SematOphyllaceae,Musci

Hikobial3：529-549.200］
MolecularphylOgenyofthegeⅢｓＢＱｊＭＤ照応〃andrelatedgenera
(SematOphyllaceae,Musci)basedOnchlOroplastmcLsequencesI
HIRoMITsuBoTA,HIRoYuKIAKIYAMA,ＴＯＭＩoYAMAGucHIANDHIRoNoRIDEGucHI
TsuBoTA，Ｈ､，AKIYAMA，Ｈ､，YAMAGucHI，Ｔ＆DEGucHI，Ｈ、２００LMolecular
phylogenyofthegenus乃伽egjsがaandrelatedgenera(Sematophyllaceae,Musci）
basedonchloroplastr6cLsequences・Hikobial3:529-549．
PhylogeneticrelationshipswithintheSematophyllaceaehavebeenthefbcusof
muchattentioninrecentyearsHerewepresentphylogeneticinferencesfbrtheSematophyllaceaeandpossiblyrelatedtaxabyinvestigatingthephylogeneticpositionof
thegeneraか伽ｅｇＭａ,Ｈｅ/emp/りﾉﾉﾉﾉ""',MmFrOpo"、and、｢i【ﾉﾉﾋね,andbytestingthe
monophylyofeachgenus,basedon63r6cLsequences・Theresultsofthepresent
smdysuggestthat(1)theSematophyllaceaewithawidesenseproposedbyTSubota
andcoworkers(2000,2001）areresolvedasamonophyleticgroup;(2)theWijki‐
oideaebyHedeniisandBuck（1999）arepolyphyletiqand（３）thegenera
刀ＭｅｇＭａ,MJsmpomaand〃kjaarepolyphyleticTheseresultsconfinnthe
suggestionproposedbyTsubotaetaL(2001）thatthecladesoftwolineagesinthe
Sematophyllaceaeareindependenｔｏｆｅａｃｈｏｔｈｅｒ，andshowtheneedfbrof
taxonomicalreviewofthepolyphyleticgenera，ｓｕｃｈａｓ刀jFmegな"α，ｊＭｍｑｐｏｍａ
ａｎｄ〃kjqWealsorecognizetheHeterophyllioideaeasaparaphyleticgroupinthe
Sematophyllaceae
H7m伽乃"6川７b〃ｏ】Zwmg"chj＆Ｈｉｍ"o'ｊＤｅｇ"c/iiPGpa7me"/q/Ｂｊｏ/ogjcaノ
Sbje"Ｃｅ,Ｇ７ａＣｍα花川､o/ｑ/此ｊｅ"Ｃｅ,ＨｉｍＳﾉﾘ〃α肋jveハノ軌hJgmm〕ﾉα〃α／-3-1,
HigasﾊﾉｰｶﾉﾉＤｓｈ〃α-sﾙﾉ,Ｈｉｍｓｈ〃α734852．上pα"・
H1グロ〕′"ﾉｔＭ町α"α,Ｍ｛se"、q/ＭＪ/"だα"cﾉＨ２/ｍａＭｃ伽伽,Ｂｉﾉ090,ItI〕ﾉojgZJoﾉhJd
Sb"〃sﾙﾉ,Ｂｉ′ogo664ﾉｺﾞｲｄＪｑｐα〃
lntroduclion
TheSematophyllaceaeareamongthemostdiversefamiliesofpleurocarpousmosses，com-
posedofca50genera(Ⅵttl984).Thefamilyis
widespreadintheAsianpaleotropicsandthe
neotropics,intrudingintosometemperateregions，
andoftenocc叩yingepiphytichabitats・CircumscriptionoftheSematophyllaceaehasbeenvariouslydiscussed,particularlyconcerningitsdistinctionfiomtheHypnaceae.Ⅵtt（1984)providedanewsystemfbrclassificationofthepleurocaIpousBryopsida・HetreatedtheSematophylla-
ceaeinawidesense,placingthefamilycloseto
theEntodontaceaeonthebasisofmorphological
andecologicalpreferenceasepiphytes．
lContributionffomtheLaboratoryofPlantTnxonomy
andEcology,HiroshimaUniversity.Ｎ・Ser・No.５１２．
SubdivisionoftheSematophyllaceaehasalso
beenproposedthroughtherecognitionofsub
familiesbyseveralauthors・Ｔｈｅsubfamilialcircumscriptionsproposeddifferffomauthortoau-
thorinthetreatmentsofthegenera乃応magな"α，
MIDmpo川〃肋,ＢＭｈｅ"ﾉﾉα,町加ｊｑ火Ilpha，
andHb花、p/iﾘﾉﾉﾉj"",Fleischer(1915-1922)rec‐
ognizedtheSematophyllaceaewithｆｂｕｒ“subfamilies，，：theClastobryeae,theHeterophyllieae，
theSematophylleaeandtheMacrohymenieae・
TheFleischersystemhasbeenwidelyadopted，
althoughsomedisputeshaveoccurredonthe
treatmentandcircumscriptionamongthesubfamilies(e､９，Tixierl977;、、＆Buckl989)．
Brotherus(1924-1925)fbllowedFleischer，ssystemandalsorecognizedfbursubfiamilies：the
Clastobryoideae，Heterophyllioideae（"Hetero-
phylloideae，，；cfRobinson＆Ｒｅｅｄｌ９６６,Ｔａｎ＆
Buckl989)，Sematophylloideae，andMacro‐
530
ＨｉｋｏｂｉａＶｂＬ１３,No.３，２００１
hymenioideaelnbothsystems,ｔｈｅｇｅｎｅｒａＧａｍ－
〃eﾉﾉﾋlMplt)ﾉc力ＱＰＳな,Ｈｂに､p/1〕ﾉﾉﾉ剛,Ｍ】,､PC川
ACα"ﾉﾉzoc/αｄｉ"碗（＝〃んｊα）ａｎｄが柳egMa
wereplacedintheHeterophyUioideaeandAcaﾙ
ﾉﾙo'伽"cA伽intheSematophylloideae・Ｔｔｍ
ａｎｄＢｕｃｋ（1989)treatedthefamilyinawider
senseinattemptinganewgroupingofthesematophyllaceousgenera・TheyrecognizedfbursubfamiliesintheSematophyllaceae：Clastobryoi‐
deae，Heterophyllioideae，Sematophylloideae
andMacrohymenioideae，excludingSj7"cAja
ffomtheSematophyllaceae,andexcluded幼ＩＤノー
Ｃｈｅ"αffomtheClastobryoideaeandplaceditin
theHeterophyllioideae、HedeniisandBuck
(1999）analyzedmorphologicalandanatomical
charactersfbrtheSematophyllaceaeandtheirallies，andproposedanewclassificationofthe
familyTheysuggestedthatthefamilyislikelyto
bemonophyleticexcludingBMAe”ﾉﾉcJandits
possiblyrelatedgenera,suchas町伽mdbIpha，
AOC/とJﾙﾉﾉZJandHbjemp/Ｍﾉﾉ川andrecognized
twosubfamiliesinthefamily：SematophylloideaeandWijkioideae・Theyclassifiedthegenera,〃肋,〃MegMaanMcα"伽７町"ｃｈｊ"m
asmembersoftheWijkioideae,showingthedifL
ferencesbetweenmembersoftheSematophyllaceae(sensuHedeniis＆Buckl999)andmembersofBm的e”ﾉﾉﾋJandalliedgenera・
PhylogeneticrelationshipswithintheSematophyllaceaeaswellasbetweentheHypnaceae
andSematophyllaceaehavebeenthefbcusof
muchattentioninrecentyears(Hedeniisl996；
TEubotaetaL1999；Arikawa＆Higuchil999；
ＤｅＬｕｎａｅｔａＬ１９９９;Hedeniis＆Buckl999;Ｄｅ
ＬｕｎａｅｔａＬ２０００;TSubotaetaL2000;BucketaL
2000;TBubotaetaL2001)nubotaetal.(2000）
madeaphylogeneticstudyonthefamilywith
particularattentiontoB'Ｄ肋e”ﾉﾉZJwhichhasever
beentreatedasamemberoftheHypnaceaeby
Seki（1968)andHedeniisandBuck(1999).TFubotaetal.(2001）showedthegenera〃Ａｊａａｎｄ
Ｈｂ/emp/1y"伽aspolyphyleticentitiesbasedon
molecularanalyses・Tsubotaandcoworkers
(2000,2001)showedonlyasinglecladeaccom‐
modatingtheSematophyllaceaeinawidersense，
inwhichBMhelqeﾉﾉﾋJ,〃ﾉﾋﾉα,Ｈｂｔｅｍｐ/1〕ﾉﾉﾉ川and
theiralliedgenera,andsomeotherspeciespreviouslytreatedasmembersoftheHypnaceaeare
alsoincludedBothsmdiesalsorecognizedno
cladescorrespondingtothesubftlmiliesproposed
byanypreviousauthorities・
かなｍｅｇｊＳｊｊａｉｓｏｎｅｏｆｔｈｅｇｅｎｅｒａintheSematophyllaceae，andismainlydistributedinthe
mountainousareasofthepaleotropicregionof
AsiaThisgenusischaracterizedbyrobustplant
size,stipes,longseta,well-developedleafborder
withteeth,andlargeandrectangularthin-walled
alarcells，togetherwithahabitatpreferenceｆｂｒ
ｇｒｏｗｉｎｇｏｎｓｏｉｌｏｒｒｏｔｔｅｎｌｏｇｓ，occasionallyon
treetrunks・Over50specieshavebeendescribed
fbrthegenus，ａｎｄｓｏｍｅｏｆｔｈｅｍｈａｖｅｂｅｅｎ
ｍｏｖｅｄｔｏｏｔhergenera，ｓｕｃｈａｓＲ/，叩/jjc/b７－
'hIy"c伽加,』cα"rho山伽加（＝〃Ajα),ＭｍＷ‐
ｐｏｍａ,Ｓｂ腕ajQp/iﾘﾉﾉﾉﾂﾞｍｏｒＨｂ/e叩/iﾘﾉ"ｊ""（Ｗｉｊｋ
ｅｔａＬ１９６９).Thephylogeneticpositionofthe
genusDなmegjlma，oneofkeygeneraonthe
subfamilialtreatmentintheSematophyllaceae，
wasalsosmdied,andthegeneraDなmeg応がαａｎｄ
〃kjadidnotfbnnasingleclade(TEubotaetaL
2001).However,therelationshipsbetweenthe
genusandMJsmpollza,Ｈｂ死''０p/iﾘﾉﾉﾉ卿ＯＭＣα"‐
ｊｍ７Ａ〕ﾉ"ｃｈｊｚ"，zarenotsufficientlydiscussedin
theirwork,sowefbcusherethephylogeneticpositionof乃応腕agiWjq
Theaimsofthissmdyare:(1)todetenninethe
phylogeneticposiｔｉｏｎｏｆＤｊＭ７ｅｇ伽ｍａｎｄｔｏｔｅｓｔ
ｔｈｅmonophylyofitandrelatedgenera;（２）to
constructaphylogenetictreefbrtheSematophyllaceae,verifyingthevalidilyofthesubfamilies
HeterophyllioideaeandWijkioideae;ａｎｄ(3)to
investigatethephylogeneticpositionsofかjw71egMa,MJurOpo川肺Aja,Ｈｅ/empﾉＭ/伽and
dcα"ﾉﾙｏｒ〃"c力伽withintheSematophyllaceae
