HikobiaHikobial3：645-665.2002 13: 645-665. 2002

MolecularMolecularphylo窪enyOfhypnobrJ/aleanmOssesasin化rredfroma phylogeny of hypnobryalean mosses as inferred from a large-scalelar淫e-scaledatasetofchlOroplastlｂｃＬ，withspecialre他rencetothe dataset of chloroplast rbcL, with special reference to the

HypnaceaeHypnaceaeandpOssiblyrelatedfnmilies1 and possibly related families I

HIROMIHIRoMITsuBoTA，ＴｏＭｏＴｓｕＧｕＡＲＩＫＡｗＡ，ＨＩＲｏＹｕＫＩＡＫＩＹＡＭＡ，ＥＦＲＡＩＮＤＥＬｕＮＡ，DoLoREs TSUBOTA, TOMOTSUGU ARIKAWA, HIROYUKI AKIYAMA, EFRAIN DE LUNA, DOLORES GONZALEZ,GoNzALEz,MASANoBuHIGucHIANDHIRoNoRIDEGucHI MASANOBU HIGUCHI AND HIRONORI DEGUCHI

TSUBOTA,TsuBoTA,Ｈ､,ARIKAwA,Ｔ,AKIYAMA,Ｈ,,ＤＥＬｕＮＡ,Ｅ,GoNzALEz,，.,HIGucHI,Ｍ H., ARIKAWA, T., AKIYAMA, H., DE LUNA, E., GONZALEZ, D., HIGUCHI, M. ４ &＆DEGucHI，Ｈ、2002.Ｍolecularphylogenyofhypnobryaleanmossesasinferred DEGUCHI, H. 2002. Molecular phylogeny of hypnobryalean mosses as inferred fromfiPomalarge-scaledatasetofchloroplastr6cL,ｗｉｔｈspecialreferencetotheHypnaceae a large-scale dataset of chloroplast rbcL, with special reference to the Hypnaceae andandpossiblyrelatedfamiliesl3:645-665． possibly related families. 13: 645-665.

▲ PhylogeneticPhylogeneticrelationshipswithinthehypnobryaleanmosses(ｉｅ,theHypnales,Leuco- relationships within the hypnobryalean mosses (i.e., the Hypnales, Leuco­ dontales,dontales,andHookeriales）havebeenthefbcusofmuchattentioninrecentyears and Hookeriales) have been the focus of much attention in recent years. HereHerewepresentphylogeneticinfierencesonthislargeclade，andespeciallyonthe we present phylogenetic inferences on this large clade, and especially on the HypnaceaeHypnaceaeandpossiblyrelatedftlmilies，basedonmaximumlikelihoodanalysisof and possibly related families, based on maximum likelihood analysis of 181l81r6cLsequences、Oursmdycorroboratesthat（１）theHypnales(ｓｓｔｒ.[＝sensu rbcL sequences. Our study corroborates that (I) the Hypnales (s. str. [= sensu VittVittl984])andLeucodontalesareeachnotmonophyleticentities、TheHypnalesand 1984]) and Leucodontales are each not monophyletic entities. The Hypnales and LeucodontalestogethercompriseawellsupportedsistercladetotheHookeriales;(2)Leucodontales together comprise a well supported sister clade to the Hookeriales; (2) theSematophyllaceae(s」at［＝sensuTsubotaetaL2000,2001a,ｂ])andPlagiothecia‐the Sematophyllaceae (s. la1. [= sensu Tsubota et a1. 2000, 2001a, b)) and Plagiothecia­ ceaeceae(s・str.［＝sensupresentDareeachresolvedasmonophyleticgroups,whileno (s. str. [= sensu present)) are each resolved as monophyletic groups, while no particularparticularcladeaccommodatesallmembersoftheHypnaceaeandCryphaeaceae;ａｎｄ clade accommodates all members of the Hypnaceae and Cryphaeaceae; and (3)(3)theHypnaceaeaswellasitstypegenusﾉﾘDlwz"川tselfwerepolyphyletioThese the Hypnaceae as well as its type genus Hypnum itself were polyphyletic. These resultsdonotconcurwiththesystemsofVitt(1984)andBuckandVitt(1986),whoresults do not concur with the systems of Vitt (1984) and Buck and Vitt (1986), who suggestedsuggestedthatthegroupswithasinglecostawouldhavedivergedfiFomthehypnalean that the groups with a single costa would have diverged from the hypnalean ancestorancestoratanearlyevolutionarystage，fbllowedbythegroupswithadoublecosta at an early evolutionary stage, followed by the groups with a double costa (see(seealsoTsubotaetall999;Bucketal2000）OurresultsfiPomlikelihoodanalyses also Tsubota et a1. 1999; Buck et a1. 2000). Our results from likelihood analyses ofofalarger6cLdatasetconfinnthesuggestionbysmallerdatasetsfiommultipleloci a large rbcL data set confirm the suggestion by smaller data sets from multiple loci firstbyDeLunaetal.(2000)andthenBucketaL(2000)thatthereisnoevidencefbrfirst by De Luna et a1. (2000) and then Buck et a1. (2000) that there is no evidence for supportingtheHypnales(ｓｓｔｒ.[＝sensuVittl984])andLeucodontales・Adetailedsupporting the Hypnales (s. str. [= sensu Vitt 1984]) and Leucodontales. A detailed reconsiderationofordinalclassihcationandfnmilialclassificationwithintheHyp-reconsideration of ordinal classification and familial classification within the Hyp­ nidaeiｓｎｏｗnecessary．nidae is now necessary.

KeyWords:HypnalesLeucodontales,Hookeriales,hypnobryaleanmosses,molecularKey Words: Hypnales, Leucodontales, Hookeriales, hypnobryalean mosses, molecular phylogeny,r6cLlargedataset,Hypnaceae,Sematophyllaceae,Plagiotheciaceae,Am-phylogeny, rbcL, large data set, Hypnaceae, Sematophyllaceae, Plagiotheciaceae, Am­ blystegiaceae

。 肋Ｄ'"ｊ７Ｍ６ｏ/ｑｄＭｒﾉﾉ℃"ＭＤｅｇ"ｃｈｊ,Depα""ze"ｔｑ/Ｂｊｏ/ogjcα/Ｓｂｊｅ"Ｃｅ,Ｇｒａｄ"α/eHiromi Tsubota & Hironori Deguchi. Department (j[ Biological Science. Graduate Sb/ioo/ｑ／血e"Ｃｅ,Ⅲ℃sﾉｶ〃α〔ﾉﾉｊｊｖｅｌＭ１ﾉ,hgamli〕ﾉα碗αノー3-/,〃jgZJSﾙﾉｰﾊjrosﾉi〃α-School o[ Science. Hiroshima University. Kagamiyama 1-3-1. Higashi-hiroshima­ Ｗ,ﾉﾌﾞIﾉ･Csﾉi〃α739-852．ﾉﾋＷ/Lshi. Hiroshima 739-8526. Japan. 7bmojiFz`9ＭﾉWmwq,Depαﾉﾌﾞ'"e"'０/Ｂｊｏ/Ogjcα/Sbje"Ｃ＆Ｆ,Ｇ７α`"α/e此/ＩＯＣ/q/ｍｉｅ"Ｃｅ，Tomotsugu Arikawa. Department of Biological Sciences. Graduate School of Science. U>ﾌﾙe'Ｍﾜﾉq/mtyo,〃O'7907-3-/,βz"7/bﾉo-k"’７bﾉb′o川-0033,上pα"・University of Tokyo. Hongo 7-3-1. BunAyo-ku. Tokyo 113-0033. Japan. 航rqMEMﾉhi〕′α',、,/Ｍｌｓａｌ''１ｑ／ﾉVb【/"'℃α'z`ﾉ仙加α'Ｍα〃jrjes,町ogo，YZlO′ojgaoAaHiroyuki Akiyama. Museum (j[ Nature and Human Activities. Hyogo. Yayoigaoka 6-cho"ze,５位"dtJ-sAj,Hyogo669-/ｺﾞｲｄﾉﾋZpα〃6-chome. Sanda-shi. Hyogo 669-1546. Japan. 〃αｍＤｅＬ""α＆ＤＣ/oresOo'1Ｚ｡/ez,ＤｅＰａｒｔａｍｅ"ｊＭｅＳＭ伽/ｊｃａﾘﾉｾge/αＭＭｍｔｏEli-ain De Luna & Dolores Gonzalez. Departamento de Sistematica Vegetal. 1nstituto ｃ/ｅＥｃｏ/ｏｇｍｌＡＣ.,ＸＭＪｐａ，比７．９/000,ＭｄＸｊｃｏ、de Ecologia AC. Xalapa. Ver. 91000. Mexico. MJSα"０６〃ｆﾉﾉgZlcAj，Depα"ｍｅ"／ｑ/ＤＣ/αノリＭＪ/jolZα/比ｊｅ"ｃｅＭ`Ｓｃ"〃，伽αAr"６０Masanobu Higuchi. Department (j[ Botany. National Science Museum. Amakubo ｲｰ/-ＩｍＭ"6α-sﾙﾉ,仏aMhH305-000LﾉﾋZpα",ＤＥＰα"ﾉ?Te"／ｑ/Ｂｉo/ogjcα/Ｓ℃/e"ＣＧＳ，4-1-1. Tsukuba-shi. 1baraki 305-0005. Japan;Department of Biological Sciences. G/ndZJα/ｅ比ﾉioo/ｑ/此ｊｅ"Ce,Ｕ>7〃e′Ｍﾜﾉｑ/、oﾉb′０，肋"ｇｏ７~3-/,Ｂ２`"ﾉtyo-ﾉ1Ｗ,Tbﾉbﾉ。Graduate School (j[ Science. University (j[ Tokyo. Hongo 7-3-1. Bunkyo-ku. Tokyo 〃3-0033,上pα〃113-0033. Japan.

'ContributionfiOmtheLaboratolyofPlantTaxonomyandEcology,HiroshimaUniversity・ＮＳｅｒ・No.S２ＬI Contribution from the Laboratory of Plant Taxonomy and Ecology, Hiroshima University. N. Ser. No. 521. 646 HikobiaVoll3,No.４，２００２Hikobia Vol. 13, No.4, 2002

