherootisarbitrarilyplacedonthebranchleadingtothePoルー "jcAacﾉellp〃sα伽arzMbllowingMagombo(2003)andTsubotaetaﾉ.(ｉｎｍｓ.)．Localbootstrapprobabilitiesmore than５０％,andthevaluesofpercentageofsupportedtopologiesthatpassedtheAUorPPtestfbreachbranchare shownonornearbranches(LBPMU7PP;iｎ％） 6４ ＨｉｋｏｂｉａＶｏＬｌ４,No.1，２００３

100/95);theDicranalescladewithinclusionof theOrthotrichaceaeVitt（1982)establishedthe theWardiaceae(66/~/-);theFissidentaceae=｡"z‐ monogenericsubfamilyDrummondioideaeinthe pAjdmmclade（94/100/100)；theRhabdoweisia‐ Orthotrichaceae,andGoffinet(ｉｎＢｕｃｋ＆GofYi- ceaeclade（90/－/－)；theDitrichaceaeclade net2000)transfierredittotheGrimmialesasthe (92/100/100)；theDrummondiaceae-Scouleria- newfamilyDrummondiaceacTheScouleria- ceaeclade(94/100/100);andtheBryoxiphiaceae ceaeconsistoftwogenera比o"/ｅｒｊａａｎｄ７１ｒｊｃﾉ、－ clade・TheDicranalescladeandRhabdoweisia- 伽"z，andtraditionallyclassifiedintheGrim- ceaecladeweresupportedwithratherlowvalues・ mialesBuckandGoffinet（2000）treatedthe WithintheGrimmialesclade，fbursmallclades fburftlmiliesGrimmiaceae，Ptychomitriaceae， wereresolved：ｔｈｅＧ７加伽a-Coscj"ocllO"-SbAjE- ScouleriaceaeandDrummondiaceaeintheGrim- rjqlf"ｍ－Ｒａｃｏ〃trj"ｍｃｌａｄｅ（87/100/100)，ｔｈｅ mialeslnthepresentstudy,theMLtreeresolves ＱＪ〃〕ﾉﾉCs/eﾉﾉ"mclade，ｔｈｅＰｌ(〕ﾉcﾉﾉo〃"ｊ"'?ｚｃｌａｄｅ 比o"/ｅｒｊａａｎｄＤ'ｗｍｍｏ"c/ｊａａｓａｂａｓａｌｌｉｎｅａｇｅ (100/100/100)，ａｎｄｔｈｅＭｊｇｅ７ｊａ－Ｂ""ｄｊａｃｌａｄｅ ｗｉｔｈｉｎｔｈｅＤicranidae，supportingthesuggestion (95/100/98)． byLaFargeeM/､(2000)．Ourresultssupportthe placementoftheDrummondiaceaeasanindepen-

Discussion dentfiamilybutdonotsupporttheplacementof thefamilyintheGrimniales,suggestinganewly 1.OrdinalreIationships placementinani､dependentorderwiththe Thepresentstudysuggeststhealignmentofthe Scouleriaceae． Grimmiales（s」at．［＝ｓｅｎｓｕＬａＦａｒｇｅｅ／α/、 (2000);sensupresent]）asaresolvedmonophyl‐ Mjge7jα/“ eticgroupwithintheDicranidae，consistingof TheSeligerialesaccommodatesonlythesingle twofamilies:theGrimmiaceae（s」at.［＝sensu familySeligeriaceae，composedofthreegenera presentDwithinclusionofmemberspreviously Mjgerjαβ/ｊＭａａｎｄ〃αc/iﾘﾉ｡ｂ"""腕．Theorder consideredasmossesbelongingtothePtychomi- ischaracterizedbysmallplantsizeanddegener- triaceae，andtheSeligeriaceae；exceptfbrthe ateperistomalteethHistorically，theSeligeria- generaGjblp/jo〃"伽,、'w加加ＣＭα,Ｓｍ此加 ceae（andsomeworkersseparatedhac/iﾘﾉ血〃 andﾚﾘﾉＺＪｍｍ・Noafflnitywasshownbetweenthe 〃masanindependentfamily)wereplacedinthe membersoftheGrimmiales(s・lat.)andmembers DicranalesVitt(1984)segregatedtheSeligeria- ofthePottiaceae,ｎｏｒｗｉｔｈＪ/ＣＭｲｸﾞﾉeﾉﾉα、 ceaeintothesuborderSeligeriineae（Schimp.） Ｇｒ〃〃α/es Vitt.，describingdifferencesinalarcellsbetween CircumscriptionoftheGrimmialeshasbeen theSeligeriineae［＝Seligeriales］andtheGrim- variouslytransfbnned(Tablel).Brotherus(1924 miineae［＝Grimmiales］Manyworkershave -1925)andNyholm(1954-1960)[alsoFleischer treatedthemasthedistinctorderSeligeriales (1915-1923)］treatedtheGrimmialeswiththe (e､9.,Vittl984[assuborderSeligeriineaelBuck nalTowestsenseonlywithinclusionoftheGrim- ＆Goffinet2000)． miaceae・BuckandGoffinet(2000)Ｃｌassifiedthe Edwards（1979)proposedagroupingofgenera Grimmialeswiththewidestsensewiththeinclu- withMjgerjatypeperistometeeth,ｉｅ.,Ｍｊｇｅ－ sionofnotonlyfamiliestraditionallyplaｃｅｄｉｎ ｒｊａ,Ｂ/ｊＭａ，Ｇ！!