withaparticularinterestontherelationship
betweenthefamilyandtheHypnaceae
TheSematophyllaceaeasdefinedbyTBubota
andcoworkers（2000,2001）areindicatedby
"(sensuTBubotaetaL).”Thedefinitionwithout
theword“sensu，，meansthetraditionalconceptualsenseortheSematophyllaceaesensulato，in-
cludingsomespeciesandgenerawhosepositions
havebeendiscussｅｄｗｉｔｈｔｈｅｆａｍｉｌｙＷｅｕｓｅｔｗｏ
ｔｅｒｍｓｉｎthephylogeneticdescription:cladeand
-lineageTheword“clade”meansthemonophyleticgroup，whichincludesamostrecentcommo、ａｎcestorandonlyallitsdescendants・Ｔｈｅ
ｗｏｒｄ“-lineage，，meansamonophyleticorparaphyleticentity，whichexcludesoneormorecomponentsofthemonophyleticgroup．
Ｈ､TsuBoTA,Ｈ・AKIYAMA,ＴＹＡＭＡＧｕｃＨＩＡＮＤＨ・DEGucHI
Materialsandmethods
Aphylogeneticstudywascarriedouttoinvestigatethephylogeneticpositionof刀jMzegjMa
anditsalliedgenerainferredffomther6cLgene
data・Thisstudyconsistsoftwosteps:（１）obtainingsequencedata(DNAextraction,ＰＣＲａｍ‐
plificationandDNAsequencing),ａｎｄ（２）data
analyses．
SDZ"℃ｅｓα"c/ｐ/α"'f〃、〃jaLs
Atotalof63mossspecieswereexaminedfbr
theanalysesasshowniｎＴａｂｌｅｌａｎｄｉｎＡｐｐｅｎ－
ｄｉｘＡ，Thelattercontainsvoucherinfbnnation
andDDBJAccessionNos・Thevoucherspecimenssequencedbythepresentauthorswerede‐
positedinHYoorHIRo・Somesequencesusedfbr
theanalyseswerealsoobtainedfi･omtheDNA
database．
D/ＶＨｅｘかαc"o",ＰＣＲα"2ﾉﾜﾉ抗cα/jo〃α"Ｃｌ/Ｄ/V4
se9"e"cmg
TbtalDNAwasextractedfi･omfTeshmaterials
(50-500mgffeshweight）ordriedspecimens
(20-300mgdryweight)bymodiHcationsofthe
CTABmethodAmpliHcationofr6cLgenesegmentsfbrdirectsequencingwasdonebypolymerasechainreaction(PCR)withTaqpolymer‐
asｅ(TaKaRa)inastandardconditionfbllowing
耐ubotaetal（1999)Reactionswereperfbnned
usingaDNAthennalcycler（ThennalCycler
9600,PerkinElmer)ｗｉｔｈＰＣＲａｍｐＭｃａｔｉｏｎｋｉｔ
(PerkinElmer)andsyntheticprimers（16-33nucleotidesinlength;Table２).Afterampliflcation
productswerecheckedandpuriHedonconcentratingfilters（TaKaRa)．DirectDNAsequence
analysesofthePCRproductswereperfbnnedby
dideoxychainteminationmethodaccordingto
themanuftlcture,sinstructionsfbllowingTbubota
etal.（1999）Thereactionswereperfbnnedin
thethennalcyclerwiththesameprimersfbrthe
PCRamplifIcations・Thesequenceswereelectrophoresedonanautomatedsequencer(ABI373A
andABIPmSM310,PerkinElmer),andanalyzedonthedata-analysissystemDetailsand
filrtherinfbnnationfbrDNAextractingprotocols，PCRprotocolsandthoseofsequencingreactionsaredescribedpreviously（Tsubotaand
coworkersl999,２０００)．
5３１
４Ｍ〕ﾉsjsc/ｂｊａ
Ｄａｍ３ｅノーThetaxatreatedintbissmdyare
shoｗｎｉｎＴａｂｌｅｌｗｉｔｈＤＤＢＪａｃｃｅｓｓｉｏｎｎｕｍbers、
Fortheanalysis,ｗｅｕｓｅｄｔｗｏｄａｔａｓｅｔｓ:ｄａｔａｓｅｔｓ
ｌａｎｄｌＬＤａｔａｓｅｔｌｉｎｃｌｕｄｅs63sequencesfior
hypnaleanmosseｓａｎｄｏｔｈｅｒｓａｓｏｕｔｇｒｏｕｐｓｔｏ
ｍａｋｅａｓｕrveyfbrthemonophylyoftheSematophyllaceacDatasetllincludes42sequencesfbr
thesematophyllaceousmossestomakeadetailed
analysisonthephylogeneticpositionof刀"M7egjs血andsomegeneratreatedintheHeterophyllioideaebyFleischer(1915-1922)andBrotherus
(1924-1925)andinWUkioideaebyHedenasand
Buck（1999).Thesedatasetsobtainedfiomour
originalsequencesandtheregisteredsequences
fbrthedatabasewereanalyzedTbedatasetsfbr
analysesinclude5speciesofDjsmegjsrjaand
another28speciesintheSematophyllaceaeand
theHypnaceae．
」ﾉﾉg"ｍｅ"ノーManualalignmentwascarriedout
fbrthedatasetagainstthoseofR/りﾉ"chos/eg/""７
pα/Ｍ/b"""?ａｎｄＢＭ/'e花ﾉﾉαｈｅ"o"jjwithaneditorprogramThesequenceswerereversedusing
ouroriginalprogram，Sclean253（byTsubota
l998-2000，ｉｎｏｕｒｈｔｔｐｓｉｔｅ，seeAppendixB)．
ＴｈｅotheroriginalprogramswritteninＰｅｒｌａｎｄＣ
ｗｅｒｅａｌｓｏａｐｐｌｉｅｄｆｂｒｔｈｅｄataprocessingThe
undeterminedsiteswereremovedfiDmtbealignmentbefbrethesubsequentanalyses．
４Ｍ〕入SjsmeﾉﾉjoQb
PhylogeneticanalyseswereperfbrmedfbllowingTsubotaandcoworkers（1999,2000,2001)．
Treeswereconstructedbythefbllowingfbur
metbods:neighbor-joining(NJ)method（Saitou
＆Neil987),minimum-evolution(ＭＥ)method
(Rzhetsky＆Neil992，1993),maximum-parsimony(ＭＰ)method(Fitchl971)ａｎｄｍａｘｉｍｕｍ‐
likelihood(ＭＬ)method(Felsensteinl981）The
treesobtainedbythefburmethodswereappraisedbythelog-likelihoodmeasurc
D伽α"ｃｅａＭ〕′sな－NJandMEtreeswereconstructedbｙＭＥＧＡｖｅｒｓｉｏｎ２０(MEGA2;Kumar
etaL2001，seeAppendixB）usingａＫｉｍｕｒａｓ
(1980）two-parametermodelfiorthedistance
estimations．
Ｐα祁伽o〃α"αl【Msmjs-MPtreeswereconstructed
bｙＭＥＧＡ２ａｎｄＰＡＵＰ＊４０b８（Swoffbrdl998)．
ＩｎａｎａｌｙｓｅｓｗｉｔｈＭＥＧＡ２，MPtreeswere
searchedbyClose-neighbor-interchange（ＣＭ）
532
ＨｉｋｏｂｉａＶｂＬｌ３,No.３，２００１
THbleLListofspeciesandfamiliesinvestigatedfbr76cLgenewithaccessionnumbersofthesequence・
MoreinfbnnationisshowninAppendｉｘＡ．
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ABO39785
Ｂｍﾉﾉie”ﾉﾉﾋJ力e"o"ｊｊ
助/odb〃r"ｂｊｃ""dｶｨｓ
Hypnaceae
ＡＢＯ５１２２５
ＡＢO51217
１１
１１
Entodontaceae
ＡＦ２３３５７２
DataSet
１１
ＡＢＯ５１２２４
ＩＩＩ１１ＩＩＩＩＩＩ１１１１ＩＩＩＩＩＩＩＩ１１ＩＩＩＩ１１１１１１１１
１１１１１１１１１１ＩＩＩＩ１１ＩＩＩＩＩＩＩＩ１１ＩＩ１１ＩＩ１１１１Ｉ
Accessionnumber
ＡＣα"ﾉﾙo〃ｿﾞ!〕′"ｃｈｊ"腕ｐｑｐﾉﾉﾉcJ'"m
Acmporj冴加ｐ"埴e"ｓ
ｄｃｍｐｏ７ｊ""１ｓかａｍｊ"e"、
〃'(〕′Ｃｈｅ〃gﾉb〃eｍ'qpmpagzｲﾉ従ｍ
Ｂｍｊ力e'qeノルＣＯ"ｌｐｍ"α、
８ｍ/ルビ”ﾉﾙ!/b1"ｒｊｅｊ
＆＆＆＆＆＆＆＆＆＆Ｉ＆＆＆＆＆＆＆＆＆＆＆＆＆＆＆＆＆＆＆＆＆＆＆＆＆＆＆ＩＩＩ＆＆１１１＆＆＆ＩＩＩＩｌＩＩＩｌＩＩ
species
Sematophyllaceae
Ｉ１１ＩＩ１１１１ＩＩ１１１１１１ＩＩ１１ＩＩＩＩｌＩ１１１１１１１１１１
Family
Ｈ・TsuBoTA,Ｈ,AKIYAMA,ＴＹＡＭＡＧｕｃＨＩＡＮＤＨ・ＤＥＧｕｃＨＩ
5３３
Ttlble2Sequences(5Ｌ３')ofsyntheticprimersusedfbrPCRamplificationandsequencingofther6cL
geneTheprimernumbersfiFom5'endcorrespondtoroughpositionsinthesequenceofMJ'℃ｈα"ｒｊａｐｏｂノー
moZphα(OhyamaetaL1986)．
ｒ
ｍ
ｅ
Ｓ
・ｎ３
．ｍ渥匡慮凪屋憾灰斌Ⅲ３３
ｐ９００４７９ｏ／４，４ＲＲ服
４
ＧＡＴＣＴＡＣＴＡＣ
ＴＴＴＡＡＴＴＣＣＴ
ＧＴ
ＡＧＴＴＧＡＡＴＣＴ
ＡＡＡＧＧＡＴＣＴＡ
Ｃ
ＴＧＴＴＧＡＡＣＴＴ
ＣＡＣＡＡＧＴＡＡＣ
Ａ
ＡＣＡＡＴＧＡＴＡＣ
ＴＧＴＴＴＧＴＴＡＴ
ＡＧ
ＡＴＡＣＣＡＡＡＧＡ
ＴＧＴＴＴＴＴＴＴＡ
ＴＡＡＧ
ＴＧＧＡＴＴＴＡＡＡ
ＧＣＴＧＧＴＧＴＴＡ
ＡＡＧ
ＧＡＡＴＣＣＴＣＣＡ
ＣＴＧＧＴＡＣＡＴＧ
ＧＡＣＧＡＴＧＣＴＡ
ＴＧＡＴＣＴＣＧＡＡ
ＧＣ
ＣＣＴＧＧＡＧＡＡＧ
ＧＣＴＡＴＧＡＴＣＴ
ＴＧＡＡＧＣＡＧＴＴ
ＡＴＴＧＣＴＴＡＴＧ
ＴＴＧＣＴＴＡＴＣＣ
ＴＧＴＣＴＴＣＧＴＧ
ＧＴＧＧＡＣ
ＧＧＴＡＧＡＧＣＴＧ
ＴＡＴＡＴＧＡＡＴＧ
ＣＡＡＴＧＣＡＴＧＣ
ＡＧＴＴＡＴＴＧＡＣ
ＣＡＴＧＧＴＡＴＧＣ
ＡＴＴＴＣＣＧＴＧＴ
Ａ
ＧＧＴＡＴＧＣＡＴＴ
ＴＣＣＧＴＧＴＡＴＴ
ＡＧＣ
ＧＡＧＡＴＣＡＴＡＴ
ＴＣＡＣＧＣＴＧＧＴ
ＡＣ
ＧＧＴＡＴＴＣＡＴＧ
ＴＴＴＧＧＣ
ＧＧＡＧＣＡＧＴＴＧ
ＣＴＡＡＴＡＧＡＧ
ＧＧＡＣＧＴＧＡＴＣ
ＴＴＧＣＴＣＧ
ＧＧＴＡＡＧＴＣＣＡ
ＴＣＡＧＴＣＣＡＡＡＣ
ＧＣＡＡＴＡＴＡＴＴ
ＧＡＴＴＴＴＣＴＴＣＴＣＣＡＧ
ＧＴＧＡＡＡＴＣＡＡ
ＧＴＣＣＡＣＣＡＣＧ
ＴＧＣＡＧＴＡＡＡＡ
ＣＣＡＣＣＴＧ
ＧＣＧＴＧＡＡＴＡＴ
ＧＧＴＣＴＣＣＡＣＣ
ＴＣＴＴＣＧＴＧＧＴＧＧ
Ｓ
ｒ
ｅ
３
３
３
ｅｏ／『ノ０〉ｎＵ（。ｏ５０〉勺Ｉ『ｊ４０／０２４，
ｐⅡｐ０ⅢＴａＪｐム肋剖ＯＢ。＆ソ升ムム肥旧冊が刀だ
遜山皿川仙川⑪山肌山田川川畑
舳沁沁沁肋あめ巾めめ伽加川物
可ｄ０『ｌ７ｌ７３５１５／０４‐５６１２〆０１２７』
蹄仙帆過罹蛙緬廼裡控趣起趣雑妙妙妙趣廸釦
▽
Ｓｅｑｕｅｎｃｅ
ＣＴＡＣＧＧＴＡＣＣ
ＡＧＣＧＴＧＡＡＴＡＴＧＧＴＣＴＣＣＡＣＣＧＧ
ＡＡＣＡＣＣＴＧＧＴ
ＡＡＡＧＡＡＡＣＣ
ＴＴＡＡＴＧＣＴＧＧ
ＣＡＴＡＴＧＣＣＡＡＡＣ
ＧＣＡＧＣＴＡＡＴＴ
ＣＡＧＧＡＣＴＣＣ
ＣＴＣＴＡＡＴＣＣＡ
ＣＴＧＡＧＣＴＡＣＡ
ＧＡＡＧＧＧＡＴＴＣ
ＧＡＡＣＣＣＴＴＧ
ＧＧＧＴＴＡＧＡＡＧ
ＧＧＡＴＴＣＧＡＡＣＣＣＴＴＧＡＣ
ＡＴＧＡＴＡＴＣＣＡ
ＣＡＴＴＣＴＴＣＡＣＡ
lMarksprimersdesignedbyManhart（1994);2marksprimersdesignedbyManhart(1994)andshortened
byTBubotaetal.（1999);3marksprimersdesignedbyTBubotaetａＬ（1999);4marksprimersdesignedby