Introduction talesandHookeriales）fbrcorroborationofthetales and Hookeriales) for corroboration of the phylogenetichypothesisofDeLunaetal.(2000）phylogenetic hypothesis of De Luna et al. (2000) Thehypnobryaleanmosses，includingtheor-The hypnobryalean mosses, including the or­ ａｎｄＢｕｃｋｅｔａＬ(2000);３)toclarifythefiamilialand Buck et al. (2000); 3) to clarify the familial dersHypnales，LeucodontalesandHookeriales，ders Hypnales, Leucodontales and Hookeriales, relationships，especiallybetweentheHypnaceaerelationships, especially between the Hypnaceae areoneofthemostdiversifiedmossgroups、are one of the most diversified moss groups. andrelatedfamilies，suchastheSematophylla-and related families, such as the Sematophylla­ AttentionhasbeenpaidtotheclassiflcationofAttention has been paid to the classification of ceaeandPlagiotheciaceae，withinthehypnobly-ceae and Plagiotheciaceae, within the hypnobry­ thehypnobryaleanmossessinceVitt（1984)pro‐the hypnobryalean mosses since Vitt (1984) pro­ aleanmosses，giventhattheLeucodontalesandalean mosses, given that the Leucodontales and videdanewsystemfbrtheBryopsida、Withinvided a new system for the Bryopsida. Within Hypnalesarenotmonophyletic;ａｎｄ４)totraceinHypnales are not monophyletic; and 4) to trace in thepleurocarpshesuggestedthat“adoublecostathe pleurocarps he suggested that" a double costa thephylogenythetransitionsofthecostafiDmthe phylogeny the transitions of the costa from ll groupasrepresentedbytheHypnaceae,Selnato-group" as represented by the Hypnaceae, Semato­ singletodouble，oneoftheimportantcharacterssingle to double, one of the important characters phyllaceaeandEntodontaceae，isdivergentfiDmphyllaceae and Entodontaceae, is divergent from fbrthefiamilialclassificationofthehypnobry-for the familial classification of the hypnobry­ ０ agroupwithasinglecosta，ｗｈｉｃｈｃａｎｂｅｒｅ－a group with a single costa, which can be re­ aleanmosses、alean mosses. ｇａｒｄｅｄａｓｔｂｅｍｏｓｔｒecentevolutionarystageofgarded as the most recent evolutionary stage of Recent，molecularphylogeneticanalysesfbrRecent, molecular phylogenetic analyses for theorderHypnales(seealsoTsubotaetall999；the order Hypnales (see also Tsubota et al. 1999; pleurocarpousmosseswerebasedonr6cLse-pleurocarpous mosses were based on rbcL se­ Bucketal2000）Subsequently,BuckandVittBuck et al. 2000). Subsequently, Buck and Vitt quencesalongwithotherdatasets,ｓｕｃｈａｓａｍＢ，quences along with other data sets, such as alpB, ￣ (1986)(1986)constluctedanotherclassificationfbrhyp- constructed another classification for hyp­ 伽Ｌ－Ｆａｎｄノアs４Ａｔｔｈｅｐｒｅｓｅｎｔｔｉｍｅ，tbere-IrnL-F and rps4. At the present time, the re­ noblyaleanmossesbasedonmorphologicalandnobryalean mosses based on morphological and sourcessourcesofr6cLsequencesarethemostcompre- of rbcL sequences are the most compre­ ecologicalecologicaldata、 data. hensivesamplingfbrthehypnaleanmosses・hensive sampling for the hypnalean mosses. Inrecentyears，phylogeneticrelationshipsIn recent years, phylogenetic relationships Alignmentofthe'6cLsequencesisalsoeasybe-Alignment of the rbcL sequences is also easy be­ withinthehypnobryaleanmosseshavebeenthewithin the hypnobryalean mosses have been the causecauseoftheirdistinctionasaprotein-codingre‐ of their distinction as a protein-coding re­ focusfbcusofmucbattentionatseveraltaxonomiclev- of much attention at several taxonomic lev­ giongioninthecodonpositions・Sqwepresenthere in the codon positions. So, we present here els(Arikawa＆Higuchil999,2002;BucketaLels (Arikawa & Higuchi 1999, 2002; Buck et al. pbylogeneticin化rencesfbrthethreeordersHyp-phylogenetic inferences for the three orders Hyp­ 2000；ＤｅＬｕｎａ＆coworkersl999，2000；Maeda2000; De Luna & coworkers 1999, 2000; Maeda nales，LeucodontalesandHookeriales，basedonnales, Leucodontales and Hookeriales, based on etetaL200qTsubota&coworkersl999,2000, al. 2000; Tsubota & coworkers 1999, 2000, themaximum-likelihood（ＭＬ）analysesofl81the maximum-likelihood (ML) analyses of 181 2001a,ｂ;VanderpoortenetaL2002)．Tberela-2001a, b; Vanderpoorten et al. 2002). The rela­ sequencessequencesofthechloroplastribulosel,5-bisphos- of the chloroplast ribulose 1,5-bisphos­ tionshipswithinthehypnobryaleanmosseshavetionships within the hypnobryalean mosses have phatephatecarboxylase/oxygenaselargesubunit carboxylase/oxygenase large subunit beenbeenevaluatedbybothmorphologyandmolecu- evaluated by both morphology and molecu­ (rbcL),(r6cL),togetherwithrecentinfbnnationonthe together with recent information on the larlarstudies(Hedeniisl995;ＤｅＬｕｎａｅｔａＬ１９９９)． studies (Hedeniis 1995; De Luna et al. 1999). molecularmolecularphylogenyofhypnobryaleanmosses， phylogeny of hypnobryalean mosses, Hedeniis（1995,1996)showedthemonophylyofHedeniis (1995, 1996) showed the monophyly of andanddiscusstheimplicationsofourfindingson discuss the implications of our findings on thetheHookeriales-Sematophyllaceaeanddiscov- Hookeriales-Sematophyllaceae and discov­ thetheHypnaceaeandtheirrelatedfamilies． Hypnaceae and their related families. erederedtheLeucodontalesasagradetaxonbasedon the Leucodontales as a grade taxon based on morphologica1data、DeLunaetaL(1999)clari-morphological data. De Luna et al. (1999) clari­ MaterialsandmethodsMaterials and methods fiedfledthehomologyofpleurocarpyandthemono- the homology of pleurocarpy and the mono­ phylyphylyofallhypnobryaleanpleurocarpousmosses of all hypnobryalean pleurocarpous mosses Phylogeneticanalysesbasedonther6cLgenePhylogenetic analyses based on the rbcL gene (the(theLeucodontales,HypnalesandHookeriales)， Leucodontales, Hypnales and Hookeriales), sequencessequenceswerebasicallyperfbnnedfbllowing were basically performed following basedbasedonacombined'6cLsequencesandmor- on a combined rbcL sequences and mor­ TsubotaTsubotaandcoworkers（1999,2000,2001a,ｂ)． and coworkers (1999, 2000, 2001 a, b). phologyphologydataset・Furtherdetailswererecently data set. Further details were recently TheThemethodconsistsoftwosteps：Dobtaining method consists of two steps: 1) obtaining の revealedrevealedbyDeLunaetaL(2000)andBucketal． by De Luna et al. (2000) and Buck et al. sequencesequencedata(DNAextractionPCRamplifica- data (DNA extraction, peR amplifica­ (2000)(2000)whoproposednewphylogeneticrelation- who proposed new phylogenetic relation­ tiontionandDNAsequencing,orDNAdatabaseho- and DNA sequencing, or DNA database ho­ shipsshipswithinthesepleurocarpsTheseworks within these pleurocarps. These works mologymologysearch);and２)dataanalyses． search); and 2) data analyses. suggestedsuggestedthattheLeucodontalesandHypnalesas that the Leucodontales and Hypnales as previouslypreviouslyunderstoodarenotmonophyletic understood are not monophyletic. Sources８℃"7calα"c/Ｐ/α"ｒＭＪＺｅｒｊａん and Plant Materials FamiliesFamiliesfroｍｔｈｅｓｅｔｗｏｏｒｄｅｒｓａｒｅｍｉｘｅｄｉｎａ from these two orders are mixed in a AAtotalofl81mossspecieswereexaminedfbr total of 181 moss species were examined for singleｓｉｎｇlelargecladesistertothehookerialeanlin- large clade sister to the hookerialean lin­ thetheanalysesasshowninTablelandinAppendix analyses as shown in Table 1 and in Appendix eage.eage、 A.A・Thelattercontainsvoucherinfbmationand The latter contains voucher information and TheTheobjectivesofthisstudyare:ｌ)toconstruct objectives of this study are: 1) to construct DNADNADatabaseAccessionNos・Voucherspeci- Database Accession Nos. Voucher speci­ aaphylogenetictreefbrhypnobryaleanmosses phylogenetic tree for hypnobryalean mosses mensmenssequencedinthepresentsmdyweredepos- sequenced in the present study were depos­ basedbasedonlarge-scaledataset;２)totestthemono- on large-scale data set; 2) to test the mono­ iteditedi、TNS,ＨＹｏ,orHIRoSomesequencesused in TNS, HYO, or HIRO. Some sequences used phylyphylyofthethreeorders（Hypnales,Leucodon- of the three orders (Hypnales, Leucodon- forfbrtheanalyseswerealsoobtainedfi･omthe the analyses were also obtained from the DNADNAdatabase． database. H.HIsuBoTA,Ｔ・ARIKAwA,Ｈ・AKlYAMA,ＥＤＥＬｕＮＡ,Ｄ､OoNzALEz,MHIGucHIANDH､DEGucHI TSUBOTA, T. ARlKAWA, H. AKIYAMA, E. DE LUNA, D. GONZALEZ, M. HIGUCHI AND H. DEGUCHI 647６４７

TableTableLListofspeciesandftlmiliesinvestigatedlbrl6cLgenewiththeaccessionnumberandvoucheror 1. List of species and families investigated for rbcL gene with the accession number and voucher or referencereferenceofthesequences、TreatmentoffamiliesinthistablefbllowsVitt（1984)ｗｉｔｈsomemodificationsby of the sequences. Treatment of families in this table follows Yitt (1984) with some modifications by BuckBuckandVitt(1986)．FurtherinfbnnationfbrthesequencesobtainedinthepresentsmdyisshowninAppendix and Yitt (1986). Further information for the sequences obtained in the present study is shown in Appendix A.Ａ、

OrderlFamily/SpeciesOrdelﾌﾟFamiIv/SDccies AccessionAccessionNo． No. References/OriginRefcrcnccs/Origin HypnalesHypnaleS(s・stⅨ） (s. str.) HypnodendraceaeHypnodendraceac HypnodendronA0lp"ocノセ"cjWmel2zjesjj(Hook.)Paris menziesii (Hook.) Paris AF23ＡＦ２３ＩＯ９３ 1093 MishlerMishIcrctal(onIyinDNA et a!. (only in DNA database)database） PleuroziopsidaceaePIeuroziopsidaceae Pleuroziopsis雄"mzjqpsな'wﾉﾉze"たα(Weinm.)Kindb・cxBritt、 rutheniea (Weinm.) Kindb. ex Britt. AB024683ＡＢＯ24683 ArikawaArikawaamdHiguchi(19”） and Higuchi (1999) FontinalaceaeFontinalaceac FontinalisFo"""α/jFampW℃"ｃｑＨｅｄｗ antipyretiea Hedw. AB050949ABO50949 TsubotamsubotaetaL(2001a） et al. (2001 a) ｡ FontinalisＦｂ"'ｊ"αﾉ応α"伽ﾉｱ'eZjcqHedw、 antipyretiea Hedw. AJ275AJ275183 183 CoxCoxctal(2000） et al. (2000) FontinalisＦｂ"/、αﾉﾉＳ“んcar/ｊｃａＢｒｕｃｈ＆Schimp・ daleearliea Bruch & Schimp. AF23AF231074 1074 DeDeLunaetaL(2000） Luna et al. (2000) Echinodiaceae EehinodiumEbﾉｸmodmll2〃"61℃s"'?z(Mitt.)Ａ・JaCgcr umbrosum (Mitt.)A.Jaeger AF233568 DeDcLunaetal(2000） Luna et al. (2000) SciaromiumSbmrommm〃jcosjcJm”(SulL)Mitt、 trieostatum (Sull.) Mitt. AB024677ABO24677 ArikawaArikawa＆Higuchi(1999） & Higuchi (1999) Thuidiaceae j6jerj"eﾉﾉﾋJa6jerj"α(Hedw.)Ｍ・FleischAbietinella abietina (Hedw.) M.Fleisch. AF005519ＡＦＯＯ５５１９ GoffinetGoffinctctal(1998） et al. (1998) Thuidium７１ﾉｶ､とﾉﾉ"ｍｃﾉｾﾉｊｃａｍｍｍ(Ｈｅｄｗ)Schimp、 delicatulum (Hedw.) Schimp. AF158177ＡＦＩ５８１７７ DcLunactaL(2000）De Luna et a!. (2000) Thuidium７１ｌｂ､ﾋﾉmmplな'ocaOMMUⅡ､HaL)AJacgcr pristocalyx (MilII.Hal.) A.Jaeger AB071416ＡＢＯ７１４Ｉ６ TsubotaTSubotactal(200ｌｂ） et al. (2001 b) Thuidiumnh"､１１"〃だcqg"""伽(Hedw.)Lindb． recognitum (Hedw.) Lindb. AB019476ＡＢＯ１９476 Akiyamactal.(2000;onlyAkiyama et a!. (2000; only inDNAdatabascXmsubotain DNA database); Tsubota etetal(2001a） al. (200Ia) LcskcaceaeLeskeaceae Bom〃αｍｊ"e"ﾉﾉ(Broth.)ＣａｒｄｏｔBoulaya millenii (Broth.) Cardot AB024963ＡＢＯ２４９６３ TsubotaTBubotactal(1999） et a!. (1999) Ｄｍ/tieﾉﾉﾋJSpecjosjFsjmqBrothexCardotDuthiella speciosissima Broth. ex Cardot ABOＡＢＯｌ９４６７ I 9467 Akiyamaetal.(2000;onlyAkiyama e! a!. (2000; only inDNAdatabasc);thiSstudyin DNA database); this study M〕′αbeαﾉﾘw"ＣｅﾉﾉﾋJ(Mitt.)BrothMiyabeafruticella (Mitt.) Broth. AB019475ＡＢＯｌ９４７５ MaedaMaedaetal(2000） et al. (2000) OkamuraeaOAcJm皿池ｅａｈａＡｏ"je"sjs(Mitt.)Broth hakoniensis (Mitt.) Broth. ABOl9477ＡＢＯＩ９４７７ MaedaMacdactal(2000） c! al. (2000) AmblystegiaceacAmblystegiaceae CalliergonellaCtJ"je堰o"eﾉﾉﾋJc"sp此ｍね(Hedw.)Loeske euspidata (Hedw.) Loeske AB024678ABO24678 Arikawa＆Higuchi(1999）Arikawa & Higuchi (1999) CratoneuronfilicinumC'、o"e"'℃"〃ｃｊ'､"(Hedw)Spruce (Hedw.) Spruce ABO95270AB095270 thisstudythis study D'℃pα"ocmdi`sadi`"c"s(Hcdw.)Warnst．Drepanocladus aduncus (Hedw.) Wamst. ABO24681AB02468I Arikawa＆Higuchi(1999）Arikawa & Higuchi (1999) ｆbｲ91℃α''161ｌＷｅｇｊ"〃'e'､x(Hedw.)JcnnHygroamblystegium tenax (Hedw.) Jenn. AF23356５AF233565 DeLunactal.(２０００）De Luna et aI. (2000) Lembophyllaceae Doﾉﾉcﾉｶo､"rjqpsjM1ve応施'ｍな(Mitt.)ＮｏｇDolichomitriopsis diversiformis (Mitt.) Nog. ABOI9465ABOl9465 Akiyamactal(2000;onlyAkiyama e! al. (2000; only inDNAdatabasc);thisstudyin DNA database); this study Ｌｅ"16qpﾉＭ/""'cﾉﾙ"/Sm"?(Hookf＆Wilson)Lindb，Lembophyllum divulsum (Hook.f. & Wilson) Lindb. AF233570 DeLunactal.(2000）De Luna et al. (2000) StcreophyIlaceacStereophyllaceae EmodO"'qpsハノＢＨＣＯＳ／２gα(Brid.)ＷＲＢｕｃｋ＆lrelandEntodontopsis leucostega (Brid.) W.R.Buck & Ireland ABO24635AB024635 Arikawa＆Higuchi(1999）Arikawa & Higuchi (1999) S1e花qpﾉ!)ﾉﾉﾉ"加沢adIczjﾉCs"、(Hook)MitLStereophyllum radiculosum (Hook.) Mitt. ABO24637AB024637 Arikawa＆Higuchi(1999）Arikawa & Higuchi (1999) Fabroniaceae A"αcα'Modb"spmch"o雄s(FroeLexBrid.)ＢｒｉｄAnacamptodon splachnoides (Froel. ex Brid.) Brid. AF231077AF23 1077 DeLunaetal.(2000）De Luna et al. (2000) 〃ﾋﾞﾉﾉcodO"""maJpjﾉﾉﾋJ”(Hcdw)AJaegerHelicodontium capillare (Hedw.) AJaeger AF23357］AF233571 DeLunaetal.(2000）De Luna et al. (2000) BrachythcciaccacBrachytheciaceae Bmc〃んecj"ｍｐ肋､Cs"、(Hedw.)Schimp、Brachytheciumplumosum (Hedw.) Schimp. AF233566 DeLunactaL(2000）Dc Luna et al. (2000) Ｂｍｃﾉi〕ﾉﾉﾙecj"mrjw`ﾉα”ＳｃｈｉｍｐBrachythecium rivulare Schimp. ABO24674AB024674 A｢ikawa＆Higuchi(1999）Arikawa & Higuchi (1999) Ｂｍｃ/Ｍｈｅｃｍｍｓα/e6ms"、(HofYinexRWCber＆Brachythecium salebrosum (Hoffm. ex F. Weber & AFl58I76AFI58176 DeLunaetal(2000）De Luna et al. (2000) DMohr)Schimp・D.Mohr) Schimp. ノMb'Ｈｍｃﾉａｄｔｗｃｍｍｏｗｊｃｚｊｊ(Borszcz.)Stcere＆ＳｃｈｏｆMyuroclada maximowiczii (Borszcz.) Steere & Scho[ ABO29389AB029389 Tsubotactal.（1999）Tsubota et al. (1999) Ｐﾉﾛﾘﾉ!〕や"jdm加吻α"o/“s(Hedw.)Dix，Platyhypnidium riparioides (Hedw.) Dix. ABO29385AB029385 Tmbotactal(1999）Tsubota et al. (1999) PseWdOsc化mpodmmpⅢ'ｗｍ(Hcdw.)Ｍ・F1cisch．Pseudoscleropodium purum (Hedw.) M.Fleisch. AF233567 DCLunaCtal(2000）Dc Luna ct al. (2000) Ｒ/i)′"c/zos/egm"'pαﾉﾉﾉ｡!/bﾉﾉz`"1(Mitt.)AJacgcrRhynchostegium pallidifolium (Mitt.) AJaeger ABO24944AB024944 Tsubotactal(1999）Tsubota ct aI. (1999) ７bme"卯""〃〃/花"s(Hedw.)LoeskeTomentypnum nitens (Hedw.) Loeske ABO24676AB024676 ArikawaandHiguchi(1999）Arikawa and Higuchi (1999) EntodontaccaeEntodontaceae E7Troc！O〃cﾉｶαﾉﾉelZgelj(Paris)CardotEntodon challengeri (Paris) Cardot ABO50993AB050993 TBubotaetal.(2001a）Tsubota e! al. (2001 a) E)?/odb"/”ｊｍ４ｓ(Gri征)AJacgcrEntodon luridus (Griff.) AJaegcr ABO50994AB050994 T1subotactaL(2001a）Tsubota ct al. (2001 a) 勘/odO""!)'”“(Hook)HampeEntodon myurus (Hook.) Hampe ABO24640AB024640 Arikawa＆Higuchi(1999）Arikawa & Higuchi (1999) EmodO"'w6jc""c/bｲs(Mitt.)AJacger＆Sauerb・Entodon rubicundus (Mitt.) AJaeger & Sauerb. ABO29386AB029386 TsubotaT1subotaetal(1999） et al. (1999) EmodO〃sca61j血"sLindbEntodon scabridens Lindb. ABO50995AB050995 TEubotaetal(200]a）Tsubota et al. (2001 a) SematophyllaccaeSematophyllaceae 化α"'んoW2〕''７chjzmzpqpj/、'"、(Ilarv.)Ｍ・FleischAcanthorrhynchium papillatum (Harv.) M.Fleisch. ＡＢＯ５１２２４AB051224 辰msubotactal(2001a）Tsubota et al. (2001 a) 化mpo"zｲｍｐｚｍｇｅ"s(Hedw.)BrothAcroporium pungens (Hedw.) Broth. ＡＦ２３３５７２AF233572 DcLunaetal(2000）De Luna et al. (2000) jcmporm〃s"α､j"e瓦加(Reinw.＆Ｈｏｍｓｃｈ)Ｍ・Flcisch．Acroporium stramineum (Reinw. & Homsch.) M.Fleisch. ＡＢＯ５Ｉ２２５AB051225 T1subotactal(2001a）Tsubota et aI. (200Ia) 648 HikobiaＨｉｋｏｂｉａＶｏＬｌ３,No.４，２００２ Vol. 13, No.4, 2002