〕lp/jo〃かj"ｍａｎｄＤｊｃＪｎ"ow伽α， ｔｈｅＧｒｉｍｍｉａｌｅｓｂｕｔａｌｓｏｔｈｅＳｃouleriaceaeand GoffIneMα/､（1998)alsosuggestedthat“though Drummondiaceae・Noguchierα/,（1987-1988)， themonophylyofagroupoftaxasharingthe Iwatsuki（1991）andDeguchi（inlwatsukieM/、 Mjge7ja-typeperistomehasnotbeencritically 2001）placedtheGrimmiaceae（s」at.）withthe examined,thenearlyidenticalperistomesofＲｈａ－ inclusionofQJ叩〕ﾉﾉosteﾉﾉz"〃ａｎｄＰｊｔ)ﾉＣＡＯ〃かj""z・ ｃＡｊｊＡｅｃｊ"腕（)，Ｑ【)lp/to〃かj"''１ OurresultssupportthetreatmentbyNoguchie／ (Ptychomitriaceae)，ａｎｄＢ""cﾉﾉα（Seligeriaceae） ａＬ（1987-1988),Iwatsuki（1991)andDeguchiin ｍａｙｂｅｓｅｅｎａｓａｎｉｎｄｉｃａｔｉｏｎｏｆｃｌｏｓｅphyloge- lwatsukierα/､(2001） neticrelationships，despitegametophyticdiffer- Ｄ７"ｍｍｏ"戒aHookischaracterizedbyveryre- ences.”ＬａＦａｒｇｅｅｒα/、（2000）madeaphy- ducedperistometeeth,andtraditionallyplacedin 1ogeneticstudyonthehaplolepidousmossesand HTsuBoTA,Ｙ・ＡGENｏ,BEsTEBANEz,ＴＹＡＭＡＧｕｃＨＩＡＮＤＨ､DEGucHI 6５ suggestedtheinclusionoftheSeligeriaceaerep- Schimper（1876),Brotheruｓ（1901-1902；1924- resentedbyBﾉﾉ"伽α伽α,Grimmiaceae,andPty- 1925),Vitt（1984),ＶｉｔｔｅｒａＬ（1998),andBuck chomitriaceaeintheGrimmiales(slat.),acon- andGoffinet(2000)includedCOsc伽伽〃and/ｏｒ clusionwesupport，andthefbrmationofoneof ＱｗＶﾌﾘﾉﾉosreﾉﾉ"minthePtychomitriaceae・Bro- basallineageofSbo"/e7jaa9"α/jcawithintheDi- therus（1924-1925）includedPltWﾉｶo〃かj"腕， cranidae，althoughthenumberoftreatedOTUs ＱｍＷ/osreﾉﾉ"ｍａｎｄＧ卯AC〃かj"minthePty- werenotenoughtoclarifythephylogeneticrela- chomitriaceaeplacingthemneartheOrthot- tionshipoftheGrimmiales、Inthepresentstudy， richaceaeinthelsobryalesJones(1933,inGrout theSeligeriaceaefbnnawellsupportedsister l928-4Dconsideredtherewasacloserelation grouptotheGrinⅡniaceae(s・lat.),althoughRhα‐ betweenGﾉｳlpho〃かｊ"ｍａｎｄＧ伽伽aontheba- c肋hecj"腕，Ｇ！!)lpho〃かj"碗，ａｎｄＤｊｃｒα"owezsJa sisofperistomecharactersDixon(1924),Nogu- donot chi（1974)，Deguchi（1979)，Churchill（1981)， BuckandGoffinet(2000)classifiedtheWardia- Iwatsuki（1991),andlwatsukiem/､(2001）have ceaeaswellastheSeligeriaceaeintheSeligeri- alltreatedtheGrimmiaceaeinthewidersense・ ales、Ourresultswouldindicatetheplacementof Noguchi（1954)reviewedJapaneseP〃c〃o〃－ onlytheSeligeriaceaeintheSeligerialesandsug- か〃mtreatingitasamemberoftheGrimmiaceae gesttheplacementoftheWardiaceaeintheDi- onthebasisoftheshapeandarrangementofleaf cranalesneartheDicranaceacTsubotaejα/､(ｉｎ cellsinsectionandsuggestedthattheBrotherus, ｍｓ.，ａｎｄｔｈｅａｂｓｔｒａｃｔｏｆｔｈｅＭＯＳｙｍｐｏｓｉｕｍ (1924-1925）ｐｌａｃｅｍｅｎｔｏｆＰ秒c〃o〃かｊ"ｍａｎｄ 2003）alsosupportthemonophylyoftheGrim- ＱＪ"Ｗ/Cs/e/j"minthePtychomitriaceaemaybe miales(slat.)basedonalargescaledatasetof derivedfiomthecharacteristicsofcalyptraandof 76cLsequences leaves,butthatitseemedtobeanunsatisfiactory Cavers（1910-1911）classifiedthemosseson oneNoguchi（1974）alsodiscussedthegenus thebasisoftheperistomeconfbnnation,dividing Raco〃かj"伽inrelationtothegeneraGr〃〃α theHaplolepideaeintofivesubdivisions，and andPltWho〃〃ｉｚ"?'，andsuggestedthatGrj"?