TBubotaetaL(2000);otherswereadditionallydesignedbyHTsubotaandYAgeno．
withsearchlevel3，ａｎｄｔｈｅＣＮＩｓｅａｒｃｈｗａｓｒｅ－
ｐｅａｔｅｄｌ５ｔｉｍｅｓstatingwithtreesgeneratedby
lO,OOOreplicationswithrandomadditionse-
quencesJnanalysesbyPAUP＊４０b８，MPtrees
weresearchedusingheuristicsearchstrategy
withthefbllowingsettingsineffect：allcharac-
terswereequallyweighted,characterstateswere
treatedasunordered,gapsasmissingdata,Startingtreesobtainedviastepwiseaddition,random
additionsequencewithlOO,O00replicates，treebisection-reconnection（TBR)branchswapping
steepestdescentoffcollapsezero-lengthbranches，
534
ＨｉｋｏｂｉａＶＯｌｌ３,No.３，２００１
andMulTreesoptions
Mm碗"碗ﾉﾙﾉﾉhoo‘α"α“曲一MLtreeswere
constructedwiththefbllowingthreeprograms：
ＮｕｃＭＬｉｎＭＯＬＰＨＹｖｅｒｓｉｏｎ２３ｂ３ｐａｃｋａｇｅ
(Adachi＆Hasegawal996),ftlstDNAmlversion
1.2.2（Olsenetall994),andPUZZLEversion
50（Strimmer＆Haeselerl996)．FortheML
analyses,HKY85model(Hasegawaetall985）
wasusedastheestimatemodel・Inanalyseswith
NucMLMLtreesweresearchedwiththelocal
rearrangementmethodffomtheNJtreebyNucMＬａｎｄＮＪｄｉｓｔＬ２Ｓ（Adachi＆Hasegawa
l996)InanalysesbyPUZZLE,MLtreeswere
searchedusingtreereconstructionlnanalyses
byfastDNＡｍｌ,MLtreesweresearcbedwiththe
globalrearrangementsoptions・Best-confbnned
transition/transversion(Ts/Tv)parameterwases‐
timatedbasedonthecalculationsbｙＮｕｃＭＬｅmployedthe"‐topt，，optionparameteras413(8.26
asNucMLnotationalsystem),ａｎｄｂｙＰＵＺＺＬＥ
ｅｍｐｌｏｙｅｄａｔ４１３，usingparameterestimation
basedonNJtreefbrsubstimtionprocessandrate
variationTheTs/Tvparameterwasthenusedas
4・l3fbrtheanalysesbyfastDNAm１．
７>でｅｃｏｌ〃αrJso〃ｗノノルノ09-ﾉﾉﾙeﾉﾉﾙＣＯＣノーTree
comparisonwiththeMLcriteriabyNucMLwas
calTiedouttoevaluateｔｈｅｒｅsultingtreesbythe
fburmethods・Severalmeasureswereusedesti-
matingthelog-likeliboodofdifferenttreetopologies,standarderrors（SES）ofthedifferencein
log-likelihCod(Kisbino＆Hasegawal989),and
thebootstrapprobabilityestiｍａｔｅｄｂｙｔｈｅＲＥＬＬ
(resamplingofestimatedlog-likelihoodofsites）
method（Kishino,ｅｔａＬ１９９０；Hasegawa＆Ki
shinol994)UsingNucML,abootstrappingtest
withlocalbootstrapprobabilities（LBPs;iｎ％）
(Adachi＆Hasegawal996)wereappliedrespec‐
tivelyfbrthebestMLtopology
Co"se"s"ｓ舵e-Astrictconsensustreealso
computedfbrthehighestlog-likelihooｄｔｒｅｅｓｏｂ－
ｔａｉｎｅｄｂｙＣＯＮＳＥＮＳＥｉｎＰHYLIP3573cpack-
ageprogramsbyFelsenstein（l980-200Linhis
httpsite,seeAppendixB)．
Results
０６/αｊ"eaSe9"e"Ｃｅ血/α
Newr6cLsequenceswereobtainedfbr7spe-
ciesbelongingto4genera：＊ＤＭ?ｅｇＭｊａｐｅｒ""
Ｃｌﾚｲ/α/α,Ⅲaff:ｃα/火ﾉｲe"川,MJsmpo"ｚａｐ"/Ｃｈｅﾉﾉα，
*ＭＭｙｊ/従r肌＊Ｍ〃"ｃｊ"(/Ｍ"碗,比ｍａｍｐ/ｉ〕ﾉﾉｰ
ﾉz"〃〃/ＣＭ/z/"7ａｎｄ＊71〃"ｊめ"ｍｐｒＭｃａｂ２ｘ,fbr
whichthe4specieswithasterisksareshownby
entiresequences・Thesequencesweredeposited
intheDDBJunderserialaccessionnumbers(Tablel)ThesequenceswereentrustedtotheDNA
databaseoftheDDBJ/ＥＭＢＬ/GenBanklnterna-
tionalNucleotideSequenceDatabaseCollaboration
OnlyasinglesitedifferenCe(＝９９９％homolo‐
gy)wasfbundbetweenthesequencesof〃jsme‐
9Ｍα〃"ぬ(〃/α(ABO51229,TBubotaetaL2001；
ftomMtDefU[＝MtPeitawu],Taiwan,AAmi〕ﾉα‐
〃ααα/､６７)ａｎｄＴｐｅ'w"clIZﾉﾉα/α(ABO71415,the
presentpaper；fiDmDoilnthanon，Thailand，
4ｋ卯ｍａｎﾉｶｰ/)．
S〃crzl花q/se9"e"Ce伽α
Thefillllengthoftheentirer6cLsequences
newlyobtainedfbrthe4speciesinvestigatedwas
L428bplon9.Thelengthprovedtobethesame
asthoseofotherhypnobryaleanspeciespreviouslyreported（Tsubotaandcoworkersl999，
2000,2001)．Eachentirenucleotidesequence
couldbetranslatedrespectivelyinto475-amino
acidsequencesandonestopcodonassumingthat
thegeneticcodeisuniversal
TheG+Ccontentsofthenucleotidesequences
wereinvestigatedfbrallknownsites，ａｎｄｔｈｅｙ
ｒａｎｇｅｄｆｉＤｍ３７､８ｔｏ40.7％、Theseproportions
provedtohavenosignificantdifferenｃｅｓｉｎＧ＋ｃ
ｃontentsonthebasisofthosepreviouslyreported
fbrhypnaleanmosses：Ｂ'Ｗｈｅ/Reﾉﾉαｈｅ"o"〃
(39.6％)，〃kjα〃omsch"cﾙﾉﾉ(39.4％),Ｈ)や""ｍ
ｃ叩砥s(/br腕ｅ(38.7％)andHﾉﾉ"c/beZgﾉﾉ(38.2％）
(Tsubotaandcoworkersl999,2000)．
Sb9"e"Ｃｅα/jg"ｍｅ"／
TWodatamatrices(datasetlandll)werepreparedfbrphylogeneticanalysesbymanualalignmentofthesequences・Therewereseveraldeletionsiteｓｉｎｓｏｍｅｓｅｑｕｅｎｃｅｓｆｉ･omthedatabase
Thenucleotidesubstitutionsfbreachposition
wereestimatedbytranslatingthemintoamino
acidsequences・Mostwerethirdpositionsbut
synonymousnucleotidesubstitutions，andmost
oftheotherswitbaminoacidreplacementwere
conservativesubstimtions
lnthedatasetsaftercompletedeletionofgaps
orundeterminedsites,wetreatedbothsequences
HTsuBoTA,Ｈ・AKIYAMA,TYAMAGucHIANDH・DEGucHI
ｏｆかｉｓ腕egjMα〃"｡tﾉﾉｍａａｎｄ７７ｐｅ７""伽ﾉﾋﾝmasa
singleoperationaltaxonomicunit(OTU),since
､osignificantdifferencewasobservedbetween
thetwospeciesinsequencedataThesubstitutionofasi､gledifferentsiteistransitional(betweenAandGinbases）andnonsynonymous
(betweenAandTinaminoacids）whichin‐
volvedanaminoacidreplacement・
Ｔｈｅｄａｔａｓｅｔｓｈａｄａ５％chi-squaretｅｓｔａｎｄ
ｐ－ｖａｌｕｅｂｙＰＵＺＺＬＥ，andallthedatapassed
thesetests・Thedatasetscanexertnonegative
influenceontheanalysesjudgingffomthecomparativelycompactdistributionofG+Ccontent
andtheresultofthetests・Nucleotidefiequencies
estimatedｆｉｏｍｄａｔａｓｅｔｌｗｅｒｅＡ＝２９．０％，Ｃ＝
17.4％,Ｇ＝21.7％ａｎｄＴ＝31.9％,ｆｆｏｍｄａｔａｓｅｔ
ｌｌＡ＝28.9％，Ｃ＝１７．７％，Ｇ＝21.5％ａｎｄＴ＝
31.9％、Homologiesamongsequencesfbrdata
setlrangedfi･ｏｍ９２３ｔｏｌＯＯ％，ｆｂｒｄａｔａｓｅｔＩＩ
ｆＴｏｍ９４２ｔｏ１００％・
Ａｔｏｔａｌｏｆｌ,Z22sitesof63r6cLsequenceｓ
ｗｅｒｅｕｓｅｄｆｂｒｔｈｅａｎａｌｙＳｅｓｏｆｄａｔａｓetLInthe
aligned76cLdatasetsitesfbrthedatasetL321
sites（＝２６３％ｉｎ1,222sites）werevariable；
namely）Z21sites(＝６８８％in321sites)werebi-
narytransitions,S9sites(＝18.4％)werebinary
transversions,and41sites(＝12.8％)weremulti‐
statesiteSAtotalofl,Ｚ５４ｓｉｔｅｓｏｆ４２'6cLse-
quenceswereusedfbrtheanalysesofdatasetII
Inthealignedr6cＬｄａｔａｓetsitesfbrthedataset
lI,Z64sites(＝21.1％ｉｎ1,254sites)werevariablejnamely，l95sites（＝73.9％ｉｎ２６４ｓｉｔｅｓ）
werebinarytransitions,４７sites(＝17.8％)were
binarytransversions,ａｎｄ２２ｓｉｔｅｓ(＝8.3％)were
multistatesites．
Ｐ/Uﾉﾉoge"erjcα"α枕応
FordatasetLaphylogenetictreewasconstructedbyNucMLbasedonthedatasetwith63
76cLsequencesandl,222bpinlengthTheNuc-
MLtreefbrtheｄａｔａｓｅｔｌｉｓｓｈｏｗｎｉｎＦｉｇＬＴｈｅ
ｔｒｅｅｓｔronglysupportsthemonophylyof“the
sematophyllaceousmosses，，defInedbyTBubota
andcoworkers(2000,2001）ａccommodatingthe
membersoftheSematophyllaceaeandasmall
numberofspeciessofarplacedintheHypnaceae
togetherwiththeEntodontaceae(s・str.）andthe
Thuidiaceae(s,str.)（100％LBP）Ｗｅ,therefiore，
madeadetailedanalysisofdatasetll
FordatasetlLphylogenetictreeswerecon-
535
structedbasedonthedatasetwith4２r6cLse-
quencesfbrmembersofthefamiliesSematophyllaceae，Hypnaceae，Entodontaceaeand
Thuidiaceae,ａｎｄ1,254bpinlengthAtotalof48
nonoverlappedtopologieswereobtainedfTom