TableTableLcontinued 1. continued.

OrderlFamily/SpeciesOrderﾌﾟFamilv/SDccies AccessionAccessionNo． No. References/OriginRcfercnccs/ｏｒ 、

Aptychella幼ＩＤ'cﾉｶeﾉﾉZzgﾉomelmqpmpqg"ﾉ娩'α(Tbyama)Ｓｃｋｉ glomeralopropagulifera (Toyama) Seki ABOSI217ＡＢＯ５Ｉ２Ｉ７ TsubotaTmbotactaL(200Ia et aL (200 I a) Helerophyllium｣5ﾉｾﾉempﾉﾘﾉﾉﾉﾉ""1〃"e(Hook.)Ｍ・F1cisch affine (Hook.) M. Fleisch. ABOSI218ＡＢＯ５１２１８ TsubotaT3ubotaetaL(2001a et aL (200 I a) Helerophylliumノ5leZemp/q〕ﾉﾉﾉﾉⅢ、〃e"Taros皿"ZBrothcxPLdelaVardc＆Th6m nemalosum Broth. ex P.de la Yarde & Ther. AB02939IＡＢＯ２９３９１ TsubotaT1subotactaL（1999） et aL (1999) IsocladiellaKsocﾉUdjeﾉﾉﾋ７３脚”"ﾉﾋJrな(Dixon)ＢＣ・T1an＆Mohamcd surcularis (Dixon) B.C.Tan & Mohamed AB039784ＡＢＯ３９７８４[asⅣｅａｃｍ‐ [as Neacro­ TsubotaT1subotactal(2000） ct aL (2000) poriumporm"Z加ｇＥﾉﾉ(/を'W"?］ flagelliferum] MaslopomaMtzsjqpo"zap皿ﾉCheﾉﾉﾋJ(Herzog)incd pulchella (Herzog) ined. AB071410ABO71410[asms- [as Tris­ TsubotaTsubotactal.(200lb） et aL (200 I b) megisliamegjsljap"ﾉb〃eﾉﾉﾋJ］ pulchella] MaslopomaMJslqpollzas皿lWZ/blwllzHorik＆Ａｎｄｏ subfiliferum Horik. & Ando AB071411ＡＢＯ７Ｉ４Ｉｌ TsubotaT1subotactaI.(200lb） et aL (200 I b) MaslopomaﾉMJS'qpo"Tα皿"cj"(/bﾉ､"'(Broth)Broth uncinifolium (Broth.) Broth. AB071412ＡＢＯ７１４１２ TsubotaT1subotactaI.(200lb） el aL (200Ib) MeiolheciumMbjoﾉﾉiecj"mmJcrocα'p"、(Hook)Mitt． microcarpum (Hook.) Mitt. AB051223ＡＢＯ５Ｉ２２３ TsubolaTmbotactal(2001a） ct aL (200 I a) PapillidiopsisＰ(Jpj"jdiqpsなｍａｃｍｓＺｊａｑ(Broth.＆Paris)Ｗ,RBuck＆ macrosticta (Broth. & Paris) WR.Buck & AB051220ＡＢＯ５１２２０[asRhqpﾙﾉﾋﾉO‐ [as Rhaphido­ TsubotaTmbotactal.(2001a） et aL (200 I a) B.C.TanＢ・Ｃｍｌａｎ slichums'ｊｃ/ｌｗｍｍａｃ'･Cs"c/"､］ macrosliclum] Pseudolrismegislia応e"ｄｏ"jm7TegjSjjaⅢ"｡iｲﾉ、α(Broth＆Ｍ・Yasuda)HAkya． undulala (Broth. & M.Yasuda) H.Akya. AB051229ABO51229[asnjs- [as Tris­ TsubolaTmbotactal.(2001a） el aL (2001a) ロ &＆Tsubota Tsubota megisliamegjs"α皿"｡！`ﾉ、α］ undulala] PylaisiadelphaDﾉﾋJな/αｄｂｌｐﾉiQco"qpm"α'α(Reimers＆Sakurai） complanala (Reimers & Sakurai) AB03978SABO39785[ａｓＢｍ‐ [as Bro­ TsubotaT1subotactaI.(2000） et aL (2000) W.R.BuckＷＲＢｕｃｋ Iherellaﾉﾉ『e”ぬＣＯ"2p/α"α`α］ complanala] Pylaisiadelphafaurieiｌｌ)ﾉﾉ､､α`/blpﾉlα/2mrjej(BcschcxCardot)ＷＲＢｕｃｋ (Besch. ex Cardot) WR.Buck AB039786ABO39786[ａｓＢｍ‐ [as Bro­ TsubotaTmbotactal.(2000） el aL (2000)

Iherellaﾉﾉze”ﾉﾉα/Zmrjej］ fauriei] 画 けﾉﾋ』な､｡b(りんαﾉｶe"o"〃(Duby)ＷＲＢｕｃｋPylaisiadelpha henonii (Duby) WR.Buck AB029167[asBro‐AB029167 [as Bro­ TsubotamSubotaetal.（1999） et aL (1999) Iherella/ｈｅ肥/ね/ie"o"ﾉﾉ］ henonii] PylaisiadelphaＲｙﾉﾋJjSmde

Ｒﾉﾘﾉﾉﾉ“α`、ＩＤﾉmMZpo"ｊｃⅢs(Reimcrs)Ｔｌ・ＫｏｐRhylidiadelphusjaponicus (Reimers) ·U.Kop. ABO39788AB039788 msubotaetaL(2000）Tsubota et aL (2000) □ Ｒ/Mjdm火ｌＩｐｈ"sﾉo1℃"s(Hcdw.)Wamst．RhYlidiadelphus loreus (Hedw.) Warnst. ABO24666A B024666 Arikawa＆Higuchi(1999）Arikawa & Higuchi (1999) Ｒ/Mjdm“lIpﾉiWss9"qrmmJs(Hcdw.)Warnst．RhYlidiadelphus squarrosus (Hedw.) Warnst. ABO24667AB024667 Arikawa＆lliguchi(1999）Arikawa & Higuchi (1999) Ｒ/Mjdmdblph"s〃""eﾉﾉWS(Hedw)Wamst・Rhylidiadelphus Iriquelrus (I-Iedw.) Warnst. ABO24668AB024668 Arikawa＆HiguchiO999）Arikawa & Higuchi (1999) PlagiothcciaccaePlagiotheciaceae LsqpZeﾉﾌｲgjqpsjS"ｍｅ比ｒｊα"ａ(Schimp)Ｚ・Iwats・lsoplerygiopsis muelleriana (Schimp.) Z.lwats. ＡＢｌ３４９４２AB134942 Arikawa＆lliguchi(2002）Arikawa & Higuchi (2002) Pmgjo'Aecj"、。ｂ"'た"/、"、(Hedw)Schimp．Plagiolhecium denliculalum (Hedw.) Schimp. ＡＢＯ２４６２３AB024623 ATikawa＆Higuchi(1999）Arikawa & Higuchi (1999) Ｐ/tJgjorAecj"、加ｿ/o"ﾉﾉ(SulL)ＥＢ・BartramPlagiothecium draylonii (Sull.) E.B.Bartram ＡＢＯ２４６２５AB02462S Arikawa＆Higuchi(l”，）Arikawa & Higuchi (1999) PﾉﾋZgjo/ﾉiec/""Te"刷りﾉ！〕ﾉﾉﾉⅢ"'(Cardot＆Th6Ⅸ)Ｚ・IwatsPlagiolhecium euryphyllum (Cardot & Ther.) Z.lwats. ＡＢO24626AB024626 Arikawa＆lliguchi(1999）Arikawa & Higuchi (1999) P/cJgjo/Aecj冴加〃ecApem雄"ｍＢｒｕｃｈ＆ＳｃｈｉｍｐPlagiolhecium neckeroideum Bruch & Schimp. ABO24630AB024630 Arikawa＆Higuchi(1999）Arikawa & Higuchi (1999) ＰﾉﾋJgjoﾉﾉiecj"ｍ〃emoraﾉe(MiU.)ＡＪａｅｇｅｒPlagiolhecium nemorale (Mitt.) AJaeger ABO29387AB029387 msubotaCtaI.（]999）Tsubota et aL (1999) ＰﾉﾋｺgjofAecjw”〃"｡）ｲﾉα'"、(Hcdw.)Schimp・Plagiolhecium undulatum (Hedw.) Schimp. ABO24634AB024634 Arikawa＆lliguchi(l”9）Arikawa & lliguchi (1999) Hypnaceae Cre"域"ｍｍｏﾉﾉ皿Sc皿"2(Ｈｅｄｗ)Mitt．Cienidium molluscum (Hedw.) Mitt. ABO24657AB0246S7 Arikawa＆Iliguchi(1999）Arikawa & Higuchi (1999) Ｏﾉossadblpﾉmsqgα/αeBroth.＆Ｍ・YasudaGlossadelphus ogalae Broth. & M. Yasuda ABO50950AB050950 T1subotactaI.(2001a）Tsubola et aL (200 I a) OoﾉﾉﾋJ"ｍｍｇｌ"Csα(Mitt.)BrothGollania ruginosa (Mitt.) Broth. ABO94341AB09434I thisstudythis study Go"α"mSp/e"ぬ"s(Iisiba)Nog．Gollania splendens (Iisiba) Nog. ABO94340AB094340 thisstudythis sludy 〃C吃qgjeﾉﾉﾋJpel．'･o6"s/α(Broth.)ＺＪｗａｔｓ・Herzogiella perrobusla (Broth.) Z.lwats. ABl34944AB134944 Arikawa＆Higuchi(2002）Arikawa & Higuchi (2002) ﾉﾌﾞＤＰ""mcz《p1℃sWbrmeHedw．Hypnum cupressiforme Hedw. ABO39674AB039674 T1Subotactal.(2000）Tsubola et aL (2000) H.HTsuBoTA,Ｔ・ARIKAwA,Ｈ・ＡＫⅣAMA,ＥＤＥＬｕＮＡ,Ｄ・GoNzALEz,Ｍ・HIGucHlANDH・DEGucHI６４９ TSUBOTA, T. ARIKAWA, H. AKIYAMA, E. DE LUNA, D. GONZALEZ, M. HIGUCHI AND H. DEGUCHI 649

TableTableLcontinued I. continued.