〃α onlytheGrimmiaceaeandSeligeriaceaewereas- andRaco〃かj"mhaveacloserelationshipto signedtothefifthgroupPlatycranoideae、Our eachotherderivedfiomacommonancestoron studysupportsthis，andthesefactssuggestthat thebasisoftheirbroadandratherthincostae， haplolepidousperistomesoftheGrimmialesand consistingofalmostunifbnncells，ａｎｄｓｉｍｉｌar Seligerialesaresimilartoeachotherwithsome penstomesHeregardedRaco'"ｊかj"masanin- modiflcations・SincethemonophylyoftheGrim- tennediategenusbetweｅｎＰ〃cho加伽"ｍａｎｄ miales(slat.)waswellsupported,includingPlt)ノー Ｇ伽〃α・Deguchi（1977）showedsmallmale- c/to〃か/""０，０J'"p〕ﾉﾉosteﾉﾉ"mandmembersofthe branchesofP〃cho〃"ｊ"marisingffomthebase SeligerialeswithintheDicranidae，wepropose ofthevaginulainsidetheperichaetialleafcircle， herethatwecouldtreattheorderinawider andsuggestedthiscryptoicousfeaturetobeone sense，ｔｏｉｎｃｌｕｄｅｔｈｅＧ７〃〃α/Se/jge7jcJtype ofthedistinctcharactersinthegenus，Deguchi peristome． (1979)revisedGm伽α,ＳｂｈＭ伽"ｍａｎｄＣｍｃｊ‐ "o血〃ｉｎJapan，andincludedPltWho〃かｉ"腕in ２Ｊａｍilialandgenericrelationships theGrimmiaceae・Heshowedmolphologicalre- Gr〃〃αｃｅａｅα"‘Ｐひcho〃か/αceae lationshipsamongthegeneraintheGrim- Thefamilialcircumscriptionproposedbypre- miaceaQandsuggestedthecloserelationship vioussystemsdiffiersintheplacementofCm7Zpl〕ﾉｰ betweenRaco〃かj"ｍａｎｄＰｌｔ〕/ＣＡＯ〃〃/"肌espe- ﾉosreﾉﾉ川CDscj"o血"，Ｇ卯/to〃耐"腕ａｎｄＲａ－ ciallyinthecharacterofperistometeeth(asindi- ｃｏ〃伽"ｍ(seeTablel)．Someworkerstreated catedbyNoguchil974),calyptraeandsetae,and Plt)ﾉＣＡＯ〃がj""zandrelatedgeneraasthedistinct concludedthatG7加伽amaybethemostprimi- familyPtychomitriaceae、BruchandSchimper tivegenusinthefamilyChurchill(1981)madea (1837-1846)，Limpricht（1885-1889)，ａｎｄＮｙ－ cladisticanalysisfbrtheGrimmiaceae，andpro- ｈｏｌｍ（1954-1960）recognizedthePtychomitri- posedarevisionoftheclassificationofthefam- aceaewithinclusionofPlt〕ﾉc/to〃かｊｚＪｍａｎｄＧｂノー ily・Healsoshowedthatcharacters，suchasthe pAo〃かj"mhavingsharedcharactersofcalyptrae． dividedperistomeoutlineandpresencｅｏｆａｂａｓａｌ 6６ ＨｉｋｏｂｉａＶｏＬｌ４,No.1,2003 membrane,areapomolphicintheGrimmiaceac hocZgkj"somae,tohaveacloseaffinitywiththe lnthepresentstudy,Ｇ７〃〃α,比ｈ伽jcﾉｍｍａｎｄ RhachitheciaceaewithintheDicranidae，witha Ｃｂｓｃｊ"odb〃constimteamonophyleticgroup;and sisterrelationshiptothePottiales，fbnninga Raco〃/伽ｍａｎｄＰｌﾜﾉcho〃かj"〃(alsoQJ"VZ〕ﾉﾉo- weakmonophyleticclade・Ourtreeshowsthat sje""腕)paraphyleticallyappearedtothecladeof Glblp〃o加州加hasacloserelationtM'℃roawith Gr加伽α,Ｓｂｈｊｓｊｊ巾""zandCbscj"oc！O"・Thissug- highsupport（93/100/100）intheDicranidae geststhattheremarkablecharactersindicatedby clade，ａｎｄリノセ"〃rjeﾉﾉZJsj"e"sjsanddMJcqpi/"" Noguchi（1974)andDeguchi（1979)areplesio- hocZgAj"so"mefbrmadistinctsingleclade molphicintheGrimmiales（s」at.）andinclude (94/100/100)sistertothePottiaceaewithstrong thesmoothsurfaceofthecalyptra；３－４ormore support(96/100/95)．Theresultsdonotsupport cell-layeredcalyptra；setaetwistedtotheright； theinclusionofGblphomかj"川nthePtychomi- peristometeethlinear-lanceolateanddeeplydi- triaceae・ThesefactssuggestthatGblpho〃かi"m videdintotwofilifbnnbranches;andpresenceof bebestplacedintheDicranaceae(slat.