theanalyses：ｌＭＬｔｏｐｏｌｏｇｙｂｙＮｕｃＭＬ；lML
byfastDNAml;ltopologybyPUZZLE1NJ,
２０ＭＥａｎｄ６９ＭＰｂｙＭＥＧＡ２ｏｆｗｈｉｃｈ２ＭＰ
ｔｏｐｏｌｏｇｉｅｓａｒｅｔｈｅｓａｍｅｏｎｅｓａｓＮｕｃＭＬａｎｄfast-
DNAml；ａｎｄｌ２ＭＰｂｙＰＡＵＰ*４Thetopology
byPUZZLEismultifUrcate，sincethequartetpuzzlingtreewasnotcompletelyresolved・More
detailedtopologiesweresearchedthroughthe
obtainedtrees（Table３).Thetopologieswere
comparedwitheachotherusingthelog-likelihoodvalueandPrindicatesbootstrapprobability
byNucMLwiththeＨＫＹ８５ｍｏｄｅｌａｎｄｕｓｅｒ,s
treeoptionConsequently，ｆａｓｔＤＮＡｍｌｂｙＭＬ
ｍｅｔｈｏｄａｎｄＭＥGA2byMPobtainedthebest
topologyTree-1inTable3isthebesttreebythe
likelihoodcriterionThemostMLtreebythe
likelihoodcriterionfbrthedatasetlIispresented
inFig2、Log-likelihoodvaluefbrtheMLtｒｅｅ
ｗａｓ－５２１３７±215.7．ThePrvalueindicatedfbr
theMLtreｅｂｙＲＥＬＬｍｅｔｈｏｄｗａｓＯＯ６０，of
whichvalueappearsalittlelow・Eachtopology
withhighlog-likelihoodvaluewasapproximately
closetooneanother・TheLBPsfbrtheclades
wereofcomparativelylowvalues(39-100％）Becauseeachtopologydidnotgainadecidedadvantageoverothertopologies，astrictconsensus
treeof29high-rankingtreeswithasterisksinTa-
ble3wasalsoobtained,andispresentedinFig3・
ＴｈｅＭＬｔreefbrmssomenotablecladesshown
ｉｎFig.２:(1)sevenbasicclades[B剛力e'℃ﾉﾉCJ-E〕ﾉｰ
ﾉａｊｓｊａｃ/bllp/ia-吻肋clade（100％LBPvaluein
MLtree);ZSoc/αdieﾉﾉZMplt)ﾉＣｈｅﾉﾉﾋJclade(63％in
MLtree);mjrheﾉﾉ"mclade;んα"/ｈｏ/γ/lMTchj"m
clade;比/e叩/（〕ﾉﾉﾉﾉ""，clade（94％inMLtree)；
Trismegistiaclade(99％inMLtree);ａｎｄＳｅ腕amp/Ｍ/z"，Ｍｃ'ｗﾌo加加-M2joj力ecj""-かjchos/e‐
/e"m-Pqpﾉﾉﾉﾉ虚Qp叩clade（100％ｉｎＭＬｔｒｅｅ)]；
(2)twoprincipalclades［BわれＭｂ－Ｅ〕ﾉﾉmsjac/ｂｌｐｈａ－〃ｋｊａｃｌａｄｅａｎｄｈｏｃ/bJcﾉﾌﾞeﾉﾉα=｡〃CheﾉﾉﾋJ
cladeuniteB川ｈｅ花ﾉﾙｰlineage（62％ｉｎＭＬ
ｔｒｅｅ);ａｎｄ比加α/Qp/Dﾉﾉｍｍ－化ｍｐｏｒｊ"腕-/Ｗｏﾉﾉｶe‐
ｃｊ""7-かjc/jos/e/e""Z-P叩ﾉﾉﾉﾉc/iOpsなclade：Ｓｅ"mmpﾉｶ〕ﾉﾉﾉz""-lineage（100％inMLtree)];(3)two
paraphyleticlineages[Ｈｂｊｅｍｐ/lﾘﾉﾉﾉﾉ"ｍｃｌａｄｅａｎｄ
介伽egMacladeuniteHCremp/』!〕ﾉﾉﾉIrz""－
ＨｉｋｏｂｉａＶｂＬ１３,No.３，２００１
5３６
TheMLtreeisindicatedas‘ＭＬ，Thediffierencesinlog-likelihoodofalternativetreesfi･omthatofthe
Table３．NucMLanalysesfbrl,254bpof42r6cLsequences・The48high-rankingtopologiesareshown
fbnnula．‘ＭＥ，meanstreewitbtheleastsumofedgeｌｃｎｇｔｈｏｐｔｉｍｉｚｅｄｂｙＭＬｗｉｔｈＮｕｃＭＬＰｒindicates
MLtree(ﾙｰﾉjMz)areshownwiththeirSEfbllowing±estimatedbyKishinoandHasegawa,ｓ（1989）
bootstrapprobabilityofatreeamong48alternativesestimatedbytheRELLmethod(KishinoetaL1990）
ｗｉｔｈ10,000replicationslbpologieswithasteriskswercusedfbrthestrictconsensustree．
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
Ｍｏｌｐｈｙ,ＭＥＧＡ２
Ｐ
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＰＡＵＰ*４
ＭＥＧＡ２
ＰＡＵＰ*４
ＰＡＵＰ*４
ＰＡＵＰ*４
ＭＥＧＡ２
ＰＡＵＰ*４
ＰＡＵＰ*４
ＰＡＵＰ*４
ＭＥＧＡ２
ＰＡＵＰ*４
ＰＡＵＰ*４
ＰＡＵＰ*４
ＰＡＵＰ*４
ＰＡＵＰ*４
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
ＭＥＧＡ２
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ＭＥＧＡ２
ＰＵＺ７ｍＦ
＊＊＊＊＊＊
ＭＥＧＡ２
Ｍ
●●●●●●●ｃ●●●●●●●の●●●●●●●●●●●●●●巳●●●●●●●●●■巳●●●●●●
０３６２－５－２６１０７５７３８１２８９１３５０２３８２９３４２４００２７０１０１３－３０１１０
６７９６４１５２３３４２００３３２１０１２１０１０１１２００００００００００００００００００００
０００００００１００００００００００００００００００００００００００００００００００００００００
００００００００００００００００００００００００００００００００００００００００００００００００
Ｓ
■●●●●●■●●●●Ｏ●●●ＣＯｃ●●●●●ＣＯ●●●●●●●●■●■０●●ＣＯ●●■●■●
７７２６６８４１４２９－８４９８８６８３２０７７７２８６５４６３５５４３２２０９４５４６５０２
１
１
１１１１１１１１１１１１１１１１１１１１１１１１１１１１１１１１１２
２６７７６７７０９０９７９４４４１４４５５５１３５５５４５４４４４４５５４５４４５５６５６７３
１１１１１１１１１１２２２２２２２２２２３３３３３３３３３３４４４４４４４４４
２３４５６７８９０１２３４５６７８９０１２３４５６７８９０１２３４５６７８９０１２３４５６７８
’一一一一一一一一一一一一一一一一一引引訓刮訓引剤訓訓訓刮引引引引訓引刈引引引ユユユュユユユ⑲
００１２２２２３３３３３４８８８９９１１１２２３３３３４５５６６６６７７８８９０１１２３３４’
●●●●●■●■●●●●＄。●●●■●●●ｐ●●●Ｂの●●●●■□●■Ｂ●●■●●●■●◆●●
５７２３２３８００１５５３６７６６９１２１４５７８７８００３１４８８１１０３５５２５４４８６４
比十一十一十一十一十一十一十一士十一十一十一十一十一十一十一十一十一士士十一十一十一十一士十一十一士士十一十一十一十一十一十一十一十一十一十一十一十一十一十一十一十一十一十一十一
ｔ
ＭＥ
Consensus
Program
＊＊＊＊＊＊＊＊＊＊＊＊＊＊＊＊＊＊＊＊＊＊＊
ＭＥＧＡ２
ＷｗｗｗｗｗｗＷｗｗｗｗｗｗｗｗｗ疵ｗｗｗｗｗ疵疵ｗｗｗ爬肥爬爬岻疵爬噸噸疵疵爬疵川確雌旺疵朏乢
Ｍ
Ｐ
ｆａｓｔＤＮＡｍＬＭＥＧＡ２
ＭＬ
Method
Pｒ
L-/MＬ
Ranking
537
HTsuBoTA,Ｈ､AKIYAMA,ＴＹＡＭＡＧｕｃＨＩＡＮＤＨ､DEGucHl
乃伽egMa-lineage；ａｎｄ４ｃα"'んｏｒ〃"ｃｈｊ"m
sensustreeinFig3alsohasatendencytoward
clade,ａｎｄ汁ＭｅｇＭａｃｌａｄｅｕｎｉｔｅＨａｅｍｐ/Ｍ－
ﾉﾉ"ｍ-刀MegMa-4cα"/ｂｏｌｍ〕ﾉ"ｃｈｊ"m-lineage]；
and(4)theSematophyllaceae(sensuTsubotaet
aL）clade（77％ｉｎＮｕｃＭＬｔｒｅｅｆｂｒｄａｔａｓｅｔｌ，
66％inMLtreefbrdatasetlI)Thestrictcon-
theMLtree
Theresultsmakeninesuggestionsthat（１）the
Sematophyllaceae（sensuTsubotaetaL）area
monophyleticentitysupportingthefamilialdefi-
nitionandcircumscriptiononthefnmilyby
鰄F:Eii='二|雲ijil灘鰯:':……
B”
リ〃’ＷＥｒ２Ｍｆ
ＦＢｍｌ舵疋ﾉル
Lsocmzﾉﾉeノルｓ"'℃"/tJrな
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刺［ⅡⅡ釧剖
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豐二i二三三三；三i三i蚕;p擁鰡ｊｌ１ｗ……
5２
三二二姜芸ii謬議Ｉｉｊｉｉ蝋溌…伽
里亡:臣=三菫i;i鱗liiiiliii';iilliii二
坐LＦ
7６
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PID〃ＰアＰ
靹偏
100
8６
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nＩＯＺ
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i三i1i i i W1i霧i izi i i i ili i i i I:１
1００
6４
8］
Zdfj〃ZdlP7I
〕Ｓ、．'αＺ Ｏ〃ＩＣ〃腕
CJJOD〃ＷＪＨ〃７Ｚ
（．－の←の、一○□。、トコのこのの）①ｍの○ｍ’’二二Ｑ○一のＥのの
100
100
,〃〃〃７ＣＨＤ'℃ＳＳＩ兀
ソSrSraaelD〃Z4nrのｐｕＩａ匹
("ＷｊａｊＺｍどわ/、Ｓｍ、
DC
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5９
4２
三堅竺iiiiillli1lmi,i,,鰯１，
ﾉﾘedWjgmcﾉﾉﾉ、ａ
DＺａＺＨＺｅＺＪ
■■
０．０１substimtions/site
FigLThebestNucMI
FigLThebestNucMLtreefbrthedataｓｅｔｌｏｆｔｈｅｈｙｐｎａｌｅａｎａｓｗｅｌｌａｓｔｈｅｓematophyllaceousr6cLgenese-
quences(HKY85model;α/6＝826;lnL＝-74272±331.7)andthecladeoftheSematophyllaceae(sensuTsu‐
botaetaL),Entodontaceae(s・str.)andThuidiaceae(s・str.）Therootisarbitrarilyplacedonthebranchlcadingto
theThuidiaceae(s・str.)Localbootstrapprobabilities(LBP;％)morethan５０％areshownabovebranches
5３８
HikobiaVb１．１３，No.３，２００１
TBubotaandcoworkers(2000,2001);(2)DMegMa,MJS/OPO川〃ｋｊａａｎｄＨａｅｍｐ/lﾘﾉﾉﾉﾉ"ｍ
ａｒｅｐｌａｃｅｄｉｎｔｈｅｃｌａｄｅｏｆｔｈｅＳｅｍａtophyllaceae
(sensuTsubotaetal.),althoughthegeneraare
notmonophyletic;(3)〃肋fbnnsacladewith
ﾉ5ﾉﾋﾟﾉemp/1〕ﾉﾉﾉﾉ""'cladewith比/emp/iﾘﾉﾉﾉﾉz"γzが"e
andMJs/ＯＰＯ〃ａｓ"航/舵'ｗｍ;(5)tbegeneraSbm‐
ｑｍｐ/iﾘﾉﾉﾙ"０，かjcﾉ，Cs/e/e川ｄｃｌＷ,o'伽,Ｐ叩ﾉﾉﾉﾉｰ
c/iqpsjyandM2jo/"ecj"mfbnnaconsistentclade，
thegeneraofwhichweretreatedasrepresentativesoftheSematophyllaceae；（６）thegenera
乃ＭｅｇＭａ，〃ﾉＩｒｊａａｎｄＡｃα"ﾉﾙo〃ﾉﾘﾉ"cAj"胸，
BM力e"ﾉﾉαand町加ｊα火わんα;(4)〃伽egMq
consiststwodistinctclades:刀jsmegjMqAo〃〃αノー
sjjvar・AorrAaとｊｊ－Ｔｂａｆｆ：ｃαﾉﾋﾉｾ”"sjU-■ｐ/ｊｃａ/α
cladeandT。〃"｡tｲﾉma-T7peノッ"c/iｲﾉα/αclade，the
fbnnerfbnnsa乃応伽ｅｇｍｊａｃｌａｄｅｗｉｔｈＭｌｍＱｐｏ－
ｍα〃"cｊ"i/b/川ａｎｄＭ〃/c/ie"α,andthelatter
membersoftheWUkioideae,donotfbnnasin-
glecladeandarenotphylogeneticallycloselyrelatedtoeachother;(7)thegrouplinkingHb/e、‐
p/iﾘﾉﾉﾉﾉ"ｍｃｌａｄｅａｎｄ〃MegMacladecorre-
ＺＩＤ
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,ＩＪ２ｎｆ