Order/Family/SpeciesOrder/Familv/SDecies AccessionAccessionNo． No. References/OriginRefbrcnces/Ｏｒｉ 1， Hypnumhbp"皿、/ｍｄｂｅ喧jjMitt、 lindbergii Mitt. AB029390ABO29390 TsubotaTmbotaetaL（1999） et al. (1999) HypnumノカplT"”ｐ此'，Ｔａｇ/bmzeWilson plumaeforme Wilson AB029384ABO29384 TsubotaTmbotactal.(1999） et al. (1999) Hypnumノh0p"""T〃jS'０－Wr雄(Broth)ＰaⅢ tristo-viride (Broth.) Par. AB024656ABO24656 TsubotaTmbotactaL（1999） et al. (1999) /sopterygiumZSqpteﾚﾂ9ｍｍに"er皿加(Swb)Mitt・ tenerum (Sw.) Mitt. AF233569AF2335６９ DeDcLunaetaL(2000） Luna et al. (2000) /sopterygiumLsqp/Clﾂｇｊ"碗Ｗ"eαノセＥＢ・Bartram vineale E.B.Bartram AB024650ABO24650 ArikawaArikawa＆Higuchi(1999） & Higuchi (1999) OrthotheciumOr/ﾉｶo肋ecj"ｍｒｚ(/bsce"s(Dicks・ｅｘＢｒｉｄ.)Schimp rufescens (Dicks. ex Brid.) Schimp. AB05095IABO50951 TsubotaT1subotactal.(ZOOla） et al. (200 I a) PylaisiaDﾉｍＳｊａｍ"にα'α(Hedwb)Schimp intricata (Hedw.) Schimp. AB024642ABO24642[asＥ〕ﾉﾉａｊ－ [as Pylai­ ArikawaArikawa＆Higuchi(1999） & Higuchi (1999) siellaｓｊｅ"αｊ""にα/α］ intricata] PlatygyriumPmOlgﾌﾉｒｊ皿加'"e"s(Brid.)Schimp repens (Brid.) Schimp. AB024645ABO24645[ａＭ〕ﾉﾉαﾉｰ [as Pylai­ ArikawaArikawa＆Higuchi(1999） & Higuchi (1999) siellasje"αpol()ﾉα"ﾉﾉ､］ polyanlha] .mxjp"y"""zao"DC"e"se(Besch)Ｚｊｗａts・ Taxiphyllum aomoriense (Besch.) Z.lwats. AB024648ＡＢＯ24648 ArikawaArikawa＆Higuchi(1999） & Higuchi (1999) Leu;:odontalesLeucodontales ClimaciaceaeClimaciaccac 、 Climaciumdｊ〃αｃｍｍｃｊｂ"｡、雄s(Hedw.)FWCber＆Ｄ・Ｍｏｈｒ dendroides (Hedw.) F.Weber & D.Mohr ABOＡＢＯｌ９４４２ 19442 AkiyamaAkiyamaetal.(200qonly et al. (2000; only ininDNAdalabase);thisstudy DNA database); this study Climaciumjaponicumｄｊｍｑｃｊ"、ﾉﾋZpo"ｊｃ"mLindb Lindb. ABOＡＢＯ１９４４３ I 9443 AkiyamaAkiyamaetaL(2000;onIy et al. (2000; only ininDNAdatabasc);thisstudy DNA database); this study NeckeraceaeNeckcrace2e BissetiaBおselm〃"gz`/α'α(Mitt.)Broth． lingulata (Mitt.) Broth. AB094789ABO94789 thisthisstudy study Homaliadelphus〃o"､"α“"ﾉｶⅢs'αrgjo"､""s(Mitt.)Dixon＆Ｐｄｅｌａ targionianus (Mitt.) Dixon & P. de la AB094792ABO94792 thisthisstudy study YardeＶａｒｄｅ Homaliodendron〃O"､ﾉﾉo`ノセ"鋤｡"Scα(peﾉﾉﾋﾞ/bﾉﾉ"、(Mitt.)Ｍ・FlCisch scalpellifolium (Mitt.) M.Fleisch. AB094788ＡＢＯ９４７８８ thisthissmdy study NeckEraノVCcAcm”"lgcmMijll､HaI、 urnigera MUII.Hal. AFI58173ＡＦｌ５８１７３ MaedaMaedactal(2000） et al. (2000) NeckEropsisノVECにrqpsn〃"jZmｍ(Mitt.)Ｍ・Fleisch nitidula (Mitt.) M. Fleisch. AB094790ＡＢＯ９４７９０ thisthisstudy study PinnatellaPm"α花ﾉﾉﾋJα"6唖α(Bosch＆SandcLac.)Ｍ､Fleisch ambigua (Bosch & Sande Lac.) M.Fleisch. AB094787ＡＢO94787 thisthisstudy study Thamnobryum７１ｈａｍ"o6ひ"ｍｍ６ｓｅｒｊａｍｍ(Mitt・exSandeLac)ＢＣＴaｎ subseriatum (Mitt. ex Sande Lac.) B.C.Tan AB094791ABO94791［as71/mmlzo‐ [as Thamno­ thisthissmdy study bryumｂひＨｍｓα"生ｊ］ sandei] Cyrtopodaceae BescherelliaBescﾉbe1℃"ｊａ６１ｗｌ/b/jaHampc brevifolia Hampe AJ275184ＡＪ２７５１８４ CoxCoxetaI.(2000） et al. (2000) BescherelliaBesc/，e”"jaeにg[J"‘jSsjmaDuby elegantissima Duby AF231097ＡＦ２３ＩＯ９７ DeDCLunactal.(2000） Luna et al. (2000) CyrtopusQ′"qp"sseros"s(Hedw.)Hookf setosus (/-Iedw.) /-Iook.f. AF231096ＡＦ２３１０９６ MishlcrctaL(onlyinDNAMishler et al. (only in DNA database)databasc） CryphaeaceaeCIyphacaccae CryphaeaQ)p"αｅａｓｊ"e"３曲Ｅ・BBartram sinensis E.B.Bartram ＡＢＯｌ９４５７ABOl9457 Macdactal(2000）Maeda el al. (2000) Cyptodontopsisqpmdm/qps町o6'"s(/b/ね(Nog.)Ｎｏｇ obtusifolia (Nog.) Nog. ＡＢＯｌ９４５８AB019458 Macdactal.(2000）Maeda el al. (2000) Ｄｏ｡’【J/apo"jcaSandeLacDozyajaponica Sande Lac. ＡＢＯｌ９４４６ABO I 9446 Maedactal.(2000）Maeda el al. (2000) Pj/O"jcﾉｶqpsjSdb"【α'α(Mitt.)BeschPilotrichopsis dentata (Mitt.) Besch. ＡＢＯｌ９４6０ABO 19460 MacdactaL(2000）Maeda et al. (2000) AnomodontaccacAnomodontaceae A"omodO〃α66死ＷａｍｓＭｉｔｔ．Anomodon abbreviatus Mitt. ＡＢＯｌ９４６８ABO I 9468 Macdactal(2000）Maeda et al. (2000) 」"omodO〃ｇｊｍ“/MUll・ＨａＬAnomodon giraldii MUII.J-Ial. ＡＢＯｌ９４６９ABO I 9469 Macdactal(2000）Maeda et al. (2000) Ａ"omodb"〃ｊ"oパHedw.)ＬｉｎｄｂAnomodon minor (Hedw.) Lindb. ＡＢＯＩ９４７ＩABOl9471 Macdactal(2000）Maeda et al. (2000) Ａ"omodb〃'wgE/〃(MiilLHaL)KeissLAnomodon rugelii (MUII.Hal.) Keissl. ＡＢＯl９４７０AB019470 Macdaetal.(2000）Maeda el al. (2000) ﾉｦﾛp/oﾉﾛ〕′ｍｅ"ｊ"腕ﾉolZgjlTerve(Broth)BrothHaplohymenium longinerve (Broth.) Broth. ＡＢＯｌ９４７２ABOI9472 Macdactal.(2000）Maeda et al. (2000) 〃ZJpﾉo/ｌ〕ﾉﾉ，Telzj"'Ｗ,semb"jS'e(MiilLHaL)Broth．Haplohymenium pseudotriste (MUll.Hal.) Broth. ＡＢＯｌ９４７３ABO 19473 Maedactal.(2000）Maeda et al. (2000) 〃ど'1pe""e"、〃roccoae(Sull.＆Lesq.)CardotHerpetineuron toccoae (Sull. & Lesq.) Cardot ＡＢＯｌ９４７４AB019474 MacdaetaL(2000）Maeda et al. (2000) Hedwigiaceae h'bclWjgjZzc"jα'α(Hcdw)PLBeauMHedwigia ciliata (Iledw.) P.Beauv. AFOO55I7AF005517 GoffinctctaL(1998）Goffinet el al. (1998) ノWiacocαノア“ｐ"'p"、Sce"s(Brid)ParisRhacocarpus purpurascens (Brid.) Paris AJ27SI71AJ275171 Coxctal.(2000）Cox el al. (2000) Leucodontaceae A"""jc"、/brmosα"αＮｏｇAntitrichiaformosana Nog. ＡＢＯＩ９４４５AB019445 MaedaetaM2000）Maeda et al. (2000) 化/仰o"eaes9mmﾉﾉﾉ(Tb6r）Ｈ・Akiy・Felipponea esquirolii (Ther.) H.Akiy. ＡＢＯＩ９４４７AB019447 MacdactaM2000）Maeda et al. (2000) ＦＤ“"Ｄｅｍｊα/とIpo"ｊｃａ(Besch)ParisForsstroemiajaponica (Besch.) Paris ＡＢＯＩ９４５０ABOl9450 Macdactal(2000）Maeda et al. (2000) FbハsZroemJα〃ecAeroj火sBrothForsstroemia neckEroides Broth. ＡＢＯI９４４９AB019449 MacdactaL(2000）Maeda et al. (2000) FblIsstmellnm〃jchollzj"ね(Hcdw)Lindb．Forsstroemia trichomitria (Hedw.) Lindb. ＡＢＯｌ９４４８AB019448 Macdactal(2000）Maeda et al. (2000) Ｌe"ＣＯ血〃α'”v"℃"ｓＮｏｇ・Leucodon atrovirens Nog. ＡＢＯｌ９４５３AB019453 Macdactal.(２０００）Maeda et al. (2000) Ｌｅ"ＣＯ巾"/"ﾉﾋIce"s(Hedw）ＳｕｌＬLeucodonjulaceus (Hedw.) Sullo ＡＦ２３ＩＯ７５AF23 1075 MishlerctaL(onIyinDNAMishler el al. (only in DNA databasc）database) Ｌｅ"ＣＯ“〃mppo"ｉｃ"sNog．Leucodon nipponicus Nog. ＡＢＯｌ９４５ＩAB019451 MaedactaL(2000）Maeda el al. (2000) Ｌe"COC/b〃ＳＣＷ,olwTsなＢｅsch・Leucodon sapporensis Besch. ＡＢＯｌ９４５２ABO 19452 Maedaetal(2000）Maeda et al. (2000) Ｌｅ"codb"３ｃｍ”jdbs(Hedw.)Schwii伊Leucodon sciuroides (Hedw.) Schwagr. ＡＢＯ９５９８８AB095988 thisstudythis study Ｌｅ"codb"secW"ぬs(Harv.)Mitt・Leucodon secundus (Harv.) Mitt. ＡＢＯl９４５４AB019454 Macdactal.(2000）Maeda el al. (2000) LeHcodb〃ｓｏﾉicZ〕ﾉａＡｊｅ"s応ILAkiy．Leucodon sohayakiensis I-I.Akiy. ＡＢＯ１９４５５AB019455 Maedactal.(2000）Maeda el al. (2000) Ｌe"ＣＯ“〃re"lpemrzJsH,AkiybLeucodon temperatus H.Akiy. ＡＢＯｌ９４５６ABOl9456 MacdactaI.(2000）Maeda el al. (2000) PtychomniaccaePtychomniaceae PO’c"ｏｍ"ｉｏ〃αcjcHﾉﾋzだ(Brid)Mitt、Ptychomnion aciculare (Brid.) Milt. ＡＦ２３３５７６AF233576 DcLunaetaL(2000De Luna et al. (2000) MyuriaccaeMyuriaceae Emy”ｍｍｓｍｊｃⅢ、(Mitt.)ＮＣ９．Eumyurium sinicum (Mitt.) Nog. ＡＢＯｌ９４６３AB019463 Akiyamaetal(ZOOO5onlyAkiyama el al. (2000; only inDNAdatabase);thisstudyin DNA database); this sludy ＭｳﾉHrjⅢｍｈｏｃﾉzsje"erj(Schimp.)KindbMyurium hochslelleri (Schimp.) Kindb. ＡＦ２３３５７５AF233575 DcLunactaL(２０００）De Luna et al. (2000) 650 HikobiaＨｉｋｏｂｉａＶｏＬｌ３,No.４，２００２ Vol. 13, No.4, 2002

TableTableLcontinued 1. continued.