[＝sensu abasalmembrane・hthepresentstudy,thePty‐ Vittl984DorRhabdoweisiaceae(sensuLaFarge chomitriaceaeappearaspoly-orpara-phyleticen- em/､2000),notintheEIpodiaceae,Ptychomitria- tities，althoughsomebrancheshavelowsupport ceaeorGrimmiaceae、 vaIues． Sbo"/ｅ７ｊａｃｅａｅ,Ｄｒ"加加o"c/Ｉａｃｅａｅα"cﾉﾘﾘﾉﾋJ'Qcﾉｍｃｅａｅ Se/jge7jaceaeα"dPlリノc/io〃rrjaceae Ｓｍ"/erjahasbeentraditionallyplacedinthe BruchandSchimper（1846）andSchimper Grimniales,althoughHooker（1837）suggested (1876）placedQJ"Ｗ/Cs/eﾉﾉ"mintheSeligeri‐ thesimilaritybetweenSＣＯ"/ｅｒｊａａｎｄﾘﾘﾉlzJ7c/、、 ａｃｅａｅｏｎｔｈｅｂａｓｉｓｏｆｉｔｓｓｍａｌｌｐｌａｎｔsize，peri- Edwards（1979）suggestedthattheproliferation stomecells，cellornamentationofthepenstome ofthecellpatternofthegenusismalkedlysimi- teeth,andsetacharactersMeanwhile,Limpricht lartotheMjgel･jatypepenstome，although (1885-1889),Brotherus(1901-1902;1924-1925)， Sbo"/erjahas32ilTegularteethwithrudimentaly andFleischer(1915-1923)showedtheaffinityof prostomiallayers，Churchill（1981）statedSboル Glblpﾉio〃か伽ａｎｄＱｍＷ/o伽/川．０J叩)ﾉﾉo- ノｅｒｊａｉｎｉｔｓｏｗｎｆａｍｉｌｙ,theScouleriaceaeChur- sje""ｍａｐｐｅａｒｅｄａｓｓｉｓｔｅｒｔｏｔｈｅＧ７加伽α－ chill（1985）placedSbo"/e7jawithintheHaplo- 此/i伽id/"m-CDscj"oc/Ｄ"二Raco〃かjzJmcladein lepideae，andtentativelyacceptedthehypothesis thｅＭＬｔｒｅｅａｎｄＰＰｃｏｎｓｅｎｓｕｓｔｒｅｅｗｉｔｈｌ００％ thatSbo"/ｅｒｊａｂｅｔｈｅｓｉｓｔｅｒｇｒｏｕｐｏｆｓｏｍｅｈａplo- support，althoughintheAUconsensustree，the lepidoustaxa，suchasthePottialesandGrim- positionofQJ"!p〕ﾉﾉosje/mmwasambiguous・Our miales;andrelatedittoahighertaxonomicleveL resultssuggestthatCtJ"１２〕ﾉﾉOsteﾉﾉ"misbestplaced Le・family，withintheHaplolepideaeexcluding intheGrimniaceae(s・lat.)． theDicranalesandFissidentales・Recently，La Gblpﾉiomか伽wastraditionallyclassedinthe Fargeem/､(2000)showedthebasalplacementof PtychomitriaceaeorGrimmiaceae（seeTablel） Sbo"んｒｉａａｎｄＤ'ｗｍｍｏ"`jaintheDicranidae・ Noguchi（1952),inhisrevisionoftheJapanese Goffinete/α/,（2001）alsodidnotsupportthe species,placedGll)lp/to〃かｊ"mintheEIpodiaceae closeaffinitybetweentheGrimmiaceae-- onthebasisofsimilarityinperistomedevelop- ceaeandtheDrummondiaceae-Scouleriaceae menttoPb"〃ｉｅﾉﾉﾋJ・Ｃｒｕｍ（1972a,ｂ)failedto basedonZps4sequences・BuckandGoffinet seethereasonfbrtransferringthegenusintothe (2000）placedtheScouleriaceaeandDrumnon- EIpodiaceae,andarguedthepositionofthegenus diaceaeintheGrimmiales・Inthepresentstudy， inthePtychomitriaceae、Edwards（1979）also Ｓｍ"/eljaappearedatthebasalpositionofthe concurredwithCrum'ssuggestionEst6baneze／ DicranidaefbrmingasinglecladewithD7zJm- α/､（2002）madehistochemicalobservationson "､"`jawithhighsupport(94/100/100)nearBr〕ノー theperistomeofseveralhaplolepidousmosses， oxjpﾊﾉ"腕，notsupportingplacementofthetwo andconfinnedthedistinctivefeaturessupporting familiesintheGrimniales、Themonogeneric thesegregationofGbip/iomrr川fi･omthePty‐ familyWardiaceaewasplacedintheSeligeriales chomitriaceae、LaFargee/α/,(2000)showedthe byBuckandGoffinet(2000)．Ｗ′､mjawastradi- EIpodiaceae，asrepresentedbyd"/αＣＯＰ""剛 tionallyplacedintheOrthotrichalesorLeucodon- HTsuBoTA,Ｙ､AGENo,BEsTEBANEz,Ｔ・YAMAGucHIANDH・DEGucHI 6７

talesAdditionalsequencesareneededtotest ＫＡＫＥＮＨＩ１１６４０６９８ａｎｄ６０１１７０１７ｔｏＨ・Degu- andresolvethephylogeneticpositionsofthe chi，10640683toTYamaguchi，ａｎｄl3740490 Scouleriaceae,DrumnondiaceaeandWardiaceae． andl5770058toHTsubota．

Conclusion Literaturecited