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,ＢｍＺｈｅ花ﾉﾉｍｑ２ｃ
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空[｣H1;WI3H;二IifZ
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100
鴻鰯il1繍鰯曰 Thuidiaceae(ｓｓｔｒ）
０．０１substitutions/sitc
Fig2Thehighestlikelihoodtreeofthealigncdr6cLgenesequences(HKY85model;0(/P＝959;lnL＝-5213.7
±Z157byNucML)andcladesThehorizontallengthofeachbranchisproportionaltotheestimatednumberof
nucleotidesubstimtionsTherootisarbitrarilyplacedonthebranchleadingtotheThuidiaceae(sstn)Localboot-
strapprobabilities(LBP;％)ｍｏｒｅthan５０％areshownabovebranches.Ｂﾉ加力eﾉでﾉﾉﾋJ-lineagemeansDo/ｈｅﾉﾍﾞeﾉﾉﾋJP〕ﾉﾉαな』α火ｊＩｐｈａ－〃kja-Aoc/αﾙﾉﾉﾋＭ〃ＣＭ/qclade,andsubdividedintotwoclades:Ｂ、ﾉﾉﾉe"ﾉﾉﾋJ-P〕ﾉ伽jadalp〃α－
脚ﾉImcladeandAocﾉαcﾉﾉeﾉ/α-A〃c/ieﾉﾉaclade；ａｎｄＳｅ"ね'０ﾉﾌﾉ１１ｿﾉﾉ""T-lineageSemα'0p/iﾘﾉﾉﾙ'，Ｍｃｍｐｏ７Ｊ""７－
ﾉＷｏ/hecj""T-Djc/ios/e/e""T-PcJpﾉﾉﾉ伽Opsjsclade．〃e/erop/（〕ﾉﾉﾉﾉz"〃ｃｌａｄｅｍｅａｎｓｔｈｅＨ巴/e叩/1〕ﾉﾉﾉﾉ"加轍"eMJs/Opo"zaswh/}/(/セノw"7-刀ＭＣｇＭａｚｌ"ロノjｲﾉα/α一正ｐｅ'w"cﾙﾉα/qclade；ａｎｄ〃MegjsﾉﾉａｃｌａｄｅＤＭｅｇＭａ
Ａｏ"ﾉﾉｑＭｖａＭｏ"ｈαﾉｌＦｊｊ－ｍａｆｆｃａ仇花'Ｍ－Ｔｐ/ｊｃａ/a-MIq1s/OPO"!α""c/"(/b/Ｍ－Ｍｐ"/Cheﾉﾙ"TcladeHe/emp/（〕ﾉﾉﾉ川一
刀MegMa-lineageuniteswithHe/emp/iﾘﾉﾉﾉﾉ""，cladeandDMegMaclade．ｆﾉﾋﾞﾉemp/】ﾘﾉﾉﾉﾉ""７－〃ＭｅｇＭａ－
Ａｃα"ﾉﾉmrr/1〕ﾉ"c胸"7-lincagcuniteswith比！e叩/iﾘﾉﾉﾉﾉ""Zclade，刀MegMQcladcanCMcα"/〃ｸﾞﾉ!〕ﾉ"c力伽clade、
Sematophyllaceae（sensuTSubotaetaL）cladeuniteswith〃o/ｈｅﾉｾﾉﾉq-lineage，Ｓｃ"、/Op/Ｍ/"m-lineage，
比'e叩/1〕ﾉﾉﾉﾉ"加一DMegMCMcα"/how!〕ﾉ"c伽m-lineageand7伽/ﾉﾌﾞeﾉﾉ"mclade・刀ＭｅｇＭａｐ"/ＣＭ/αHerzogis
shownhereasMmOpo"、〃/bんe"""1(ined),andthenewcombinationwillbepublishedinaseparatepape正
５３９
HTsuBoTA,Ｈ・ＡＫＩＹＡＭＡ,ＴＹＡＭＡＧｕｃＨｌＡＮＤＨ､ＤＥＧｕｃＨｌ
spondstotheHeterophyllioideaeproposedbyDiscussion
Fleischer(1915-1922)andBrotherus(1924-1925）
Thesuhjectstobediscussedonthefamilialcir‐
exceptfbr〃肋,andtheWijkioideaeproposed
cumscriptionoftheSematophyllaceaeinclude（１）
byHedeniisandBuck（1999)wasnotsupported；
thedistinctionbetweentheSematophyllaceaeand
(8)Ｈｂ/emP/１〕ﾉﾉﾉﾉ"ｍ-刀ＭｅｇＭａ刊cα"'んorl御/（〕ﾉ"－
theHypnaceae;(2)thesubfamilialcircumscription
cﾙﾉ"m-lineageasaparaphyleticentitytothe比ｍ－
ａｍｐﾉi1)ﾉﾉﾉ""z-lineageuniteswithHb/emp/,ﾘﾉﾉﾉﾉ"moftheSematophyllaceae:(2-a)thecircumscription
clade,〃MegMacladeanMca肋。,町"chj"腕oftheHeterophyllioideae（orWUkioideae),and
clade,althoughsometopologiesshowtheline-（2-b)thecircumscriptionoftheClastobryoideaa
ageasmonophyletic；ａｎｄ（９）７ＺＭＭｊ"ｍａｐ－ａｎｄ（３）thesurveyofthephylogeneticrelaｔｉｏｎ－
ｐｅａｒｓｔｏｂｅｓｉｓｔｅｒｔｏｔｈｅＢ'、ﾉﾌﾞe'､eﾉﾉa-lineage，shipsbetweenthefamilySematophyllaceaeand
althoughthepositionisnotstableantheorderHookeriales・Concerningthesubject
８ｍ//zα℃ﾉﾉａｃｏｌ"p/α"α/α
B'ｗ〃e”ﾉﾉαノロ”jej
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LFOp/erygj"加花"e'ｗｍ
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Zsoc/αd7eﾉﾉａｓ"”"ﾉαr/kF
．店xjlheﾉﾉumclade
i霧蕊1,;iii]曰……伽
①⑦の①Ｅ｜‐ミミ仁。Ｅ当仁巨○名巨町○式
‐の這辺ロ①Ｅの堂トーミコミ雪仁Ｑｐ一色の工
露Ｍ１幻;；
‐←Ｅご一洲索識宇喀←このの
Mbjo/hecj"碗ｍｊｃｍｃａｚｐｍｍ
ＳｂｍａｍＰ/1〕ﾉﾉﾉ""ｚＰ"/c/zeﾉﾉ皿"？
『００Ⅱ‐ⅡⅡⅡⅡⅡⅡⅡⅡ８１Ⅱ］
7zp[ﾉﾉﾉzeﾉﾉ"ｍｐﾉα""、
Ａｃｍｐｏｒｍｍｓかα、ｊ"e"m
AcmPorJ"ｍＰ""ge"s
Pqpjﾉﾉﾉc/jOpsjs'ｍｃｍｓ"c/α
刀jchosje/e"ｍｐ叩ﾉﾉﾉ０s"加
乃ｊｃ/'Cs/eﾉe""Ｔｓ伽叩/（〕ﾉﾉﾉ"m
（．｜、］の⑩一○ロゴのトコのこのの）の、の。囚一云二Ｑ○一ｍ〔Ｅのの
の①、①Ｅ｜‐聖の」の名。」□
け/msjα火(りんαｊｅ""ｊ''０s"ｉｓ
〃ﾉIFja力or"Sc肋c肋
LOp""加伽ｊｏ－Ｗｊ火
鰯護(iilililiiIinmm…(…，
Fig.３．Thestrictconsensustrccfbr29trceswithhigh-rankinglog-likelihoodvalues・Thcrootisarbitrarilyplaced
onthebranchleadingtotheThuidiaceae(sstr.)．
540
ＨｉｋｏｂｉａＶＯＬｌ３,No.３，２００１
(1)mentionedabove，Tsubotaandcoworkers
(2000,2001）proposedtotreatthelhmilyina
widersense，andthepresentanalysisalsosupportsthatpropositionThepresentinvestigation
discussesonlythesubject(2-a)．
LGeneraintheSematophyllaceae
かむmGg伽ｊαα"c/MJsmPo腕ａ
Ｔｈｅｇｅｎｅｒａか応meg伽ｊａａｎｄＭｚｌ/qpo〃αwere
placedintheHeterophyllioideaebyFleischer
(1915-1922）andBrotherus（1924-1925）orin
theSematophylloideaebyTnnaｎｄＢｕｃｋ(1989）
Thepresentanalysisshowsthespliｔｏｆかjlmeg伽jaintotwodistinctclades：ｎｋｏ〃/iaZsノノvar・
ﾉﾋo〃ノロハノトＴｌａｆｆｃａﾉﾋﾉｾﾉｳﾞe"sな-Ⅲｐ/ｊｃａ'αｃｌａｄｅａｎｄ
Ⅵ〃"伽/α/a-mpeﾉｗ"伽/LJjaclade・Thefbrmer
cladeissistertothecladeMmQpomα〃"ｃｊ"加一
ﾉﾉ""?-Ｍ〃/Cheﾉﾉ""0．Mostspeciesof〃な"Zeg/sjjaandMJsmpo碗afbnnasinglecladebutsister
tothｅＳｂｍα'０p/1〕ﾉﾉﾙm-lineagcMeanwhile,the
latterissistertothecladeMJsmpomas"b/i/旅一
'w腕,andstronglysupportedThe正〃"伽/α/α-Ⅲ
pe'w"〃αｒＱ－ＭＭﾂﾞ】/舵'wmcladeissisterto
HC/empﾉiﾘﾉﾉﾉ伽〃"ｅｗｉｔｈ９４％LBRSeveral
morphologicaldifferencescanberecognizedin
theleafshape,leafundulationandalarcellsbetween正ﾉﾋo〃/iaZsjjvar､肋ｒｒｈａＡｊｊ，ｎａｆｆ:ｃα/cノセノー
e"ｓｊＳａｎｄＺｐ/ｊｃａ/α，ａｎｄＴＭ/"伽/α/αａｎｄＺｐｅ－
'w"ぬ/α/α(seeAkiyama＆mubota2001).The
presentinvestigationsuggeststhateachofthese
generaispolyphyletic，andthetwodistinct
cladesincludingsomespeciesofthosegenera
mentionedabovewererecognizcdnearthepositionofsuchtypicalgeneraoftheSematophyUaceaeas比加α'０p/1)ﾉﾉ伽anMcmpo'川(Figs・Z
and3）
DjsmegjMα〃"ぬ/mawasdescribedbyBrotherus（1926)fi･omTniwan,ａｎｄＦｐｅｌｗ"伽〃α
byDixon(1936)ffomLaosHorikawaandAndo
(1964)transfelTedⅢpe7""伽/αＺａｔｏＭｍＯｐｏｍａ
ｏｎｔｈｅｂａｓｉｓｏｆｔｈematerialfiomMt､Doilntha-
non,Thailandintheirfloristicworklnthepresentinvestigation,the76cLsequencesofn〃"｡iｲｰ
ﾉaZafi･omMtPeitaw,Taiwan(４A(〕′α腕αera/､６７）
andT7pe'ｗ"dbﾉﾉα/αfromMt・Doilnthanon,Thai-
land（』ﾉhリノａｍａ７１ｈ－Ｉ）respectivelydifferfrom
eachotheｒｏｎｌｙｉｎａｓｉｎｇｌｅｓｉｔｅｉｎｔｈｅｉｒｒ６ｃＬｓequencedata(ｌｂｐｐｅｒ1,428ｂｐ;99.9％inhomol‐
ogy).Thediffbrencewasatransitionalsubstitu‐
tioLbutanon-synonymouscodonthａｔｈａｓａ
changeoftheaminoacid・ArikawaandHiguchi
(1999)reportedintraspecificsequencevariations
of'6ｃＬｇｅｎｅｆｂｒＰ/cJgjoﾉﾉiecj""’ｅ"Ｗﾌﾉlﾘﾉﾉﾉ"ｍ
(ｌbase),ﾉ？〃ｅｍｏｍ１/ｅ（１base),ＰＭｂ城c"〃"〃
(４bases),没〃eckeMcノセ"ｍ(Sbases),ａｎｄＲｈ〕ﾉﾉﾉｰ
dia火llP伽かj9"e〃８（１base).Comparedwith
thesevariations,thedifferenceof76cLsequence
betweenz〃"伽ﾉｍａａｎｄＺｐｅｒ""｡ｉｲﾉﾋＪｍｉｓｓｍａｌｌ
ａｎｄａｔｔｈｅｓａｍｅｌｅｖｅＬＴａｘonomicrevisionfbr
thesetwospeciesisneeded,particularlybasedon
themorpbologicalfeamres．
HCjemp/iﾘﾉﾉﾉﾉz"〃
RobinsonandReed(1966)revisedHe/emph〕ﾉﾉｰ
ﾉﾉ"腕,andtreatedthegenusasmonotypicconsistingofHが"ｅ(Hook.)Ｍ､FleischSeki（1968）
reviewedtheJapaneseSematophyllaceae，and
treatedthefamilyinanarrowerSense，placing
"e/emp/i〕ﾉﾉﾉﾉ""ZintheHypnaceae､Nishimuraet
al（1984)proposedanewcircumscriptionofthe
HypnaceaeandplacedHeje叩ﾉiﾘﾉﾉﾉ川inthe
Hypnoideaeofthefamily・HedeniisandBuck
(1999)alsotreatedthegenusasamemberofthe
HypnaceacMeanwhile，ＴａｎａｎｄＢｕｃｋ（1989）
madeasynopticreviewofPhilippineSematophyllaceae，andtreatedHe'emp/Ｍﾉﾉ"masa
memberoftheHeterophyllioideaeintheSematophyllaceaelEubotaetaL（2001）suggestedthat
"e/emp/iﾘﾉﾉﾉﾉ"masrepresentedbyHが"ｅａｎｄ
Ｈ〃e"、伽""7ＢｒｏｔｈｅｘＲｄｅｌａＶａｒｄｅ＆Th6r・is
notmonophyletic,ａｎｄＨｑがi"eissistertothe