OrderlFamily/SpeciesOrder/FamiIv/SDccies AccessionAccessionNo． No. References/OriginRcferCnccs/Origin Wardiaceae Wardiaﾚｲblz/ねﾉﾘlgl･ome〃jcaHarv.＆Hook hygrometriea Harv. & Hook. AJ275ＡＪ２７５Ｉ７０ 170 CoxCoxetaL(2000） et al. (2000) Prionodontaceae P"o"oｄｏ"火"s"s(SwbexHedw.)Mijll・Ｈａ１．Prionodon densus (Sw. ex Hedw.) Miill.Hal. AFI58174ＡＦｌ５８１７４ DcLunaetaL(2000）De Luna et al. (2000) Taiwanobryum７ZJjwα"o6ひ"mspecJoszJmNog・ speeiosum Nog. ABOl9466ＡＢＯ１９４６６ AkiyamactaL(2000;onlyAkiyama et al. (2000; only inDNAdatabasc);thisstudyin DNA database); this study PterobryaceaePtcroblyaccae P'em6ひo〃qr6Wsc"mMitt・Pterobryon arbuseula Mitt. ＡＢＯｌ９４６１ABOl9461 MaedaMacdactaI.(2000） et al. (2000) Prem6ひo〃。b"sWmHornschPterobryon densum Homsch. AFI58175ＡＦｌ５８１７５ DcLunaetal(2000）De Luna et al. (2000) P/ero6ひqpsjlForje"'αﾉﾉSsubspj′Ｈ２""α"e"s心(Broth)ＮｏｇPterobryopsis orientalis subsp. yuennanensis (Broth.) Nog. ＡＢＯ１９４６２ABOl9462 MaedaMacdactal(2000） et al. (2000) TraehylomaｍＪｃﾉＭｏｍａｊ"ぬczJmMitt・ indieum Mitt. ＡＢＯ１９４6４ABO 19464 AkiyamaAkiyamactaI.(Z00qonIy et al. (2000; only inDNAdatabasc);thissmdyin DNA database); this study TrachypodaceaeTrachypodaceaｅ

Traehypodopsis７》zzc/1)podbps応α皿"cヅノ､α(Mitt)Ｍ・F1cisch aurieulata (Mitt.) M.Flcisch. AB024682ABO24682 ArikawaArikawa＆lliguchi(]999） & lliguchi (1999) □ Traehypus７》zJcﾉＷ,Ⅸｓ６ｊｃｏﾉorRCinw.＆Homsch bieolor Reinw. & Hornsch. AF233577 DeDeLunactaL(2000） Luna et al. (2000) MeteoriaceaeMcteoriaccac P[Zpj"αrjZJcノピppej(HornschexMUlLHaI.)AJaegerPapillaria deppei (Hornsch. ex MiilI.Hal.) A.Jaeger AF158172ＡＦｌ５８Ｉ７２ DeDeLunaetal.(2000） Luna et al. (2000) Hookeriales DaltoniaccaeDaltoniaccac qＰ Lepj［わ〃/ｍｍｓ"ｒｊ"αme"seMijⅡ､Hal､Lepidopilum surinamense Miill.Hal. AF233578ＡＦ２３３５７８ DeDcLunactal(2000） Luna et al. (2000) Hookeriaceae 〃OoAerjaacm/bﾉﾉαＨｏｏｋ＆Orev、Hookeria aeutifolia Hook. & Grev. AFI58170ＡＦｌ５８１７０ DeDcLunactaI.(2000） Luna et al. (2000) HypoptcD/giaccacHypopterygiaceae ノbpqpje奴gjHmZamarなcj(Sw.)Ｂ｢id・exMUlI・Hal、Hypopterygium tamarisei (Sw.) Brid. ex Miill.Hal. AFI58171ＡＦｌ５８１７１ DcLunactal(2000）De Luna et al. (2000) ハ0ｨpqp'Cl〕ｌｇｊ"〃/α/zj/e"seAngstrOmHypopterygium tahitense Angstr6m AF23ＡＦ２３１０９５ 1095 MishleretaI.(onIyinDNAMishler et al. (only in DNA database)databasc） BryalesBryaIes BryaceaeBIyaceae MJeﾉﾉcﾉzAq/brjaeﾉolZgzJ'α(Hoppc＆Ｈｏｍｓｃｈ)Nccs＆Mieliehhoferia elongata (Hoppe & Hornsch.) Nees & AF232693 CoxCoxctaI.(2000） et al. (2000) Hornsch.Hornsch OrthodontiumO〃ﾉｍｄｂ"""〃ﾉj"eα”Schwiigr・ lineare Schwagr. AJ275174ＡＪ２７５Ｉ７４ CoxCoxctaL(2000） et al. (2000) ＰＣｶﾉmclwcﾉﾋJ(Hedw.)LindbPohlia eruda (Hedw.) Lindb. N275175ＡＪ２７５１７５ CoxCoxctal(2000） et al. (2000) LcptostomataccacLeptostomataceac LepmsjomＷｍｍａｃｍｃα'1p"、(Hcdw.)Bach・Pyl、Leptostomum maeroearpum (Hedw.) Bach.Pyl. AJ275178Ａｊ２７５１７８ CoxCoxctaL(2000） et al. (2000) MniaccacMniaeeae A化j""ﾉﾉjomso"ｊｊＳｃｈｉｍｐMnium thomsonii Schimp. AF005518AFOO55I8 GoffinetOofYinclctaI.（1998） et al. (1998) Ｐｍｇｊｏｍ"ｊ"、cⅢSpj血/"ｍ(I-ledw.)1XLKop・Plagiomnium euspidatum (Hedw.) T.1.Kop. U87082U87082lasM"ｊ"m [as Mnium LewisLewisctal（1997） et al. (1997) cuspidatumcWqpjdbm'?？］1 Ｐｍｇｊｏｍ"ﾉﾂﾞﾉ，，/ZJpo"jc"、(Lindb)1Ｊ・Kop・Plagiomniumjaponieum (Lindb.) ·U.Kop. AB050992ＡＢＯ50992 TsubotaT1subotactaL(2001a） et al. (200 I a) RhizogoniaceaeRhizogoniaccac Bw･ﾉio6〃""2Ｖαﾉﾉﾊｰｇｍｒｍｅ(HampccxMUlLHaL)ManuclPyrrhobryum vallis-gratiae (Hampe ex MiilI.Hal.) Manuel AJ275179ＡＪ２７５１７９ CoxCoxetaI.(2000） et al. (2000) BartramBartramiaccac iaceac Bar"αｍｊａＰｏｌｍ/brmjSHedWBartramia pomiformis Hedw. AB024620ABO24620 ArikawaArikawa＆Higuchi(1999） & Higuchi (1999) Lejomeﾉｨz6ar"αmjo此尤s(Hook.)ＰａｒｉｓLeiomela bartramioides (I·look.) Paris AF478238 Magombo(2002）Magombo (2002) Ｐﾉﾉﾉﾉo"o油α"ぬ"α(Mitt.)Ａ・JacgcrPhi/onoUs andina (Mitt.) A.Jacgcr AF478240 MagomboMagombo(2002） (2002) AulacomniaceaeAulacomniaccac AulaeomniumA"ﾉacom"j皿、！zｨ垣竝《"(WahIcnb)Schwiigr・ turgidum (Wahlenb.) Schwagr. AJ275ＡＪ２７５１８０ 180 CoxCoxctal(2000） et al. (2000) RaeopilaceaeRacopiIaceac RaeopilumRacqpj/ｌＪｍｃｏｍ'０ﾉⅢ/αcezｨ"(ＭＵⅡ､llaL)Rcichardt convolutaeeum (Miill.llal.) Reichardt AF23ＡＦ２３１０９４ 1094 Mishlcrctal(onlyinDNAMishler et al. (only in DNA database)database）

OutgroupOutgrouptaxa taxa 、 ErpodiaceaeErpodiaccac VenturiellaI'b"'2J"eﾉﾉﾋJsj"elTsLs(Vent.)MUllHal sinensis (Vent.) Miill.Hal. AF005546ＡＦＯＯ５５４６ GoffinetGofYinctctal（1998） et al. (1998) SplachnaceaeSpIachnaccac Tayloria７h〕ﾉﾉoljα〃"９２`/α/α(Dicks.)Lindb lingulata (Dicks.) Lindb. AF005515ＡＦＯＯ５５１５ GoffinetGofTinctctal.（1998） ct al. (1998) OrthotrichaceacOrthotrichaccac DrummondiaD'ｗｍｍｏ"`地０６'2Ｗｂ/jaMUllllaL obtusifolia Miill.Hal. AF232697 CoxＣｏｘｃｔａｌ(2000） et al. (2000) ノＭｒ"ｍｊ"j"Ｗ"c"rv