Ourstudycorroboratesthat(1)theGrimniaceae Adachi,』＆Hasegawa,Ｍ１９９６．ComputerScience (slat.[＝sensupresent])withtheinclusionofthe Monographs，Ｚ８ＭＯＬＰＨＹｖｅｒｓｉｏｎ２３、Programs generaG7加加α，Ｓｂｈな〃ｊ"肌Cbscj"oｄｏ"，Ｒａ－ fbrmolecularphylogeneticsbasedonmaximum ｃｏ〃〃j"腕，Ctm2ZZ〕ﾉﾉosteﾉﾉ"腕，Ｐｌリノcﾉto〃/伽mare likelihoodpp・’５０．InstimteofStatisticalMathe- monophyletic;(2)membersofgenerapreviously matics,Tokyo、 「 consideredasbelongingtothePtychomitriaceae Akiyama,Ｈ､,Tsubota,Ｈ､,Yamaguchi,Ｔ､＆Suleiman， andrepresentedbythegeneraPlt〕ﾉＣＡＯ〃/加川ａｎｄ Ｍ、2003．ThenewgenusBe"ｊｊｏｍ"ｊα(Daltonia- ＱＪ'１Ｗ/osZeﾉﾉ"''２，withtheexclusionofGﾉｳlpho〃‐ ceae,B1yopsida）fiPomMt・Kinabalu・Bryologist かj"、，didnotfbnnasingleclade，ａｎｄａｒｅｐａｒａ－ lO6:454-459． ｐｈｙｌｅｔｉｃｔｏｔｈｅＧ７加加a-Sb力加城""z-COscj"o- Arikawa，Ｔ＆Higuchi，Ｍ1999．Phylogenetic dm-Raco〃か伽clade;(3)Gblpﾉ）｡〃か伽showed analysisofthePlagiotheciaceae（Musci）andits acloseaffinitytodMoaintheDicranales,notto relativesbasedonr6cLgenesequencesCIypto- theGrimniales；（４）membersoftheSeligeria‐ gamie,BryoL20:231-245． ceae,representedbyMjgmaandBﾉﾉＭα,fbnna Arnott,Ｗ、１８２５．Dispositionm6thodiquedesespeces distinctcladesistertothecladeoftheGrim- demousses・Paris・ miaceae(slat.[＝sensupresent])includingmem‐ Brotherus,VFl901-1902Musci(Laubmoose）In bersofPlD′cho〃/伽manditsalliedgenera;（５） Engler，Ａ＆Prant１，Ｋ（eds.)，Dienamrlichen theGrimniales(s・lat.[＝sensupresent]),accom- PfIanzenfamilien，Ｅｄ・LWilhelmEngelmann， modatingtheGrimmiaceae(slat.)andtheSeli‐ Leipzig．［Ｍｊｇｅｌｊａ,β""伽（1901);Grimmiaceae genaceae，aremonophyleticwithoutmembersof （1902)］ theDrumnondiaceae，ｔｈｅScouleriaceaeorthe l924-192aMusci(Laubmoose）InEngler， Wardiaceae;ａｎｄ（６）thecladeconsistingofthe Ａ（ｅｄ),DienamrlichenPHanzenfamilien,Ｅｄ２， DrummondiaceaeandScouleriaceaeappearedat VoLlO-1LWilhelmEngelmann，Leipzig． thebasalpositionofthehaplolepidousmoss ［Seligeriaceae,Grimmiaceae（1924);Ptychomitria- groupDicranidae，nexttotheBryoxiphiaceac ceae(1925)］ Theprevioustaxonomicalcontroversyregarding Bruch,Ｐ．＆Schimper,Ｗ､Ｐ、1837-1846．Bryologia theGrinmialesisreHectedintheirphylogenetic Europaea・Smttgartiae．［Pl(〕′c〃o〃か川,Gblpho- relationships・ 〃かｊ"腕,Ｃｂｓｃｊ"oｄｏ"(1837);比〃Ｍｄ/"〃,Ｇ伽〃α， WithintheGrimmiales,additionaldataofmore Ｒａｃｏ〃/血沈（1845);Mjgerja,Ｂﾉﾉ"｡iα,Ｃの?Ｗ/o- rapidlyevolvedsequences，suchasmarKand sre/加加(1846)］ lTSorofuntreatedgenera(e,ｇＢｍｃ/ｉｙ血"〃ｍ Buck，Ｗ、Ｒ、＆Goffinet，B2000Morphologyand ａｎｄ乃汕"〃腕),wouldhelptoclarifytherela- classincationofmosses・ｈＳｈａｗ,ＡＪ.＆Goffinet， tionshipsamonggeneraorspecies・Furtherphy- Ｂ（eds.),BryophyteBiology,pp71-123・Cam- bridgeUniversityPress,Cambridgc logeneticinvestigationonrelationshipsamong theorderswithintheDicranidaeisalsoneeded Cavers，F1910-191LTheinter-relationshipsofthe bryophytesl-XLRepublishedasNewPhytologist Acknowledgements Reprintno4,ｐｐ、１－２０３，７２figs・Cambridge，１９１１． Churchill,Ｓ．Ｐ．１９８１．Aphylogeneticanalysisclassi‐ WewouldliketoacknowledgeProfR.ＤSep- Hcationandsynopsisoｆｔｈｅｇｅｎｅｒａｏｆｔｈｅ peltfbrredressingourEnglishandfbrtheircom- Ｇｒｉｍｍｉａｃｅａｅ(Musci)．ＩｎＦｕｎｋ,ｙＡ＆Brooks， mentsandsuggestionsonanearlierversionof 、Ｒ(eds.),AdvancesinCladisticsProceedingsof thispaper；Ａ，Tanaka，Ｎ・Nishimura，ａｎｄM theFirstMeetingoftheWilliHennigSociety,pp Itougafbrtheirgreathelpinprovidingplantma- l27-144NewYorkBotanicalGarden,ＮｅｗＹｏｒｋ teriaLResearchonmolecularsystematicsof １９８５．Thesystematicsandbiogeographyｏｆ mosseswasfimdedbythefbllowinggrants： 6８ HikobiaVoll4,No.1,2003