cladeDなmeg伽ｊａｐ/ｊｃａＺａ－ｒｋｏ〃/iaLsjjvar・ｋｏｒ－
ﾉﾙα肋(ａｓＴｋＭｈａＭ)intheMLtree,although
itspositionisnotstable・Inthepresentstudywith
moresupplementalsequencedataofthegenera
〃ＭｅｇＭａａｎｄＭⅣOpo”α,Ｈ〃"ehaspro‐
ventohaveastablepositiontogethｅｒｗｉｔｈｔｈｅ
〃jlD腕eg伽jα〃"｡bﾉﾉｍａ－Ｔｐｅｒ""ＣｌﾚﾉﾉﾋJ/a-MzSrQPomas"b/)/旅'wmcladeThecladeismorphologicallycharacterizedbythescalarifbnnalarcells
ofleafOntheotherhand,Ｈ〃e〃α/Cs"ｍｉspositionedintheBMﾉﾉe”ﾉﾉα-町/α奴ａｄｂｌｌｐｈａ－〃kja
cladewithstrongLBPs・Thisresultshowsthe
polyphylyofthegenus比ｊｅｍｐ/iﾘﾉﾉﾉ伽，supportingthetaxonomictreatmentbyRobinsonand
Reed(1966).Ourresultssuggestthatthegenera
ﾉｈｅｍｐ/jﾘﾉﾉﾉﾉ"腕，〃ＭｅｇＭａａｎｄＭＭＱｐｏ"、be
placedintheHeterophyUioideae(sensupresent）
intheSematophyllaceae,differingffomthetreatmentsbySeki（1968),NishimuraetaL（1984）
ＨＴｓｕＢｏＴＡ，Ｈ､ＡＫＩＹＡＭＡ,Ｔ・ＹＡＭＡＧｕｃＨＩＡＮＤＨ、ＤＥＧｕｃＨＩ
5４１
andHedeniisandBuck(1999)．
thegenus．
ＡＣα"ﾉﾊorr/ＩＷＣ/zj""7α"c/ＴＩＭ/Mzl"T
HedeniisandBuck(1999)suggestedthaMcaﾙ
ﾉﾊo〃/Ｍｃﾙﾉ"ｍｉｓｃｌｏｓｅｌｙｒｅｌａｔｅｄｔｏｔｈｅｇｅnera
DMegMaand〃kjaintheircladisticanalysis
basedonmorphologicalandanatomicalcharacterslBubotaetaL(2001)sbowedthatthegenera
化α"/AC〃/【〕ﾉ"c/Zjz"'７ａｎｄ７ＺＭｈｅﾉﾉz/"Ｔａｎｄｔｈｅ
ＢＭ/ie花ﾉﾉα-町/αＭＩＣ/ｂｌｌｐ力α-〃kja-Lyoc/α`jeﾉｰ
ﾉa-z4plﾜﾉc力e"αcladeareplacedsidebysideinthe
mostMLtreaalthoughtheirpositionｓａｒｅｎｏｔ
２.ThefaInilyandsubfamilycIassification
Hae叩/l(〕ﾉﾉﾉﾉo雄αeα"ロノ〃kjoj伽e
Fleischer（1915-1922）dividedtheSemato‐
phyllaceaeintofbursubftlmilies，andBrotherus
(1924-1925)fbllowedthesystemJntheBrotherusssystem，thesubfamilyHeterophyllioideae
arerepresentedbyseveralgenera,ｓｕｃｈａｓＨａｅノー
叩/Ｍｊ川乃ＭｅｇＭａ,MJsmﾉﾌﾞo"ｍａｎ。〃Ａｊｑ
(Fi９４，Ａ）Seki（1968)circumscribedthefiamily
ｓｔａｂｌｅｉｎｔｈｅｏｔｈｅｒｔｒｅｅｓｌｎthepresentstudy，
ＡＣα"ﾉﾙor'/iWcﾊﾉ"腕didnotfbnnasingleclade
togetherｗｉｔｈ〃/ｔｊａｎｏｒ〃ｊｓｍｅｇＭａ，ａｎｄｉｓ
ｐｌａｃｅｄａｔｔｈｅmostbasalpositiontotheSCmaｍ－
ｐ/Ｍｍ腕‐lineageandHb/emp/】ﾘﾉﾉﾉﾉz"'7-〃M7egls‐
/ja-lineageintheMLtreeJnthepresentanalyses，ＡＣα"ｊｈｏｒ'今/lﾘﾉ"ｃｈｊ"腕wasalsoplacedinthe
Heterophyllioideae(sensupresent)intheSematophyllaceae、７hxj'んeﾉﾉ"ｍａｐｐｅａｒｓｔｏｂｅｓｉｓｔｅｒｔｏ
ｔｈｅＢｍｊｈｅだ"a-lineage，althoughthepositionis
notstable．
〃kja
Thegenus〃kjahasbeenplacedunderthe
varioussuMlmiliesinthefamilySematophyllaceae；theHeterophyllioideae（ｅｇ，Brotherus
l924-1925),theSematophylloideae(ｅｇ,Ｔａｎ＆
Buckl989)ortheWUkioideae(Hedeniis＆Buck
l999).HedeniisandBuck（1999)treated〃kja
asthemostcloselyrelatedgeｎｕｓｔｏＤなｍｅｇＭｊａ・
Tsubotaandcoworkers(1999,2000)reportedthe
sequenceonlyof〃ノノor"sch"cﾊﾉﾉ（Ｄｏｚｙ＆
Molk)HACrumfbr〃ﾉﾋjaandshowedthespe‐
ｃｉｅｓｔｏｂｅｐｌａｃｅｄｉｎｔｈｅＢ'Ｗﾙe'､e"αclade,where
mostspeciesofB'Ｗ力e/qeﾉﾉaarealsopositioned
TEubotaetal.（2001）showedthepolyphylyof
吻肋asrepresentedbythreespeciesof〃ん！α：
〃ｄ白/7e兀加ﾉﾉα（MittexRenauld＆Cardot）
Ｈ・ＡＣｒｕｍ，〃ノio'"SCA"cﾙﾉﾉａｎｄ〃〃αノセJ"ＬＷﾊﾉﾉ
(Broth.）ＨＡＣｒｕｍ，andsuggestedthegenus
mightbecloselyrelatedtoBMhe花ﾉﾉaandPI〕ﾉ〃‐
ｓｊａ火Ilp肋ratherthanto〃ＭｅｇＭａｏＭｃａﾙ
ﾉﾊo〃/iWcﾙﾉ"mlnthepresentinvestigation，the
polyphylyof〃AjaisalsosupportedFurtherinvestigationisneededonthephylogeneticpositionsof〃肋,incorporatingmorespeciesofthe
genus，especiallythetypespecies，爪ｅｍｅ""α/α
(Brid)ＨＡＣｒｕｍ,fbrabetterunderstandingof
inthenarrowestsensewhichcorrespondsｔｏｂｏｔｈ
ｔｈｅＳｅｍα/叩/Ｍ/z""-1ineageandapartoftbe
hotAelneﾉﾉﾋJ-lineageasrepresentedbytheLＳＯＣ/ａ－
ｄｊｅﾉﾙｰﾉ4pji〕ﾉＣｈｅﾉﾉａｃｌａｄｅａｎｄｔｈｅＴｈＭｈｅﾉﾉ""７
cladeinthepresentanalysis(Fig.４，Ｂ)Hetreat‐
edsomegenerawhichwereplacedintheHeteropbyllioideaeinthesystemofFleischerand
Brotherus,suchasHaemp/iﾘﾉﾉﾉﾉｚｌｗａｎｄＷﾉｌｉﾉﾊﾉα,as
membersoftheHypnaceae・Ｎｉｓｈｉｍｕｒａｅｔａｌ
(1984)placeｄﾉﾘC/e、ﾉﾌﾉＭｊ"川ntheHypnoideae
oftheHypnaceae,proposinganewcircumscriptionoftheHypnaceacTanandBuck(1989)fbl‐
lowedBrotherus（1924-1925）andrecognized
thefburfamilies，althoughsomemodifications
weremadeintheirproposal，ｓｕｃｈａｓｔｏｅｘｃｌｕｄｅ
幼l(〕ﾉc"eﾉﾉafi･omtheClastobryoideaeandtoplace
itintheHeteropbyllioideae(Fig.４，Ｃ)Hedentis
andBuck（1999)treatedthefiamilyinanarrower
Sense，placingBMhe”//α，ＨＣ/e叩Ａ１〕ﾉﾉﾉﾉ""’ａｎｄ
itsalliedgeneraintheHypnaceae（Ｆｉｇ４,Ｄ)．
Theyrecognizedtwosubfamilies：theSematophylloideaeandtheWijkioideaeintheSematophyllaceae,andrefierredtothelatterapartofthe
Brotherus，ｓ（1924-1925）Heterophyllioideae
TheysuggestedthattheWUkioideaearelikelyto
bethemoreancestralsubfamilyoftheSemato‐
phyllaceaeandthegeneraofthesubfamilyare
mostlikelymembersoftheHypnaceac
ThepresentanalysisshｏｗｓｔｈａｔｔｈｅＨＣ/e/℃‐
ﾉﾌﾉ１１〕ﾉﾉﾉﾉ""７－〃MegMa-lineageandtheHe/e、‐
Ｐ/11〕ﾉﾉﾉﾉ""?－胴smegMa-Acα"//7o〃/iγ"c肋"??-line‐
ageareparaphyleticgroupstotheSemamp/Ｍ‐
/"碗‐lineage（seeFigs、２）Ｔｈｅ〃巴lfemp/i1〕ﾉﾉﾉﾉz"７，‐
胴sﾉ"egjs/ja-lineagecorrespondstoapartof
Brotherus，ｓ（1924-1925）Heterophyllioideae
(Fig.４，Ｅ）Ourresultsdonotrigorouslysupport
theproposalmadebyHedentisandＢｕｃｋ（1999）
whoplaccdthethreegenera吻仙,刀ＭｅｇＭａ
ａｎｄＡｃα"/〃o〃/Ｍ７ｃﾙﾉ"mintheWijkioideae．
①
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切
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Ｅ"'０．ｍｃ/、此"ｇＰｒｊ
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乃jsmegな"α〃"伽/αね,ＺＹｐｅ'w"伽ﾉα/ａ
MJS/cpomaM2βﾉ雄'wm
Hbjerqp/｛〕ﾉﾉﾉj"加城"Ｂ
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-
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勘joqlo〃'ｗ６ｊｃ""diｨＳ
DzrodO〃sca67jdb'１ｓ
＆rodO〃ﾉzWdbUs
乃町ｍｅｇＬｓｒｊａﾉto〃/zaLsjjvar・ＡＣ〃ｈａＬｓｊｊ
乃応megLmZJP/ｊｃａｍ
ＭＪｓｊｑｐｏｍａＰｚｲﾉb/ze/mm
Mzsrqpomaz`"ｃｊ"(/Ｍ"加
四一六○亘画く。
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比加α叩ﾉ!〕ﾉﾉﾙ"zpzｲﾉＣｈｅﾉﾙｍ
Ｓｂｍα/cp/１〕ﾉﾉﾉ""2S幼ﾉz""2ﾉﾉesubsp./叩o"ｊｃ"腕
乃なmegなｔｊａａＨｃα/dbだ）、ｓ
Zzx"ｈｅﾉﾉ皿加ｐ/α"zJm
Acmporj"碗ｓ〃α"/"ezJm
Acrqpormmp""9℃'１ｓ
Ｐｑｐﾉﾉﾉﾉdiqpsな、αc'℃ｓｔｊａａ
刀ｊｃ/ｍｓｊｅ陀皿ｍｐｑｐﾉﾉﾉCs皿加
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Acrqpo7jHms〃αｍｊ"e"m
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PcJpﾉﾉﾉjdIqpsjsmacmszjcza
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DjChosleﾉe""WjSSqpﾉﾘﾉﾉﾙｍ
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Se"zaZqp/Ｍ/皿加ｐ"/Cheﾉﾉ皿加
比､α/qp/Z〕ﾉﾉﾉ""zsz`6/z"'Mesubsp./叩o"jC""，
刀jsmegな"αaff:ｃα/C1b”"sな
DjsmegjsrjaAo〃haLFjjvar・肋〃/zaZsjj
〃jsmegjSrjap/jca1a
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544
ＨｉｋｏｂｉａＶｂＬＩ３,No.３，２００１
However,thedefinitionofthesubfamilythatincludesthegenerawhicharelikelytobe
themostprimitiveonesoftheSematophyllaceae，
thathavetheleastspecializedperichaetiaand
sporophytes,isconsiderablebecauseoftheirplesiomolphiccharacterstates､ThegeneraAcα"/hocﾉα成""?（＝〃ｋｊα),ａPひＣＭﾉﾋﾝ，Ｃ/川o6r〕ﾉeﾉﾉａ
ａｎｄＺＺｍﾉﾉｶeﾉﾉ"mhavebeenexcludedfiDmthe
SematophyllaceaebySeki(1968).Inthepresent
analyses,thesegenerawereplacedintheBMルー
ｅＭ/a-lineageorthe7Zzx肋eﾉﾉ""Tclade,outsideof