ＤＭ欧伽crjo",ＰＣＲ。〃/抗cαﾉﾉo〃α"c/DMDNA Extraction, peR Amplification and DNA aligned,thosebelongingtothesamesequencearealigned, those belonging to the same sequence are Sb9"e"cmgSequencing representedbyasinglesequenＣｅ，andanyun-represented by a single sequence, and any un­ TotalDNAiswereextractedfi･omfieshsam-Total DNA's were extracted from fresh sam­ alignablesequencewasomittedfiomthedataalignable sequence was omitted from the data plesorherbariumspecimensbymodificationsofples or herbarium specimens by modifications of sets．sets. theCTABmethod(Murray＆Thompsonl980；the CT AB method (Murray & Thompson 1980; Arikawa＆Higuchil999;TsubotaetaL2000)orArikawa & Higuchi 1999; Tsubota et al. 2000) or A"αbﾉsjsjWAoqlsAnalysis Methods phenol-chlorofbrmmethod(TsubotaetaL1999）phenol-chloroform method (Tsubota et al. 1999). TreeswereconstructedbythefbllowingthreeTrees were constructed by the following three PCRamplificationsof76cLgenesegmentsandPCR amplifications of rbcL gene segments and methods:theneighborjoining(NJ)(Saitou＆Neimethods: the neighbor-joining (NJ) (Saitou & Nei directseqUenceanalyseswerecarriedoutusingdirect sequence analyses were carried out using l987)，themaximum-parsimony（ＭＰ）method1987), the maximum-parsimony (MP) method standardconditionsaccordingtothemanufactur-standard conditions according to the manufactur­ (Fitchl971),andthemaximum-likelihood(ＭＬ）(Fitch 1971), and the maximum-likelihood (ML) ersinstructions、Thereactionswereperfbrmediners instructions. The reactions were performed in method(Felsensteinl981).Thetreesobtainedbymethod (F elsenstein 1981). The trees obtained by thethermalcyclerswithPCRamplificationkitsthe thermal cyclers with PCR amplification kits thethreemethodswereappraisedbythelog-like-the three methods were appraised by the log-like­ andsyntheticprimers・Thesequenceswereelec-and synthetic primers. The sequences were elec­ lihoodmeasurewithNucMLiｎＭＯＬＰＨＹ２３ｂ３lihood measure with NucML in MOLPHY 2.3b3 trophoresedonautomatedsequencers，andana-trophoresed on automated sequencers, and ana­ (Adachi＆Hasegawal996)．(Adachi & Hasegawa 1996). 、伽α"ｃｅａＭ〕ﾉsjs--ANJtreewasconstructed 。 lyzedonthedata-analysissystemThesequenceslyzed on the data-analysis system. The sequences Distance analysis-A NJ tree was constructed obtainedinthepresentstudyweresubmittedtoobtained in the present study were submitted to byNuｃＭＬａｎｄＮＪｄｉｓｔｉｎＭＯＬＰＨＹｖｅｒｓｉｏnby NucML and NJdist in MOLPHY version ｔｈｅＤＤＢＪ/ＥＭＢＬ/GenBanklntemationalNucle-the DDBJ/EMBLIGenBank International Nucle­ 23b3package（Adachi＆Hasegawal996,see2.3b3 package (Adachi & Hasegawa 1996, see otideSequenceDatabaseCollaborationotide Sequence Database Collaboration. AppendixB)usingHKY85model(HasegawaetAppendix B) using HKY85 model (Hasegawa et DetailsandiilrtherinfbrmationfbrDNAex-Details and further information for DNA ex­ aL1985)fiorthedistanceestimations．al. 1985) for the distance estimations. tractingprotocols，amplificationprimers，PCRtracting protocols, amplification primers, PCR Ｐα/Ｍ，､〃α"αlbMS-MPtreeswerecon‐Parsimony analysis - MP trees were con­ protocolsandthoseofsequencingreactionsareprotocols and those of sequencing reactions are structedbyPAUPRat（Sikes＆Lewis2001)，structed by PAUPRat (Sikes & Lewis 2001), describedpreviouslyinArikawaandHiguchidescribed previously in Arikawa and Higuchi ｗｈｉｃｈｉｓａｔｏｏｌｔｏｉｍｐｌｅｍｅｎｔｔｈｅＰａｒｓimonywhich is a tool to implement the Parsimony (1999)，ＤｅＬｕｎａｅｔａＬ（1999)，ＭａｅｄａｅｔａＬ(1999), De Luna et al. (1999), Maeda et al. Ratchetsearches(Nixonl999)withPAUP*,overRatchet searches (Nixon 1999) with P AUP*, over (2000),(2000),andTsubotaandcoworkers（1999,2000, and Tsubota and coworkers (1999, 2000, PAUP＊4.0blO(Swoffbrd2002)JnanalysesbyP AUP* 4.0b 10 (Swofford 2002). In analyses by 2001a,ｂ)．2001a, b). PAUPRat,MPtreesweresearchedusingthePar－PAUPRat, MP trees were searched using the Par­ simonysimonyRatchetsearchstrategyintwenty200it- Ratchet search strategy in twenty 200 it­ DataＤａｊＭ"αｌ１ｙｓｅｓ Analyses erationerationruns runs. DataＤａｒａｓａ－Ａｔｏｔａｌｏｆｌ８１７６cLgenesequences set-A total of 181 rbcL gene sequences Mzx加川/MjAoodq"αｳﾉsな－MLtreeswereMaximum likelihood analysis - ML trees were werewereexamined(TableLwithDDBJ/EMBL/Gen- examined (Table 1, with DDBJ/EMBLIGen­ constructedconstructedwiththeNuｃＭＬｉｎＭＯＬＰＨＹｖｅｒ‐ with the NucML in MOLPHY ver­ BankBankaccessionnumbers)．Thisdatasetcontains accession numbers). This data set contains sionsion23b3package(Adachi＆Hasegawal996)． 2.3b3 package (Adachi & Hasegawa 1996). nineninesequencesobtainedfiomourlaboratoryand sequences obtained from our laboratory and ForFortheMLanalysesHKY85model(Hasegawa the ML analyses, HKY85 model (Hasegawa 172l72registeredsequencesafterBLAＳＴｓｅａｒｃｈｅｓ registered sequences after BLAST searches etetaLl985)wasusedastheestimatemodeLML al. 1985) was used as the estimate model. ML inｉｎｔｈｅＮＣＢＩｈｏｍｅｐａｇｅｆｂｒtheDNAdatabase・ the NCB! homepage for the DNA database. treestreeswereconstructedwiththelocalrearrange- were constructed with the local rearrange­ TheThematrixincludeslOOspecies（l020TUs)of matrix includes 100 species (102 OTUs) of mentmentsearchfiomtheNJtreebyNucMLand search from the NJ tree by NucML and thetheHypnales,S1speciesofLeucodontales,４spe- Hypnales, 51 species of Leucodontales, 4 spe­ NJdistNJdist1.2.5(Adachi＆Hasegawal996);ａｎｄＭＰ l.2.5 (Adachi & Hasegawa 1996); and MP ciesciesofHookeriales，andothersasoutgroups，to of Hookeriales, and others as outgroups, to treesｔｒｅｅｓｂｙＰＡＵＰＲａｔ・Best-confbnnedtransition／ by P AUPRat. Best-conformed transition/ makemakeasurveyfbrthephylogeneticrelationships a survey for the phylogenetic relationships transversiontransversion（Ts/ＴＶ）parameterswereestimated (Ts/Tv) parameters were estimated betweenbetweenthehypnobryaleanmosses．〃"伽α/lリノー the hypnobryalean mosses. Funaria hy­ basedbasedoncalculationswiththeHKY85modelby on calculations with the HKY85 model by grometricaヅomeかｊｃａａｎｄＰ/iWo〃かeﾉﾉａｐａｒｅ"swereused and Physcomitrella patens were used NucMLNucMLemployingthe“-topt，，optionparameter、 employing the" -topt" option parameter. asasthemostdistantoutgroups． the most distant outgroups. Tree乃ｅｅｃｏ"ZparjSo〃ＭﾉﾉMog-ﾉﾉﾉﾋﾋﾞﾉﾉ/ＩＯＯＣ/αﾉ?‘sev- comparison with log-likelihood and sev­ Alignment-Manual』/jg71me"ノーManualalignmentwascarriedout alignment was carried out erale7α〃esな一TreecomparisonwiththeMLcriteria tests-Tree comparison with the ML criteria forfbrthel81sequencesagainstthoseofPﾉiﾘﾉSCO〃－ the 181 sequences against those of Physcomi­ bybyNucMLwascarriedouttoevaluatetheresult- NucML was carried out to evaluate the result­ trella〃e"αｐａｔｅ"ｓａｎｄ此"co6r〕/"碗ｓｃａ６/ｗ"zwithan patens and Leucobryum scabrum with an ingingtrees、Severalmeasureswereusedestimating trees. Several measures were used estimating editoreditorprogramandouroriginalprogram，Sclean program and our original program, Sclean thethelog-likelihoodofdifferenttreetopologies， log-likelihood of different tree topologies, 2.81281(byTsubotal998-2002,inourhttpsite,see (by Tsubota 1998-2002, in our http site, see standardstandarderrors(SES)ofthedifferenceinlog-like- errors (SEs) of the difference in log-like­ AppendixAppendixB）Theotheroriginalprogramswrlt- B). The other original programs writ­ lihoodlihood(Kishino＆Hasegawal989),andtheboot‐ (Kishino & Hasegawa 1989), and the boot­ tenteninPerlandCwerealsoappliedfbrthedata in Perl and C were also applied for the data strapstrapprobabilityestimatedbytheＲＥＬＬ(resam- probability estimated by the RELL (resam­ processing.processingUndeterminedsites，gaps，andre- Undetermined sites, gaps, and re­ piingplingofestimatedlog-likelihoodofsites)method of estimated log-likelihood of sites) method gionsgionsnotclearlyalignablefbrallsequenceswere not clearly alignable for all sequences were (Kishi（KishinoetaLl990;Hasegawa＆Kishinol994） no et al. 1990; Hasegawa & Kishino 1994). excludedexcludedfi･omtheanalyses、Amongsequences from the analyses. Among sequences TheTheprogrampackageCONSELO1e（Shimo- program package CONSEL O.le (Shimo- 652652 HikobiaHikobiaVoLl3,No.４，２００２ Vol. 13, No.4, 2002 dairadaira＆Hasegawa200Dwasalsousedtocalcu- & Hasegawa 200 I) was also used to calcu­ PhylogeneticＰ/iﾘﾉﾉoge"αｊｃａＭ〕入Fes analyses lateslatesthep-valuesoftheconfidencefbrthebifUr- the p-values of the confidence for the bifur­ TheThedatamatrixwaspreparedfbranalysesof data matrix was prepared for analyses of catingcatingcandidatetopologiesusingseveraltesting candidate topologies using several testing phylogeneticphylogeneticrelationshipsofthehypnobryalean relationships of the hypnobryalean procedures:procedures：theApproximatelyUnbiased（AU） the Approximately Unbiased (AU) mosses.mosseSAtotalof2,223topologies，ofwhich A total of 2,223 topologies, of which testtest(Shimodaira2000,2002)usingthemultiscale (Shimodaira 2000, 2002) using the multi scale onlyonly7treetopologieswerebifilrcate，ｗｅｒｅob- 7 tree topologies were bifurcate, were ob­ bootstrapbootstraptechniquqtheKishino-Hasegawa(ＫＨ） technique; the Kishino-Hasegawa (KH) tainedtainedfTomfburanalyses：１ＭＬａｎｄｌＮＪｔｏｐｏｌｏ－ from four analyses: I ML and I NJ topolo­ testtest（Kishino＆Hasegawal989)；andthe (Kishino & Hasegawa 1989); and the giesｇｉｅｓｂｙＮｕｃＭＬ;ａｎｄ２，２２１ＭＰｂｙＰＡＵＰＲａｔｏｖｅｒ by NucML; and 2,221 MP by PAUPRat over Shimodaira-HasegawaShimodaira-Hasegawa（SH）test（Shimodaira＆ (SH) test (Shimodaira & PAUP*.ＰＡＵＰ*・Onthe7bifUrcatingtopologies，local On the 7 bifurcating topologies, local HasegawaHasegawal999;GoldmanetaL2000)． 1999; Goldman et al. 2000). rearrangementrearrangementsearcheswereconstructedfbrthe searches were constructed for the BootstrapＢｏｏｍｍｐ卿ｊ－ＵｓｉｎｇＮｕｃＭＬ，abootstrap- test - Using NucML, a bootstrap­ obtainedobtainedtopologies・Atotalofl2bifhrcatingto- topologies. A total of 12 bifurcating to­ pingpingtestwithlocalbootstrapprobabilities(LBPs； test with local bootstrap probabilities (LBPs; pologiespologieswereusedfbrthefbllowinganalysis、 were used for the following analysis. iniｎ％)(Adachi＆Hasegawal996)wasappliedfbr %) (Adachi & Hasegawa 1996) was applied for MoreMoredetailedtopologiesweresearchedthrough detailed topologies were searched through ▼ thethebestMLtopology・LBPisarelativebootstrap best ML topology. LBP is a relative bootstrap thetheobtainedtreesusinglog-likelihoodmeasure obtained trees using log-likelihood measure frequencyfi･equencyobtainedffomtopologysearchbylocal obtained from topology search by local andandseveraltests(Table２）HKY85modelwas several tests (Table 2). HKY85 model was rearrangementsrearrangementsofNucMＬＴｈｅｖａｌｕｅｗａｓｃｏｍ‐ of NucML. The value was com­ usedusedastheestimatemodelwiththe as the estimate model with the 勺 parableparablewithavalueofFelsenstein1s(1985)boot- with a value of Felsenstein's (1985) boot­ transition/transversiontransition/transversionparameterestimatedけom parameter estimated from strapstrapprobability,anditisalittlelargerthanthe probability, and it is a little larger than the datadatasetfbrtheMLtreeas339・Thehighestlike- set for the ML tree as 3.39. The highest like­ standardstandardbootstrapprobability bootstrap probability. lihoodlihoodtreeofhypnobryaleanmosseswasob- tree of hypnobryalean mosses was ob­ ConsensusCb"Ｓｃ"s"ｓ／ree-Astrictconsensustreelbr tree - A strict consensus tree for tained,tained,andshowninFigLTheLog-likelihｏｏｄ and shown in Fig. I. The Log-likelihood thethebifUrcatingtopologieswithhigh-ranking bifurcating topologies with high-ranking valueｖａｌｕｅｆｂｒｔｈｅＭＬｔｒｅｅｗａｓ－１５４４５８±794.6． for the ML tree was - 15445.8 ± 794.6. log-likelihoodlog-likelihoodvaluesthatpassedtheAUtestwas values that passed the AU test was TheＴｈｅＬＢPsfbrthecladeswerecomparativelylow LBPs for the clades were comparatively low alsoalsocoｍｐｕｔｅｄｂｙＰＡＵＰ*． computed by PAUP*. valuesvalues（41-100％）Becauseeachtopologydid (41-100%). Because each topology did notnotgainadecidedadvantageoverothertopolo- gain a decided advantage over other topolo­ gies, a 50% majority-rule consensus tree of Results gies，ａ５０％majority-ruleconsensustreeof high-rankinghighrankingtreeswasalsoobtained，ａｎｄpre- trees was also obtained, and pre­ ObtainedＯ６ｊａｊ"ecl/se9"e"cedma sequence data sentedinFig2Thistreealsohasatendencyto-sented in Fig. 2. This tree also has a tendency to­ Newr6cLsequenceswereobtainedfbrelevenNew rbcL sequences were obtained for eleven wardtheMLtree・ward the ML tree. species:species：ＢｊｓＭｊａノノ"g"〃α，ＣＭＯ"e"'o〃ﾉｶﾞﾉﾉcノー Bissetia lingulata, Cratoneuron jilici­ TheMLandconsensustreesfionnnotableThe ML and consensus trees form notable """０，Ｇｏﾉﾉα"/α'wgj"Csα，Ｇ､Ｓ〃た"｡e"s,ｆ/ｏ"zα/ja-num, Gollania ruginosa, G. splendens, Homalia­ clades:(1)twomaiorcladesareshowninFiglclades: (I) two major clades are shown in Fig. I cﾉellp〃sjaXgj"o"ｊα""８，〃ｂ"､ﾉﾉo〃dro〃scallpeﾉｰdelphus targinonianus, Homaliodendron scalpel­ [the[thecombinedHypnales（s・ｓｔｒｌ＝sensuVitt combined Hypnales (s. str. [= sensu Vitt ﾉ(/bﾉﾉ川Ｌｅ"COC/ｂ〃Sc〃ｏｊ伽,ノVbcke'QpsjMjjj-lifolium, Leucodon sciuroides, Neckeropsis niti­ l984D-Leucodontalesclade(87％LBPvaluein1984])-Leucodontales clade (87% LBP value in 伽ﾉﾋJ,Ｐｊ""areﾉﾉαα'、g"α、ﾉjα"z"o6'測加ｓ"6se‐dula, Pinnatella ambigua, Thamnobryum subse­ MLtree,８２％supportinconsensustree);andtheML tree, 82% support in consensus tree); and the γj川"ＭｎｄＷ/iiﾉﾉImco"cav(/Ｍα，For〃ﾉhiaco"riatum, and Wijkia concavifolia. For Wijkia con­ Hookerialesclade(64％LBP,７３％)]inthehyp‐Hookeriales clade (64% LBP, 73%)] in the hyp­ cav肋/jaanentiresequencel,428bplongwascavifolia an entire sequence 1,428 bp long was nobryaleanclade(96％LBP,100％);ａｎｄ(2)４８nobryalean clade (96% LBP, 100%); and (2) 48 obtainedThesequenceshavebeensubmittedtoobtained. The sequences have been submitted to basicclades[Sematophyllaceaeclade(96％LBP，basic clades [Sematophyllaceae clade (96% LBP, theDNAdatabaseundertheserialaccessionthe DNA database under the serial accession １００％)；Ｅ川血〃clade（96％LBP，100％)；100%); Entodon clade (96% LBP, 100%); numberｓａｓｓｈｏｗｎＴａｂｌｅＬnumbers as shown Table I. Ｅ"joc/､〃ノｗ６ｊｃ""c/"sclade；７１ﾉﾉ"城""TcladeEntodon rubicundus clade; Thuidium clade (100％LBP，100％);肋ｊｅｔｊ"eﾉﾉａａ６ｊｅｊｊ"αclade；(100% LBP, 100%); Abietinella abietina clade; 比9"e"Ｃｅαﾉﾉg"ｍｅ"jSequence alignment AOpjerygj叩sjs-O"hotAecj"ｍｃｌａｄｅ（94％LBP，Isopterygiopsis-Orthothecium clade (94% LBP, ThedatamatrixwaspreparedfbrphylogeneticThe data matrix was prepared for phylogenetic 100％);ＥＣ"/ｑＷ〃"e"jjclade;Hypnaceaeclade100%); Boulaya mittenii clade; Hypnaceae clade analysesbymanualalignmentofthesequences・analyses by manual alignment of the sequences. (71％LBP，５５％)；血"COC/ｂ"-ne7o6r〕ﾉo''－０〕ノー(71% LBP, 55%); Leucodon-Pteroblyoll-Cry­ Ｔｈｅｄａｔａｓｅｔｓｗｅｒｅｓｕｂｍｉｔｔｅｄｔｏａ５％chi-squareThe data sets were submitted to a 5% chi-square p力aeaclade(89％LBP,５５％);Ａ"o"ZodO〃'wgeﾉﾉﾉphaea clade (89% LBP, 55%); Anomodon rugelii testbyTREBPUZZLE50(Strimmer＆Haeselertest by TREE-PUZZLE 5.0 (Strimmer & Haeseler clade；Ｈｅ”ogje"αｐｅ〃o6"sjaclade；んOp/eﾉﾌﾉｰclade; Herzogiella perrobusta clade; Isoptery­ l996),andallthedatapassedthetest・Forthe1996), and all the data passed the test. For the gj""ﾊﾉﾉ"eα/eclade;Prjo"odO〃(ノセ"s"sclade;。"gium vineale clade; Prionodon dens us clade; An­ analyses,atotalofLO92sitesofl8176cLse-analyses, a total of 1,092 sites of 181 rbcL se­ /伽cﾊﾉα／b""Csα"αclade；Ｒﾉﾉ〕ﾉ肋α火llpA"s-titrichia formosana clade; Rhytidiadelphus­ quenceswereusedNucleotideｆｉ･equenciesesti-quences were used. Nucleotide frequencies esti­ fMoco〃"m-LoGWo6/Ｗｍ－Ｃｔｅ"/血"mclade(８９Hylocomium-Loeskeobryum-Ctenidium clade (89 matedftomthedａｔａｓｅｔｗｅｒｅＡ＝29.2％，Ｃ＝mated from the data set were A = 29.2%, C = ％LBP,100％);ＭﾉＭ"碗-sb〃o〃"mclade(８８% LBP, 100%); Myurium-Sciaromium clade (88 17.5％，Ｇ＝22.2％，ａｎｄＴ＝31.1％．Inthesites17.5%, G = 22.2%, and T = 3l.l %. In the sites ％LBP,１００％);伽c/iDlp"s-P叩ﾉﾉﾉarjq-伽c/1)ノー% LBP, 100%); Trachypus-Papillaria-Trachy­ ofthealignedr6cLsequences,４１７sites(＝３８２％of the aligned rbcL sequences, 417 sites (= 38.2% podopsjs-D"/ﾊﾉeﾉﾉａｃｌａｄｅ（89％LBP，100％)；podopsis-Duthiella clade (89% LBP, 100%); inLO92sites)werevariable．in 1,092 sites) were variable. P/cJgjotAecj""ｚｃｌａｄｅ（98％LBP，１００％)；Plagiothecium clade (98% LBP, 100%); Ｈ・TsuBoTA,Ｔ､ARIKAwA,Ｈ・AKIYAMA,ＥＤＥＬｕＮＡ,Ｄ・GoNzALEz,Ｍ・HIGucHIANDH・DEGucHIH. TSUBOTA, T. ARIKAWA, H. AKIYAMA, E. DE LUNA, D. GONZALEZ, M. HIGUCHI AND H. DEGUCHI ６５３653