Ｓｍ此"αHook．（Musci：Scouleriaceae)．Lind‐ CircumscriptionandphylogenyoftheOrtho- ｂｅｒｇｉａｌｌ：S9-7L trichales(Bryopsida)inferredfTomr6cLsequence Cox,ＣＪ.，Goffinet,Ｂ,Newton,Ａ､Ｅ，Ｓｈａｗ,Ａ・』.＆ analyses・Ａｍ・ＪＢｏｔ,８５：１３２４－１３３７． Hedderson,Ｔ・ＡＪ・Z000Phylogeneticrelation- ，ＣＯＸ,ＣＪ.，Ｓｈａｗ,Ａ・I＆Hedderson,Ｔ､Ａ､Ｊ、 shipsamongthediplolepideous-alternatemosses Z00LTheBryophyta（)：systematicand （Bryidae）inferredfiomnuclearandchloroplast evolutionaryinfbrencesfi･ｏｍａｎノヴps4gene DNAsequencesBIyologistlO3:224-24L （cpDNA)phylogeny,AnnBot87:191-208． Crum，Ｈ・l972aAtaxonomicaccountofthe Grout,ＡＪ、1936．SeligeriaceachAJ.Ｇrout(ed)， ErpodiaceaaNovaHedwigia23:201-224. １９２８－１９４１，MossFloraofNorthAmerica,northof 1972bThedubiousoriginofGblpho〃/伽加 MexicoVoLLPublishedbytheauthor,Newfane． ｃα"α火"seMitt・IBryo1.7：165-168． ［Facsimileeditionl972,HafherPublishingCom‐ ＤｅＬｕｎａ,Ｅ，Buck,Ｗ､Ｒ､，Akiyama,Ｈ､，Arikawa,Ｔ， ｐany,NewYork］ Tsubota,Ｈ､,Gonzalez,，.,Newton,Ａ､Ｅ＆Ｓｈａｗ， Hascgawa,Ｍ,Kishino,Ｈ､＆Yano,Ｔｌ985．Dating ＡＪ・Z0000rdinalphylogenywithinthehypno- ofthehuman-apesplittingbyamolecularclockof

bryaleanpleurocalpousmossesinferredfiPom mitochondriaｌＤＮＡ．』・ＭｏＬＥｖｏＬ２２：160-174 ］ cladisticanalysesofthreechloroplastDNAse- ＆ 1989．Confidencelimitsonthe quencedatasets:jmL-EZps4and76cLBryologist maximum-likelihoodestimateofthehominoidtree lO3:242-256． fTommitochondrial-DNAsequencesEvolution43： Dcguchi,Ｈ１977．Smallmale-branchesofPlWAo〃かｊ"ｍ 672-677. （Grimmiaceae）arisedfiomthebaseofvaginula ＆１９９４．Accuraciesofthesimple insidetheperichaetialleafcircle・Misc・BryoL methodsfbrestimatingthebootstrapprobabilityof LichenoL7：177-179（InJapanesewithEnglish amaximumlikelihoodtree・ＭｏＬＢｉｏＬＥｖｏＬ１１： Figs.） l42-l4a 1979(1978）ArevisionofthegeneraGr加伽α， Hooker,Ｗ､Ｊ・’837．）Pizmm,anewgenusofmosses， ＳＭＭ伽"ＩａｎｄＣＯｓｃｊ"o血〃(Musci)ofJapanJ discovcredinSouthernAfTica・Companiontothe SciHiroshimaUniv.,ｓｅｒｂ,ｄｉｖ２，１６(2)：121- Bot・Ｍａｇ２：１８３－１８４＋ｐ１２５． Z56 Iwatsuki，Ｚ１９９１．CatalogofthemossesoｆＪａｐａｎ Dixon，Ｈ、Ｎ、1924．TheSmdent1sHandbookof Ｐｐ、１８２．HattoriBot・Ｌａｂ,Nichinan BritishMosses,Ｅｄ３、London［Facsimileedition ，Deguchi，Ｈ、＆Fumki，Ｔ２００ＬＮｉｈｏｎｎｏ l970,Wheldon＆Wesley,London］ yaseishokubutsu,Koke.[Mossesandliverwortsof Edwards,Ｓ､Ｒ・’979．Taxonomicimplicationsofcell JapanlPp３５５＋ｐＬ１９２，Heibonsha,Tokyo．（In patternsinhaplolepidousmossperistomesln Japanese） Clarke,ＯＣＳ＆Duckett,Ｊ､Ｏ(eds),Bryophyte Jones,ＯＮ1933．Grimmiaceae・ｈＡＪＧｒｏｕｔ(ed)， Systematics,pp317-346AcademicPress,London・ １９２８－１９４１，MossFloraofNorthAmerica,northof Est6banez,Ｂ,Tsubota,Ｈ､,Yamaguchi,Ｔ､＆Deguchi， MexicoVolZPublishedbytheauthor,Newfane． Ｈ、2002．Histochemicalobservationsonthe ［Facsimileeditionl972，HafherPublishing 「 peristomeofseveralhaplolepidousmosses Company,NewYork］ Hikobial3:667-677． Kishino,Ｈ､＆Hasegawa,Ｍ１989．Evaluationofthe Felsenstein，Jl98LEvolutionarytreesfiPomDNA maximumlikelihoodestimateoftheevolutionary sequences：amaximumlikelihoodapproachJ・ treetopologiesfiPomDNAsequencedata，ａｎdthe MoLEvol」７:368-376. branchingorderinHominoidea．Ｊ・ＭｏＬＥｖｏＬ２９： １９８５．Confidencelimitsonphylogenies：an 1７０－１７９． approachusingthebootstrap・Evolution39： ，Miyata,Ｔ､＆Hasegawa,M1990Maximum 783-79L likelihoodinferenceofproteinphylogeny，andthe Fitch，Ｗ・Ｍｌ９７ＬＴｏｗａｒｄｄｅｆＩｎｉｎｇｔｈｅｃｏｕｒｓｅｏｆ originofchloroplastsJ・Mol､ＥｖｏＬ３１：ｌ５１－ｌ６０ evolution：Minimumchangefbraspeciflctree ＬａＦａｒｇｅ，Ｃ､，Mishler,ＢＤ.，Wheeler,ＪＡ.，Wall，Ｄ、 topology・Syst・Zoo1.20:406-416． Ｐ.，Johannes,Ｋ､，Schaffbr,Ｓ＆Ｓｈａｗ,Ａ､Ｊ、２０００ Fleischer，Ｍ１915-1923．DieMusciderFloravon Phylogeneticrelationshipswithinthehaplo- BuitenzorglＶ・Leiden lepideousmosses・ＢryologistlO3:257-276. Ooffinet，Ｂ，Bayer，ＢＪ．＆Vitt，，．Ｈ・’９９８． ,Ｓｈａｗ，Ａ・Ｉ＆Vitt，，．Ｈ、2002．Ｔｈｅ HTsuBoTA,Ｙ・ＡGENｏ,BEsTEBANEz,TYAMAGucHIANDH・DEGucHI 6９