theSematophylloideae（sensupresent）orthe
Heterophyllioideae(sensupresent).Wealsodisa‐
greewithTanandBuck(1989)whoplacedゆり
CheﾉﾉaintheHeterophyllioideac
Weproposeheretwolineageswhichrepresent
twodistinctsubfamilies:(1)the比"mjqpﾉiﾘﾉﾉﾙ"?‐
lineageasthesubfamilyofamonophyleticentity,andiscomparabletotheSematophylloideae
ofBrotherus(1925)andtheSematophyllaceaein
thenarrowersenseofSeki（1968)andHedeniis
andBuck(1999),placingBMie”ノルanditsal‐
liesintheB'℃肋e”"a-lineageinthecladeofthe
Sematophyllaceae；ａｎｄ（２）theHB/e叩h〕ﾉﾉﾉj"－
刀MegMa-z4cα〃/io'”"ｃｈ伽-lineageasthe
subfamilyofaparaphyleticentity(maybemonophyletic),andiscomparabletotheHeterophyl‐
Heterophyllioideae．
Ｍ､'pho/ｏｇｊｃα/cﾉﾉα'αc/e酒ｑ/ﾉﾉｹＣｆノセ花畑p/Ｍ/ojdbae
Sporophyticcharacterssuchasperistometeeth
andexothecialcellscanbetaxonomicallysignificantfbrtherecognitionofspecies,generaorsubfamilies・TbemembersbelongingtotheSematophylloideae（sensupresent)ｏｆｔｈｅＳｅ碗amph)ノノー
ルm-lineagearecharacterizedbylesswelldevelopedinnerperistometeeth，andbyshort
androundcollenchymatousexothecialcells，as
previouslypointedoutbySeki(1968)andHedeniisandBuck（1999).Somecharacterstatesincludingalongsetaandcharacteristicalarｃｅｌｌｓ
ｃａｎｂｅｆｂｕｎｄｉｎｓｏｍｅｏｔｈｅｒｓｅｍａtophyllaceous
generaincludingBm/力e花ﾉﾉﾋ７，whichwasplaced
intheHypnaceaebySeki（1968）andHedeniis
andBuck（1999).Ｔｈｅwell-developedalarcells
arealsooneofthemoststrikingcharactersto
recognizetheSematophyUaceaeasreferredtoby
Nishimuraetal(1984)andTsubotaetaL(2001）
Theecologicalpreferenceasepiphyteswasalso
importanttorecognizetheSematophyllaceaeand
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Fig.４Continued．
andtheinclusionoMcα"伽'町"c伽"'inthe
ミ冒旨・ヘュミ昌邑：ミミミ員ミミミｓごミ男
EPresentstudy
lioideaeofBrotherus（1924-1925）withtheex‐
clusionof〃ＡｊＱｔｏｉｎｔｈｅ脇oﾉﾉje"ﾉﾉﾋJ-lineage
HTsuBoTA,Ｈ､AKIYAMA,ＴＹＡＭＡＧｕｃＨＩＡＮＤＨ､DEGucHI
theEntodontaceaeasstatedbyVitt（1984）and
TsubotaetaL(2001)．
TheWUkioideaewererecognizedbyanumber
ofcharacterstatesthatareplesiomorphicwithin
theSematophyllaceae,excluding町o/ｈｅ花ﾉﾉaand
itsalliedgenerawhichareplacedintheHypnaceaeassuggestedbyHedeniisandBuck（1999)．
TheHeterophyllioideae（sensupresent)arealso
recognizedbycharacterstatesthatareplesiomorphouswithintheSematophyllaceae（sensuTsu‐
botaetaL).WecanrecognizetheHeterophyllioi‐
deae（sensupresent）asaparaphyleticentity，
whichhaveprimitivecharacterstatesinthefami1y・ThesubfamilyHeterophyllioideaecanbedistinguishedfi･omtheSematophylloideaebyhavingthefbllowingfeatures：（１）scalarifiormand
collenchymatouscellsinalarceUsorjuxta-alar
cel1s；（２）dimorphicleaves（betweenstemand
branch);(3)weU-developedandlongperistomes；
and（４）alongseta,However,Ｂ川/ieだ"αand
somemembersoftheHypnaceaearealsocharacterizedbysomeofthecharacterstates・TheresultssuggestthatB'Ｗ〃e”"αａｎｄitsalliesconstimteaspecializedgroupoftheSematophyllaceae，
andthecharacterstatesamongthemａｒｅａｌｓｏ
ｃｏｍｍｏｎｔｏｓｏｍｅｍｅｍｂｅｒｓｏｆtheHypnaceae・
ThesecharactersmaybeprimitiveintheHypnales,butnotwithintheHeterophyllioideaeSo，
wetreattheSematophyllaceaeinawidersense，
andconsiderthatthewell-developedalarcells
tingedwithbrownandtheparticularecological
requirementsareworthyoftaxonomicrecogni‐
tionastheSematophyllaceaaaspreviously
pointedoutbyTBubotaetal(2001）
Cﾉﾉ℃""7scrlipﾉﾉo"SCWﾌﾉ!〕ﾉﾉogc"e/ｊｃｇｍぃｗｊ/ﾙﾉ〃
/ｈｅＳｅｍａｍｐ/iﾘﾉﾉﾉﾋJceae
ThepresentresultsshowasinglecladeaccommodatingtheSematophyllaceaeinawidersense，
includingＢ'Ｗhe花//α，〃kjaandtheirallied
genera,andsomeotherspeciescurrentlytreated
asmembersoftheHypnaceae(e､９，Hyp"""〃ｉｓ‐
/o-yかｊ火,Aqpjer〕lgj""ｚ／e"ｅｌｗｍａｎｄ比/emp/】!〕ﾉﾉｰ
ﾉﾉ""z〃emamsz"").ThedefinitionoftheSemato‐
phyllaceae(sensupresent)iscongruentwiththat
oftheSematophyllaceae（sensuTsubotaetaL)，
andthepresentcircumscriptioniswiderthanthat
madebyNishimuraetaL(1984)andVitt(1984）
TWostablelineageswerefbund,corresponding
tothepreviouslyrecognizedSematophylloideae
5４５
andHeterophyllioideaepreviouslyrecognized
Theresultsofthepresentstudydonotconcur
withthesystemproposedbyHedeniisandBuck
(1999),wheretheSematophylloideaeweredivid‐
edintotwogroups,〃oﾉﾙe花ﾉﾉaanditsalliedtaxa
wereplacedintheHypnaceaaandthegenera
〃Ａｊａ，ＤＭ７ｅｇＭａａｎｄ化α"'んｏＷＭ７ｃ〃jzJmin
theWijkioideaeHowever,apointoftheWUkioi‐
deaeasaprimitiveentityoftheSematophyllaceaeshouldbeconsidered,sincetheHeterophyllioideaeandmemberｓｏｆｔｈｅＢ'Ｗ"e'､eﾉﾉa-lineage
aredistinguishedfi･omeachotherbysomeofthe
characterstatesmentionedbyHedenasandＢｕｃｋ
(1999).ＷedefinetwosuMumiliesSematophyl‐
1oideaeandHeterophyllioideae：thefbrmerasa
monophyleticentity,andthelatterasaparapby‐
letic(ormaybemonophyletic)entityOntheba‐
sisofthephylogenetictreethatshowstheHeter‐
ophyllioideaeareparaphyletic，ormonophyletic
insomehigh-rankingtrees，wetentativelyincludesuchspeciesandgeneraasHCZemp/11〕ﾉﾉﾉﾉ""？
q"7"eandthegenera〃ＭｅｇＭｑ,MJsmpo碗α
and化α"/力ｏｒ町）"c/ｉｊ"minthatsubfamilyofthe
Sematophyllaceac
WecannotdeflneheretheB'Ｗ/ieﾉｲe"a-lineage
andtheTZIxjjheﾉﾉ""７clade，sincethedatafiorthe
ClastobryoideaeseemtobeinsufflcienttobeanalyzedWC,however,considerthattheBmﾉﾙe花ﾉｰ
ﾉa-lineageandthe7hxﾉﾉﾉﾉeﾉﾉz/mcladeshouldneverbeplacedintheHypnaceaaalthoughthe
Hypnaceaeareapolyphyleticentity(seeHedeniis
l995;TsubotaetaL1999;DeLunaetaL2000)．
Conclusion
lnconclusion,oursmdysuggeststhat（１）the
Sematophyllaceaewithwidersenseproposedby
TsubotaandcoWorkers(2000,2001)areresolved
asamonophyleticgroupandwellsupported;(2)
theWijkioideaeproposedbyHedeniisandBuck
(1999）arenotmonophyletic，andtheHetero‐
phyllioideae（sensupresent）arewellsupported
inthecladeoftheSematophyllaceae；ａｎｄ（３）
ＤＭｅｇＭａ,Ｍｌ１ｓｊｑｐｏｍａａｎｄ〃んｊαhaveproven
tobepolyphyleticinthecladeoftheSematophyllaceae・
Inthepresentinvestigation，thenecessityof
taxonomicalrevisionfbrthepolyphyleticgenera
かＭｅｇＭａ,MzljOpo"、,Ｂ川/je”ﾉﾉαand〃kja
issuggestedPhylogeneticrelationshipsbasedon
moreprecisemoleculardatashouldbesurveye．
5４６
HikobiaVOl．］３，No.３，２００１
betweentheClastobryoideae，includingdmm6M"〃ａｎｄ肌Jge"ｊｅ"し7,andtheothersuMlmilies
oftheSematophyllaceae、Furtherdetailedsmdy
mightsupporttheindependenceofthetwo
clades，whichwouldsuggesttherecognitionof
twosubfamiliesoroftwoindependentfamilies
correspondingtotwoclades:Ｂ川〃eだ"a-lineage，
andthelineageoftheSematophylloideaeandthe
Heterophyllioideae．
Tsubota,Ｈ､，Gonzalez,，.,Newton,ＡＥ.＆Ｓｈａｗ，
ＡＪ、2000．OrdinalphylogenywithinthehypnobryaleanpleurocarpousmossesinferredfromcladisticanalysesofthreechloroplastDNAsequence
datasets:／r"L-EZps4and76cLBryologistlO3：
242-256.