Table2Compansonoflog-likelihoodscoresamongthel2resultingbifilrcatingtopologiesamongtheTable 2. Comparison of log-likelihood scores among the 12 resulting bifurcating topologies among the hypnobryaleanmossesfbr76cLsequenceswiththeHKY85model(HasegawaetaL1985）Thelog-likelihoodhypnobryalean mosses for rbcL sequences with the HKY85 model (Hasegawa et al. 1985). The log-likelihood valueswerecalculatedbyNucML(Adachi＆Hasegawal996),andallthep-valuesbyCONSEL(Shimodaira＆values were calculated by NucML (Adachi & Hasegawa 1996), and all the p-values by CONSEL (Shimodaira & Hasegawa2001）fiPomlO,O00repetitions、Thelog-likelihoodvaluesofthehighestlikelihoodtreesaregiveninHasegawa 200 I) from 10,000 repetitions. The log-likelihood values of the highest likelihood trees are given in anglebrackets,andthedifferencesinlog-likelihoodofalternativetreesfiomthatoftheMLtreeareshownwithangle brackets, and the differences in log-likelihood of alternative trees from that of the ML tree are shown with theirstandarderrors(SE)fbllowing±・Thep-valuesthatarenotsignifiCantata＝OO5areemphasizedinboldtheir standard errors (SE) following ±. The p-values that are not significant at a = 0.05 are emphasized in bold typeTopologieswithasterisksintheAUcolumnwereusedfbrtheconsensustree(Ｆｉｇ２）type. Topologies with asterisks in the AU column were used for the consensus tree (Fig. 2).

£-Values Ｒａnｋ△/士ＳＥRank t:,/± SE MethodProgramMethod Program ＮｏｔｅNote ＡＵＢＰＫＨＳＨAU BP KH SH 。ｇ傘４白５●６竃３傘８●３戦０傘７△４。２３ 砂茄佃瓠⑬州Ⅳ列ｍ刈りⅡ ００００００００００００ ２００１０１００００００ ３８９１５４４４５７４０ ９０２５５６８５７８５０ ００００００００００００ ６３３３２３２２２２１０ ５４０５９４８７７５７３ ４６５８０８３７６３９０ ００００００００００００ ９８８８８７７７７６４０ ５９５４１７９７３０８４ ７５８６２９５４０１１６ 皿ⅢⅢⅢⅢⅢｗｗｗⅢⅢ川 ００００００００００００ 引叺２６６０２３孔０４０ ５０シ７５０シ０シ５（Ｕ１４００”５ 斜士士士士士士士士士士士 ５２２４３５３３４０６〆０８ ８６８Ⅱ０４９８Ⅱ。Ｌ０Ｏ ＲＲＲＲＲＲＲＲＲＲＬＬ 馳即岬山川川岼岼岬岼 醐蜘ＵＭ皿皿刀刃勺刃 ＬＬ 眺臥恥恥恥恥叺叺恥恥叩、 ｍｍｗｍｍｍｗｍｍｍ叩珊 ＋＋ｌ＋＋＋ｌＩくＩＮ シ７〈Ｕ今Ｊ７７６（５６４，句Ｊ７』 ｔｔｔｔｔｔｔｔｔｔ く－１１１２２つ』２４‐５６ ａａａａａａａａａａ ●●●●●●●●●●●● I <-15445.8> ＭＬ(Fig.1）ML(Fig. I) 0.699' 0.239●●●●●●●。●●●● 0.654●●●●●●●●●●●● 0.957 ML PAUPRat + NucML LRSfiPomMP-lLRS from MP-l ２３４５６７８９０１２ 2 -9.9 ± 26.7 0.564' 0.080 0.346 0.895 ML PAUPRat + NucML LRSfiDmMP-2LRS from MP-2

3 ’’’’’’一｜｜刊 -12.7 ± 28.0 0.435' 0.092 0.305 0.858 MP PAUPRat(MP-l) 4 -16.5 ± 47.3 0.516' 0.115 0.358 0.846 ML PAUPRat + NucML LRSfiPomMP-4LRS from MP-4 5 -16.9± 30.7 0.433' 0.055 0.290 0.812 ML PAUPRat + NucML LRSfiFomMP-3LRS from MP-3 6 -20.9 ± 54.7 0.488' 0.146 0.348 0.779 ML PAUPRat + NucML LRSfiPoｍＭＰ－５LRS from MP-5 -22.5 ± 39.6 0.373' 0.048 PAUPRat (MP-2) ◆ 7 0.283 0.795 MP 8 -23.0 ± 38.8 0.360' 0.045 0.277 0.774 MP PAUPRat (MP-3) 9 -27.1 ±47.6 ＭＥME 0.297' 0.057 0.276 0.730 MP PAUPRat (MP-4) 10１１１ -40.4 ± 61.4 0.264' 0.078 0.253 0.601 MP PAUPRat (MP-5) 11 -54.9 ± 60.3 0.192' 0.045 0.179 0.481 ML NucML ＬＲＳｆｉＰｏｍＮＪ－ｌLRS from NJ-l ｄ Ｊ 川 Ｊ ＋ 12 -160.5 ± 80.2 0.003 0.000 0.030 0.046 NJ NucML + NJdistＳ (NJ-l)

Ｎｏｔｅ－ＡＵ:Approximateunbiasedtest(Shimodaira2000,2002),ＢＰ:thebootstrapselectionprobabilityofFelsenstein（1985）Note. - AU: Approximate unbiased test (Shimodaira 2000, 2002), BP: the bootstrap selection probability of Felsenstein (1985) estimatedbytheRELLresamplingmethod(KishinoetaLl990;Hasegawa＆Kishinol994),ＫＨ:Kishino-Hasegawatest(Kishinoestimated by the RELL resampling method (Kishino et al. 1990; Hasegawa & Kishino 1994), KH: Kishino-Hasegawa test (Kishino &＆Hasegawal989),ＳＨ:Shimodaira-Hasegawatest(Shimodaira＆Hasegawal999;GoldmanetaL2000);ａｎｄＬＲＳ:thelocal Hasegawa 1989), SH: Shimodaira-Hasegawa test (Shimodaira & Hasegawa 1999; Goldman et al. 2000); and LRS: the local reanPangementsearch(Adachi＆Hasegawal996)ｂｙＮｕｃＭＬrearrangement search (Adachi & Hasegawa 1996) by NucML.

Pleurozium-HylocomiastrumP/e"rozj"ｍ-町/ＯＣＣ〃ａｓ伽"ｚｃｌａｄｅ（82％LBP， clade (82% LBP, OurOursmdycorroboratesthatl)theHypnales(s study corroborates that I) the Hypnales (s. 82%);82％);Brachytheciaceaeclade(66％LBP,１００％)； Brachytheciaceae clade (66% LBP, 100%); str.strl＝sensuVittl984]）andLeucodontalesare [= sensu Vitt 1984]) and Leucodontales are ClirnaciaceaeClimaciaceaeclade(99％LBP'100％);Ｐ/e"rozJ- clade (99% LBP, 100%); Pleurozi­ notmonophyleticentities，whiletheHookerialesnot monophyletic entities, while the Hookeriales opsisOpsLsclade;Stereophyllaceaeclade（１００％LBP， clade; Stereophyllaceae clade (100% LBP, withwiththeinclusionofthePtychomniaceaeara2） the inclusion of the Ptychomniaceae are; 2) 100%);100％);lVboC/O/jc/io"ｚｊｊｒａＷ""α"e"sjlyclade;〃 Neodolichomitra yunnanensis clade; To­ thecombinedHypnales（ｓｓｔｒ．［＝sensuVittthe combined Hypnales (s. str. [= sensu Vitt mentypnumｍｅ"卯""ｍ〃"e"sclade；Amblystegiaceaeclade nitens clade; Amblystegiaceae clade 1984])l984]）ａｎｄLeucodontalescompriseawellsup‐ and Leucodontales comprise a well sup­ (99%(99％LBP,100％);Ｈｊｌｐ""、ｃ叩ressl/M71eclade； LBP, 100%); Hypnum cupressiforme clade; portedsistercladetotheHookeriales;ａｎｄ３)theported sister clade to the Hookeriales; and 3) the Trachyloma伽c/iﾘﾉﾉo加ａｊＭｃ"mclade；Neckeraceae-FoKMo- indicum clade; Neckeraceae-Forsstro­ familiesfamiliesEntodontaceae（s・str．［＝sensuVitt Entodontaceae (s. str. [= sensu Vitt