circumscriptionoftheDicranacｃａｅ（Bryopsida） logenetictreeselectionSyst､ＢｉｏＬ５１：492-508. basedonthechloroplastregionsか"Ｌ－か"Ｆａｎｄ ＆Hasegawa,Ｍ２００ＬＣＯＮＳＥＬ:fbrasses- Ｚｐｓ４Ｓｙｓｔ・Bot､２７:435-452． singtheconfidenceofphylogenetictreeselection Limpricht，Ｋ、Ｇ、1885-1889．DieLaubmoosQIn Bioinfbnnaticsl7：1246-1247． RabenhorstisKryptogamen-Flora,４vols、Leipzig Sikes,，．Ｓ、＆Lewis,Ｐ､０．２００ＬＰＡＵＰＲａｔ:ＰＡＵＰ＊ Magombo，ＺＬＫ、2003．Thcphylogenyofbasal implementationoftheparsimonyratchet,betasoft- penstomatemosses：evidencefiomcpDNA，and ware,versionLDepartmentofEcologyandEvolu- implicationfbrperistomeevolution・Syst､Bot28： tionalyBiology,UniversityofConnecticut,Storrs、 ２４卜38． Swoffiord,Ｄ､ＬＺ００ＺＰＡＵＰ*PhylogeneticAnalysis Murray，ＭＧ．＆Thompson，Ｗ、Ｆ１９８０．Rapid UsingParsimony(*andOtherMethods）Version4 isolationofhighmolecularweightplantDNA、 SinauerAssociates,Sunderland,Massachusetts ５ Nuc・ＡｃｉｄｓＲｅｓｌ９:4321-4325． Tsubota,Ｈ､，Nakao，Ｎ､，Arikawa,Ｔ，Yamaguchi，Ｔ， Nixon，Ｋ・Cl999TheParsimonyRatchet，anew Higuchi,Ｍ,Deguchi,Ｈ､＆Seki,11999．Apre-

／ methodfbrrapidparsimonyanalysisCladistics liminaryphylogenyofHypnales(Musci)asinfelTed l5:407-414 ffomchloroplastr6cLsequencedataBryoLRes、 Noguchi,Ａ、１９５２．MusciJaponici・nEIpodiaceae．』・ ７:233-248. HattoriBotLab8:５－１７． ,Yamaguchi，Ｔ，Seki，Ｉ＆Deguchi， 1954．MusciJaponici．Ⅳ、Ｔｈｅｇｅｎｕｓ Ｈ・Z000Preliminaryphylogeneticrelationshipsof ＰｌﾘﾉCAO〃かｊ"ｍＪ・HattoriBot,Lab､１２:1-26. thegenus〃ｏｒA”eﾉﾉaanditsalliedgenera(Hyp- 1974．MusciJaponiciX・Ｔｈｅｇｅｎｕｓ nales,Musci)basedonchloroplast76cLsequence Ｒａｃｏ〃かｊ"ｍＪ・HattoriBot・Ｌａｈ３８:337-369. data．Ｊ・HattoriBotLab､８８:79-99. ,Iwatsuki，Ｚ＆Yamaguchi，Ｔ・’987-1988． －，Akiyama，Ｈ､，YamagUchi，Ｔ＆Deguchi，Ｈ、 IllustratedmossfloraofJapan，Ｐａｒｔｓｌ＆２．，ｐｐ 2001aMolecularphylogenyoftheSematophyl- ｌ－２４２＆Z43-49LHattoriBot・Ｌａｂ,Nichinan laceae(Hypnales,Musci)basedonchloroplastrbcL Nyholm,Ｅ1954-1960111ustratedMossFloraof sequences・JHattoriBotLah90:２２１－２４０．

Fennoscandia・IIMusciFasc､１，２，４．Ｌｕｎｄ l ＆２００１bMolecular ［Seligeriaceae(1954);Grimmiaceae(1956);Ptycho‐ phylogenyofthegenus刀jMoegjs/jaandrelated mitriaceae(l960H genera(SematophyllaceaeMusci)basedonchloro- Oguri,Ｅ，Yamaguchi,Ｔ，Tsubota,Ｈ、＆Deguchi，Ｈ、 plastr6cLsequencesHikobial3:529-549. 2003．Apreliminaryphylogenicstudyofthegenus ，Arikawa，Ｔ，Akiyama，Ｈ､，ＤｅＬｕｎａ，Ｅ， Le"co61W畑（,Musci）ｉｎＡｓｉａａｎｄ Gonzalez，，.，Higuchi，Ｍ＆Deguchi，Ｈ、2002． ｔｈｅＰａｃｉｆｉｃｂａｓｅｄｏｎＩＴＳａnd76cLsequences Molecularphylogenyofhypnobryaleanmossesas Hikobial4:45-53. inferredffomalarge-scaledatasetofchloroplast 01sen,ＧＪ.，Matsuda,Ｈ､，Hagstrom,Ｒ､＆Overbeek， 76cL,withspecialreferencetotheHypnaceaeand Ｒ・ｌ９９４ｆｉａｓｔＤＮＡｍｌ：Atoolfbrconstructionof possiblyrelatedfamilies・Hikobial3:645-665． phylogenetictreesofDNAsequencesusingmaxi- Vitt，，．Ｈ・ｌ９８２ＳｐｈａgnopsidaandBryopsida・In mumlikelihoodComput・ApplBiosci・１０:41-48． Parker,ＳＰ.（ed)，Synopsisandclassificationof Saitou,Ｎ＆Ｎei,Ｍ、1987．Theneighborjoining livingOrganisms，ＶｏＬｌ，pp307-336・McGraw-