，Newton，Ａ、Ｅ，Withey,Ａ､，Gonzalez，，．＆
Mishler,ＢＤ・l999Thetransitiontopleurocarpy：
Aphylogeneticanalysisofthemaindiplolepidous
lineagesbasedonr6cLsequencesandmolphology・
BryologistlO2:634-650．
Acknowledgements
WewouldliketothankH・Mohamed，Ｓ－Ｈ．
Ｌin，WTabachaiandMMohamedfbrtheir
greathelpincollectingmaterials,TArikawa,Ｄ・
GonzAlezandEDeLunafbrtheircollaboration，
YAgenofbrherhelptodesignnewpriｍｅｒｓＷｅ
ａｌｓｏｔｈａｎｋＷＲ・BuckfbrcheckingtheEnglish
textandfbrsuggestionsFinancialsupportsfi･ｏｍ
ＭＥＳＳＣｇｒａｎｔｓｌｌ６４０６９８ｔｏＨ、Deguchi；
Ｏ９０４１１６５ａｎｄｌ３３７５００３ｔｏＪＭｕｒａｔａ；and
Dixon，Ｈ、Ｎ・l9360nacollectionofmossesけom
LaosAnnalesBryologici9:61-72．
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TsubotaetaL（1999);〃o/helｲeﾉﾉα/jer6aceaSakuraiex
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ABO39789(as及汕!〃α火」Ip〃αｊｅ""〃s"ｉｓ),TEubotaet
5４８
ＨｉｋｏｂｉａＶｂＬｌ３,Ｎｏ．３，２００］
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q'7"ｅ（Hook）MFleisch,ABO51218,Tsubotaetal
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Varde＆Th6r.,ABO29391,TSubotaetal（I999MFoc/αcﾉｸﾞeﾉﾉａｓｚ"℃z'〃/Ｓ（Dixon）ＢＣＴａｎ＆Mohamed，
ABO39784（ａｓ/VeQc'叩Ｍ２1脚/7ageﾉﾉ(/、""),Tsubota
etaL（2000)；MJS/Oﾉﾌo"7α〃/Ｃｈｅﾉﾉａ（Herzog）ｉｎｅｄ，
ABO71410(ａｓ刀．/Ｍ７ｅｇＭＱ〃/b〃eﾉﾉqHerzog),Malaysia,Sabah,Ttlwau,Ｈｄﾉhi〕ﾉａｍＱＭＪﾉﾉα"-50Ｊ（ＨＹＯ)，
thisstudy;ＭＩＳ'０pol"ａｓ"肋/舵""〃Horik＆Ando，
ABO71411,Thailand,ChangMaLDoilnthanon,Ｈ
Ａｋｌｉ〕ﾉα"?αル２(HYO),thissmdy;ＭＩＦ/Opo"mzl"ｃｊ'7ノー
/Ｍｚ"〃（Broth）Broth,ABO71412,Malaysia,Sabah，
KinabaluNationalPark,fMk〃αﾉ"ｑＫﾉﾙﾉ0/６(ＨＹＯ)，
thissmdy；Ｍｂｊｏﾉﾙｃｃ/z/"7〃7/ｃｍｃＱ'〃"7（Ｈｏｏｋ）Mitt.，
ABO51223,TsubotaetaL(2001);PclIpj//MOpsjs〃7ａｃ－
'ＤＭＣ/ａ（Broth＆Paris）ＷＲＢｕｃｋ＆ＢＣＴan，
ABO51220（ａｓＲ〃叩/7ﾉﾛﾉＤＭＣ/7z/"７〃7αcﾉ℃Mclfz/"1)，
TsubotaetaL（2001)；け/αｊｓｊａｄｂ伽α／e"zｲﾉﾉＤｓ/'７ｓ
(Ｂｍｃｈ＆ＳｃｈｉｍｐｅｘＳｕｌＬ）ＷＲＢｕｃｋ,ABO51219,
TsubotaetaL（2001)；Ｓｅ”α/0p/Ｍｍ"，ｐ"/c〃eﾉﾙﾉ'７
(Cardot)Broth,ABO7141Mapan,Honshu,Hiroshimaken,Hatsukaichi-shi,Ｈ肋6０，３７３６（HIRO),ｔｈｉｓ
study；Ｓｅ"，α/叩/iyﾉﾙﾉ〃sw6〃z"Ｍｅ（MtjllHaL）M
Fleischsubspノ叩o"jα"〃（Broth）Seki,ABO39675,
Tsubotaetal．（2000)；五Ｍｈｅﾉﾉ""リノα"""７（Brid）
Mitt.,AF233573,DeLunaetal(2000);刀ｊｃ〃Cs/e/e""T
MssOp/Z〕ﾉﾉﾉz"〃（Ｈａｍｐｅ＆MijlLHal）AJaeger，
ABO51226,Ｔｓｕｂｏｔａｅｔａｌ（2001)；７Wc/josje/e"ｍ
ｐ叩ﾉﾉﾉCs"〃（Ｈｏｍｓｃｈ)AJaegeLAF233574,ＤｅＬｕｎａ
ｃｔａＬ（2000);*刀ＭｅｇＭｑａｆｆ：ｃα/火花'伽（SulL）
Broth,ABO71414Malaysia,MalayPen,Cameron
Highlands,TanabRata,Ｈ乃"Ｍｑイ387(HIRO),this
study;刀ＭｅｇｊＭＭｏ'7AαＭ(Dozy＆Ｍｏｌｋ）Broth
vaMorﾉﾉ，α肋,ABO51227(as77ko"〃αＭ),Tsubotaet
al（2001)；かﾊﾟﾉ"egMαノブe'w"`zﾉﾉα/αDixon，
ＡＢＯ７１４１５，Thailand，ChangMai，Doilnthanon，H
4kli〕ﾉα"?α７Ｗ(ＨＹＯ),thissmdy;ＤＭｅｇＭＱｐ/ｊｃａ/α
(ｍｓ),ABO51228Jsubotaetal（2001)；刀ＭｅｇＭｑ
ｚﾉﾉ、ﾉﾉｲﾉﾋJ'αBroth＆Ｍ､Yasuda,ＡＢＯ５１２２９，Tsubotaet
al.（2001)；〃Ａｊｑｄｂ化x(/b/ｊＱ（Mitt・exRenauld＆
Cardot)ＨＡＣｍｍ,ABO5122LTsubotaetaL(2001)；
〃Ａｊα〃o'wschz/cﾉﾘﾉﾉ（MFleisch）ＨＡＣｒｕｍ，
ABO29383,ｈｕｂｏｔａｅｔａＬ（1999)；〃ﾊﾉα〃αAα"ｊＷﾊﾉﾉ
(Broth）HACrumABO512Z2［ａｓβ'℃/ｈｅ'℃ﾉﾉα
"qkcJ"/Ｗｋｊｊ]，ＴｓｕｂｏｔａｅｔａＬ（2001)；Entodontaceae
E"/oQlb〃ｃﾉﾉα此"geｸﾞノ（Paris）Cardot，ABO50993,
TsubotaetaL（2001)；Ｅ"'odb〃ノ姻伽ｓ（GrifD
A・JaegenABO50994,ＴｓｕｂｏｔａｅｔａＬ（2001);Ｅ"/odb〃
"Ｍ"ＷＳ（Ｈｏｏｋ）Hampe，ABO24640,Arikawa＆
Higuchi（1999);Ｅ"jodb〃ﾉw6jc""ぬ/ｓ(Mitt.)AJaeger，
ABO29386,Tsubotaetal（1999);Ｅ"'０．b〃sca6r枕"s
Lindb.,ABO50995,Tsubotaetal（2001);Hypnaceae
H〕抑"〃ｃ叩'でSSi/blwleHedw.,ABO39674,Tsubotaet
aＬ（2000)；Ｈ)Ip"""ｿﾞﾉﾉ"Ｃｌ/beZgノノMitt，ABO29390,
Tsubotaetal.（1999);ﾉIyp""、ﾉﾜﾉ"ｍａＱ/blwleWilson，
ABO29384,ＴｓｕｂｏｔａｅｔａＬ（1999)；ノリ)p""、伽'０－
Ｗﾉｿﾞ火(Broth)Paris,ABO50991,TsubotaetaL(2001)；
んOP/er〕/ｇｊｚｌ１腕／e"e"ｲ腕（Ｓｗ.）Mitt.，AF233569，De
LunaetaL(2000);ｏｒﾉﾉ､ﾉﾙec/"/、W/ibsce'7s(Dicks・ｅｘ
Ｂｒｉｄ）Ｓｃｈｉｍｐ,ABO5095LTBubotaetaL(2001);け‐
/αｊＦｊａｊ"/ﾉﾌﾞcα/α(Hedw）Ｓｃｈｉｍｐ,ABO24642[asけ/ｑｊ－
ｓｊｅﾉﾉα/"Ｗｃｑ/α(Hedw.)Ｇrout],ArikawaandHiguchi
(1999)；Thuidiaceae肋ｊｅ""eﾉﾉαα6/eﾉﾉ"ａ（Hedw.）
MFleisch,AFOO5519,Goffinet,Bayer＆Vitt(1998)；
71/7"/吻加ｑＭｃα/"/"〃（Hedw.）Ｓｃｈｉｍｐ,AFl58177，
DeLunaetal(2000);Tﾙ"/伽mplMcqbAx(MUllHaL）
AJaeger,ＡＢＯ７１４１６，Japan，Kyushu，Kagoshimaken，YakushimaL，ＨＺＭｉ６ｏ/α３８６２（HIRO)，ｔｈｉｓ
study；７ﾙ"ｊｃ//z"ﾌﾌﾉでcog"〃"？（Hedw.）Lindb，
ABO19476,ＭａｅｄａｅｔａＬ(2000);LeskeaceaeBo"/ＱＷ
〃"e"ﾉノ（Broth）Cardot,ABO24963,ＴＢｕｂｏｔａｅｔａＬ
(1999);PlagiotheciaceaeP/ZZgjoﾉﾉiecj"〃〃ｃｃ舵'Djdbzｲ〃
Schimp.,ABO24630,Arikawa＆Higuchi(1999);Ｐ/cJg/o/ｈｅｃｊＭ７〃emoraﾉピ（MittjAJaeger,ABO29387,
TsubotaetaL（1999);AmblystegiaceaeCMje'190"e"α
c叩Ｍ７ｊｑ(Hedw.）Locske,ABO24678,Arikawa＆Hi-
guchi（1999)；BrachytheciaceaeMﾉz"℃c/､ﾉﾋ珈叩"１owjcz〃（OGBorshch）Steere＆WBSchofield，
ABO29389,TsubotaetaL(1999);Ｐ/､()ﾉﾉ０１p"肋/、ﾉＴｐＱｒ－
ｊＭ/bｓ（Hedw.）Dixon，ABO29385,Ｔｓｕｂｏｔａｅｔａｌ
(1999);Ｒ/'〕ﾉ"c〃Cs/egjzl"7pα/Ｍ/bﾉﾉz/ｍ(Mitt)AJaeger，
ABO24944Tsubotaetal（1999);HylocomiaceaeR/iﾘﾉｰ
肋αc/ＣＩＰ/ｊＷｃＷ"ｊｃ"s(Reimers)TJKop.,ABO39788,
TsubotaetaL（2000);FontinalaceaeFo"""αﾉﾉＭ"ﾉﾉｰ
ﾉ〕ｗｅ//ｃＱＨｅｄｗ，ABO50949,ＴｓｕｂｏｔａｅｔａＬ（2001)；
MniaceaeM7/z"Ｗｈｏ"7so"ｊｉＳｃｈｉｍｐ.,AFOO5518,Gof
HnetetaL（1998);Ｐ/LJg/ｏｍ"ｊｚｲ〃α叩加/"'〃(Hedw.）
TJKop,Ｕ87082(asMiW"ﾉ"c"Sp肋/"〃),Lewisetal
(1997)；Ｐ/cJg/０ｍ"ｊ"〃ノcZpo"/c"腕（Lindb）ＴＪＫｏｐ，
ABO50992,TsubotaetaL(2001);HedwigiaceaeHedL
Mg/αcﾉﾉﾉα/α(Hedw）PBeauv.,AFOO5517,Goffmetet
aL（1998)；OrthotrichaceaeU/o/α/"/eａ（Hookf＆
Wilson)Mitt,AFOO5540,GoffinetetaL(1998）
ＡｐｐｅｎｄｉｘＢＨｏｍｅｐａｇｅＵＲＬｓａｎｄＦＴＰｓｉｔcsofsofiwarefbrthephylogeneticanalysisanddatabase
Sb/iwα”
ｆｔｌｓｔＤＮＡｍｌ＜http://geta・life・uiuccdu/-9ary/＞
ＭＥＧＡ２＜http://www・megasoftwarenet/＞
ＭＯＬＰＨＹ＜ftp://ftpjsm・acjp/pub/ISMＬＩＢ／
ＭＯＬＰＨＹ/＞
ＰＨＹＬＩＰ＜http://evolutiongenetics・washington
edu/phyliphtml＞
*ThisspecimenistentativelyidcntｉＨｃｄａｓ刀･jsmegJs-
"αｃα此ルノセノＭｓ,sincethespecimenexaminedhasaleaf
characterdeviatingfromtheusualfbrmofthespecies：
theleafbordersarelesswell-developed
ＨＴｓｕＢｏＴＡ,Ｈ､AKIYAMA,ＴＹＡＭＡＧｕｃＨＩＡＮＤＨ、ＤＥＧｕｃＨＩ
ＰＵＺＺＬＥ＜http:〃ｗｗｗ・tree-puzzlcde/＞
Ｓｃleanandourprogramsdownloadsite
＜http://homehiroshima-uacjp/chubo/＞or
linkedwith＜http://homehiroshima-uacjp／
miyajima/＞
Ｄａｍ６Ｑｓｅ
ＤＤＢＪ＜http://www・ddbLinigacjp/＞
Sequencedatadownloadsite
＜http://ftp2ddhjnigacjp:8000/cgi-bin/getLentry・
ｐｌ?Accessionnumbeか（Forexample,alinktoan
entrywithaccessionnumｂｅｒＡＢＯ７１４１６ｉｓ
＜http://ftp2ddbj・nigac・jp:8000/cgi-bin/get-entry・
ｐｌ?ABO71416>）
Ａｃｃｅｐｔｅｄ３０ＩＸ,２００１
5４９
近年,ナガハシゴケ科(Sematophyllaceae)内の系統関
係について盛んに議論がなされている．本研究では，
刀jsmegjsﾉﾉα属やモリクサゴケ属("c/emp/1〕ﾉﾉﾉﾉ""7)，Ｍ１Ｋｓ－
ｍｐｏｍａ属，トゲハイゴケ属(〃仇)の系統学的な位置づ
けとその単系統`性を確認するため,合計63種のr6cL遺伝
子の塩基配列を用いて検討を行い,ナガハシゴケ科とそ
の近縁分類群についての系統学的な知見を提示した.そ
の結果,Tsubota他(2000,2001)で提案された広義のナガ
ハシゴケ科の単系統性が支持された.またHedeniisand
Buck(1999)により提案されたトゲハイゴケ亜科(Wijkioideae)は多系統群であることが示された．さらに乃/ｓｍｅ－
ｇＭａ属とMJS/OPO'"α属，トゲハイゴケ属は多系統群とな
った．これらの結果はTsubota他(2001)で示されたナガハ
シゴケ科内の二つのクレードを支持するとともに多系統
々
坪田博美･秋山弘之･山口富美夫･出口博則：葉緑体
となった属の再検討の必要,性を示した．また,モリクサゴ
r6cL遺伝子に基づく乃ＭｅｇＭａ属とその近縁属(ナガハ
ケ亜科(Heterophyllioideae)をナガハシゴケ科内の側系
シゴケ科)の分子系統学的位置
統群として認識した．