、 method：anewmethodfbrreconstructingphylo- Hill,NewYork． genetictrees・ＭｏｌＢｉｏＬＥｖｏＬ４:406-425． －，１９８４．ClassificationoftheBryopsidaln Schimper,ＷＰ・’876．SynopsisMuscommeuropaeolum Schuster,ＲＭ.（ed),NewManualofBryology， praemissaintroductionedeelementisbryologicis Vol､２，pp696-759・HattoriBot・Ｌａｂ,Nichinan tractante,Ｅｄ,２．ｐｐ８８６・ESchweizerbard,Stuttgart・ ，Goffinet，Ｂ、＆Hedderson，Ｔ、1998．Ｔｈｅ Schwarz，Ｇ、１９７８．Estimatingthedimensionofa ordinalclassificationofthemosses・Questionsand modeLAnn.Ｓtat､６:461-464． answersfbrthel990s・InBates,１Ｗ.，Aston,Ｎ、 Shimodaira，Ｈ、2000．Anothercalculationofthe Ｗ＆Duckett，ＪＧ（eds)，Bryologyfbrthe p-valuefbrtheproblemofregionsusingthescaled Twenty-firstCentury,ppll3-123・Maneyandthe bootstrapresamplings・TechRep・Ｎｏ．2000-35． BritishBryologicalSociety,Leeds． StanfbrdUniv.,PaloAlto,Califbrnia． 2002.Ａ、approximatelyunbiasedtestofphy－ 7０ HikobiaVoLl4,No.1,2003

AppendixA・hfbrmationonspecimensffomwhich alt.,Ｊｕｎｅ24,1998,HTsubota2326（HIRO);Ｓｂｈｉｓ－ DNAwasextractedinthissmdy・ ﾉﾉ伽"加伽α"〃（Turner）LoeskeexMartemsson， Hereisalistoftaxawhose'七cLscquenceswereuti- AB125583,Japan,Honshu,Hiroshima-ken,Yamagata- lizedinthissmdywiththeiraccessionnumbers,show- gun，Geihoku-cho，YahataHighlands，ｃａＢＯＯｍａｌｔ.， ingtheirsourcesandvoucherspecimeninfbnnation July16,1996,HDeguchi33775(HIRO);Seligeriaceae VoucherspecimensarekeptinHRo・ Mjge7mα"sかｊａｃａＴ・Schauer，AB125589，Japan， Thesequenceofinfblmationis：ｔｈｅｎａｍｅｏｆｔａｘｏｎ， Honshu，Nagano-ken，Ina-gun，Ooshika-mura，ca accessionnumber，localityandspecimennumber，or L570malt,August21,2000,HTsubota3842(HIRO)； referencesource・Grimmiaceaeanditsallies：QJ77qp〕ﾉｰ Bﾉﾉ"ぬαノ叩o"jcaBroth.，AB125590，Japan,Honshu， ﾉosreﾉﾉ"腕scmco/ａ（ＦＷｅｂｅｒ＆DMohr）Bruch＆ Hiroshima-ken,Yamagata-gun,Togouchi-cho,Sandan- Schimp.，AB125584，Japan，Honshu，Tottori-ken，Mt kyo；510-600ｍａｌｔ.，August２３，１９９６，Ｈ・Deguchi ザ Daisen,ｃａ９５０ｍａｌｔ.,August28,2000,Ｔ､Yamaguchi 34189(HⅨo);EIpodiaceae凡"/帆eﾉﾉﾋＩＳｉ"e"sな(Ven- l8851（HIRO);Cbsci"o血刀c7j6ros"ｓ(Hedw.）Spruce， mri)MiillHaL,ABl25591,Japan,Honshu,Hiroshima- AB125575，Japan,Kyushu,Kagoshima-ken，Yakushi- ken，Aki-gu､，Kaita-cho，Ｃａ、Omalt.，February25， maIsL，ｃａｌ,800ｍａｌｔ.，Ｍａｙ２９，２０００，Ｈ・Deguchi 1999,HTsubota2580(HIRO);PottiaceaePo"ｊａｊ"花アー 35024（HIRO)；Ｇ伽伽ａｑｐｉｃ"〃αHornsch， ｍｅｄｉａ（Turner）Fijmr.，AB125592，Japan，Honshu， AB125576，Japan，Honshu，Nagano-ken，MtShiro- Hiroshima-ken，Aki-gun，Shimokamagari-cho，April uma,Ｃａ､2,600ｍａｌｔ.,August12,2000,Ｙ,Ａｇｅｎｏ２８ 23，2000,HTsubota3850（H凪o)，Leucodontaceae (HⅢo)；Ｇ伽〃αｅ/o"gzzmKaulf,AB125577,Japan， DoZ〕ﾉαノ叩o"jcaSandeLac.，AB125593，Japan，Ｋｙ‐ Honshu,Nagano-ken,MtShirouma,Ｃａ､2,600ｍａｌｔ.， ushu,Kagoshima-ken,YakushimaIsL，1,380-1,600m August12,2000,ＹＡｇｅｎｏ２９(H皿o);Ｇ伽伽ａｏｌｂｊ－ alt.,October19,2000,HTsubota4019(HⅢo） ｃ"/α伽Bruch,AB125578，Spain,Ｍadrid,ArgandaIel Rey,Dehesa,ElCalrascaL660malt.,December25, AppendixBIntemetresourcesfbrthephylogenetic 1999,Ｙ・Ageno57(H皿o);Ｇ伽伽apj/舵mPBeauM， analysisanddatabaSc． AB125579,Japan,Honshu,Hiroshima-ken,Saeki-gun， Sﾋﾂﾞiwa7e Miyajimals1.,Ｍt・Misen,ｃａｊＯＯｍａｌｔ.,Apri］13, CONSEL＜http://www・islnacjp/~shimo/prog/consel／ 2000,Ｙ､Agenol9(HIRO);Ｇ伽〃α〃/W"α、(Hedw.） indexｈｔｍｌ＞ Ｓｍ.,AB125580,Spain,Madrid,ArgandalelRey,De- MOLPHY＜fttp://ftpjsmacjp/Pub/ISMLIB／ hesa,ElCarrascaL660malt.,December25,1999,Y・ ＭＯＬＰＨＹ/＞ Ageno56(Hmo);GbipAo加加加〃"〃ﾉﾉ〃"腕(Mitt.） PAUP*