Ｐ emiae〃αclade(86％LBP,100％);Lembophyllaceae clade (86% LBP, 100%); Lembophyllaceae 1984;l984;TsubotaetaL2000]),Fontinalaceae,Ｓｅ‐ Tsubota et al. 2000]), Fontinalaceae, Se­ cladeclade（82％LBP，100％)；nJxjp/iし〕ﾉﾉﾉ"m-G/Cs‐ (82% LBP, 100%); Taxiphylium-Glos­ matophyllaceae（s」at.［＝sensuTsubota＆co-matophyllaceae (s. lat. [= sensu Tsubota & co­ sadelphus-Miyabea-Bissetia-HomaliadelphusSα(/elp/i"s-Mi〕ﾉa6ea-肋Ｍｊａ－Ｈｏｍα/ｊａｄｅ/p伽 workersworkers2000,2001a,ｂ]),Thuidiaceae(ｓｓｔｒ.[＝ 2000, 2001a, b]), Thuidiaceae (s. str. [= cladeclade（90％LBP,７３％);Anomodontaceaeclade (90% LBP, 73%); Anomodontaceae clade sensusensuTsubotaetaL2000DandPlagiotheciaceae Tsubota et al. 2000]) and Plagiotheciaceae (70%(70％LBP，１００％)；Fontinalaceaeclade（100％ LBP, 100%); Fontinalaceae clade (100% (s.(s・str.［＝sensupresent]）areresolvedasmono- str. [= sensu present]) are resolved as mono­ LBP,LBP，100％)；Hookerialesclade（64％LBP， 100%); Hookeriales clade (64% LBP, phyleticphyleticgroups、MembersoftheAmblystegia- groups. Members of the Amblystegia­ 73%);73％);LeY）/Cs/ｏｍ"mmacrocaZp"mclade;Ｂａ7〃－ Leptostomum macrocarpum clade; Bartra­ ceae,ceae，HypnaceaaandLeucodontaceaearedis- Hypnaceae, and Leucodontaceae are dis­ mia〃ａｐｏ加加'伽clade；Cyrtopodaceaeclade pomiformis clade; Cyrtopodaceae clade persedpersedamongseveralclades． among several clades. (100%(100％LBP，100％)；HDlp"ode"Q/、〃〃?e"z/asi/ LBP, 100%); Hypnodendron menziesii clade;clade；RacOpj/""ｚｃｏｍ′ｏｍ/ace""Zclade；けr‐ Racopilum convolutaceum clade; Pyr­ Discussion rhobryumr/io6M"、ﾉαﾉﾉjs-g7aZjaec/αcノビ,．』"/ZJco"?"j""?／z"‐ vallis-gratiae clade; Aulacomnium tur­ gidumgjc/""zclade；Ｏ〃hoc/Ｄ"〃"１ノノ"eα'でclade； clade; Orthodontium lineare clade; 1.LOrdinalrelationships Ordinal relationships Philonotis-LeiomelaP/iﾉﾉo"otjs-Leｊｏｍｅ/αclade（97％LBP，100％)； clade (97% LBP, 100%); OurOurphylogeneticanalysiswith76cLlarge-scale phylogenetic analysis with rbcL large-scale MielichhoferaＭｅﾉﾉcﾊﾉjq/ｂｒａｅ/o"gajeclade；PAO/ｊａｃｊｗ血 elongate clade; Pholia cruda datadatasetdidnotrecovereitherthemonophylyof set did not recover either the monophyly of clade;cladaMniaceaeclade（100％LBP,１００％);Hed‐ Mniaceae clade (100% LBP, 100%); Hed­ thetheorderHypnales(s・str.[＝sensuVittl984]),or order Hypnales (s. str. [= sensu Vitt 1984]), or wigiaceaewigiaceaeclade(100％LBP,１００％);ａｎｄＰＷｔＪｍｌｊａ clade (100% LBP, 100%); and Wardia thethemonophylyoftheLeucodontales；although monophyly of the Leucodontales; although hygrometricaｈ〕lgromeかjcaclade］ clade]. thetheanalysisrecoveredthemonophylyofthe analysis recovered the monophyly of the 654654 HikobiaＨｉｋｏｂｉａＶｏｌｌ３,No.４，２００２ Vol. 13, No.4, 2002

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ｐＨ '0.01０.O1substitutions/sltE substitut{onS/site FiglThehighestlikelihoodtreefbrthealignedl,Ｏ９２ｂｐｏｆｔｈｅｌ８］ノカcLgenesequences(HKY85model;α/β＝Fig. I. The highest likelihood tree for the aligned 1,092 bp of the 181 rbcL gene sequences (HKY85 model; a! {3 339;ｌｎＬ＝-154458±794.6ｂｙＮｕｃＭＬ);andthemajorclades:theHypnales(s」at.)clade[＝thecombined3.39; In L = -15445.8 ± 794.6 by NucML); and the major clades: the Hypnales (s. lat.) clade [= the combined Hypnales(s・str.[＝sensuVittl984])-Leucodontalescladc],andtheHookerialesclade・ＴｈｅｈｏrizontallengthofHypnales (s. str. [= sensu Vitt 1984J)-Leucodontales clade], and the Hookeriales clade. The horizontal length of eachbranchisproportionaltotheestimatednumberofnucleotidesubstitutions・Therootisarbitrarilyplacedontheeach branch is proportional to the estimated number of nucleotide substitutions. The root is arbitrarily placed on the branchleadingtothenl"αｒｊａﾉl【Woll1eﾉﾉ化αａｎｄﾉ､ﾉﾘWol"/Ｍ/αﾉﾌqZe"s、Localbootstrapprobabilities（LBP;％）branch leading to the FUllaria hygrometrica and Physcomitrella patens. Local bootstrap probabilities (LBP; %) morethan50％areshownaboveornearbranchesShadedboxesidcntifythetraditionalLeucodontales．more than 50% are shown above or near branches. Shaded boxes identify the traditional Leucodontales. Vl Vl 2: 0-, o m ~ ~ ~ ~ ~ ~ ~ ~ ~ t-< 2: -< z Ci Cl o c o m a n !7l ::r: :» :» ~ ~ ~ tIl R C 1:: > ;J ;l ~ ~ HTsuBoTA,Ｔ・ARIKAwA,Ｈ・ＡＫＩＹＡＭＡ,ＥＤＥＬｕＮＡ,Ｄ・GoNzALEz,ＭＨＩＧｕｃＨＩＡＮＤＨ・DEGucHI６５５J> J> J> （・←、一・の）の①而亡・夛工 の①ｌｍＥ○つＯＮ一匹 の①而己囚 ‐（・』←の・の）の①一⑩亡旦夛エ］ 。… lat.) str.)- (s. (s. Bryales 認’三

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100 崖 Ｊ【Ｊ【Ｊ「Ｊ【二「【】信 Entodontaceae(ｓｓｔ｢.） 一一一 73 蝋ilj鑿iil識w二二＝．,----_...... ThuidiumThuﾉdjUmclade clade smuel/ｍａｎａ 梛liiﾄ|(;lmll;;』 ｍねscens-．_____....11 Isopterygiosis-OrthotheciumﾉsOpfelygjOsjS-O'tholhec/umclade clade 鑿蕊llii鑿二m;~~;;: ~~::;~~~nne HypnaceaeHypnaceaeclade clade ~:~d~~~~m;n~s~======~ Leucodon sohayakiensis DOZYB jBponica Felipponsa esquirolii LeucOdon蝋i霧ｌＩＩｉ曇り率 sciuroides Leucodon nipponicus Leucodon sapporensis t~~~g~ }~r,:g:~tus Leucodon-Pterobryon-CryphaeaLeucodOn-PleloblyoかＣ'yphaeacIade clade 繍iiiii1il1ji`。Leucodon SBcundus Cryphaea蕊繍 sinensisけ〃ごｌｓ 55 ｉｊｓｏｂ "ａ ~fol~g~c:It;~ec;,~::t:;fo'iaｄｂｎｔ Pferobryon arbusculaｂｕｓＣ， pterobryon densum，ｎＳＵＦ PferobryopsisＩｏｎｅｌ ori(:!nlalis subsp.subspyuennanensls yuennanenSIS omoi:Jon rugs/IIｇｅ〃 erz jella psrrobustaｇｎＤｂ vinealeＶＴｎｅ届 ensus A､"ｍＣｈｊａｌｂ'mosanannosana １００ ００Ｆ R hussquarros~u 555５ ~ hus ~r?o~~cus Rhytidiade/phus-Hy/ocomium- 2182 Rhyl/diade/Phus-ﾉヵﾉbcomjUm‐ Hljillll讓蕊露i簔 s Loeskeobryum-CtenidiumLoeskeoblWm-Clenjdjumclade clade 65119° Loes~eobryum cavifql,,,iu,,,m======:; 」』且 'tl;~r::;::;;~::t~s,}gfften 1 Myurium-SciaromiumMyu"um-Scjammjumclade clade １００ ００「 Trachypus-Papillaria­77achypus-Papjｿlalja- １００ Trachypodopsis-Duthiella７７achypodOpsjS-Dulhje"aclade clade ，１，ii1； I PlagiotheciumP/agjOfhec/umclade clade 嘩礎砿靜 鑿 '~-======4 I Pleurozium-HylocomiastrumP/euloz/um-hMOcom/as伽ｍｃｌａｄｅ clade 鑿|篝iiiiili義！ ~=====l 鑿霧護讓襄 BrachytheciaceaeBrachytheciaceaecIade clade I ClimaciaceaeC1imaciaceaecIade clade 鑿iiliiiiii雲霧~====:::::I StereophyllaceaeStereophyIlaceaeclade clade

Ｆｉｇ２Ｔｈｅ５０％majority-ruleconsensustreefbrthellbifUrcatingtopologieswithhigh-rankinglog-likelihoodFig. 2. The 50% majority-rule consensus tree for the II bifurcating topologies with high-ranking log-likelihood valuesthatpassedtheAUtest、Therootisarbitrarilyplacedonthebranchleadingtothe〃"Ｍα/1Ｗｏｍｅｒ７ｊｃａａｎｄvalues that passed the AU test. The root is arbitrarily placed on the branch leading to the Funaria hygrometrica and Ｐ/1〕ﾊFCC〃Ｍ/〃α/e"s、Physcomitrella patens. H.Ｈ・TsuBoTA,ＴＡＲＩＫＡｗＡ,Ｈ・AKIYAMA,ＥＤＥＬｕＮＡ,Ｄ・GoNzALEz,Ｍ・HIGucHIANDH・DEGucHI６５７ TSUBOTA, T. ARIKAWA, H. AKIYAMA, E. DE LUNA, D. GONZALEZ, M. HIGUCHI AND H. DEGUCHI 657

＊ 55５５*

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鍵鵜 Neckeraceae-ForsstroemiaNeckeraceae-FO'ssf71oemjaclade clade ’ ＵＩＢ「ｺ 828２ ﾌｅｍｌ．、■ ｡ｎｍO月月行kＢｍ 、｡"ｃｈｏｍｌｆｎｏＤ :===::::::::; LembophyllaceaeLembophyIlaceaecIade clade ;鰯liil菫iil;蕊 Taxiphyl/um-G/ossade/phus-Miyabea-耐xﾉPhyﾉﾉUm-G/Ossadelphus-Myabea- 828２ ~====~.I Bissetia-Homa/iade/phusB'Sse脂一HOma脂deゎhuscIade clade AnomodontaceaecIade 1001００ IAnomodontaceae clade !!===~ FontinalaceaeFontinalaceaeclade clade 555５ ＵＵＫＵ『０．回ＵｕＩＩＩＵ ~====4 ｉ鰯jH轍!’ HookerialesHookerialesclade clade 555５ ii1鱗ill蝋…, MaclDmlmUmjncuMわ"ｕｍ ＭａｃｍｍｊｍＵｍｊｎｃｕｒＷｂﾉﾉﾘ、 霧

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1001００ I------""'C 鰯lljll:鰹H1鯏但竺昌ニニニニニニコＰ'Wbno施一LejOme/aclade------,1 Phi/onotis-Leiomela clade 糒噸一顎 階鰐WI:|i;!；a｡lbngaね 100１００ I MniaceaeMniaceaeclade clade ;;s====~ HedwigiaceaecIadeHedwigiaceae clade 100，．.「 |j篝鍵鍵il1篝 |蕊lii霧i篝！｝ Fig.２．continuedFig. 2. continued. ,

HookerialesThecombinedHypnalesandLeuco-Hookeriales. The combined Hypnales and Leuco­ somespeciesoftheBryales,Hookeriales,Leuco‐some species of the Bryales, Hookeriales, Leuco­ ▲ dontales，however，compnseawell-supporteddontales, however, comprise a well-supported dontalesandHypnaleSDeLunaetaL(2000)anddontales and Hypnales. De Luna et al. (2000) and sistercladetotheHookeriales・Ourresultsalsosister clade to the Hookeriales. Our results also BucketaL(2000)revealedthenon-independenceBuck et al. (2000) revealed the non-independence showedthemonophylyofallhypnobryaleanshowed the monophyly of all hypnobryalean ofthetraditionalordersHypnalesandLeucodon-of the traditional orders Hypnales and Leucodon­ mossesaspreviouslydiscoveredbyDeLunaandmosses as previously discovered by De Luna and tales,andsuggestedthenecessitytobereconsid-tales, and suggested the necessity to be reconsid­ coworkers(1999,2000),andBucketaL(2000)．coworkers (1999, 2000), and Buck et al. (2000). eredasasinglemajorlineageered as a single major lineage. Inthepresentstudywithalarge-scaledataset，In the present study with a large-scale data set, HDlp"α伽α"`Le"ＣＯ血"〃/“Hypnales and Leucodontales monophylyofeachorderHypnales(s・strI＝sen-monophyly of each order Hypnales (s. str. [= sen­ Tsubotaetal．（1999）showedapreliminaryTsubota et al. (1999) showed a preliminary suVittl984DorLeucodontaleswasnotrecov-su Vitt 1984]) or Leucodontales was not recov­ phylogeneticrelationshipoftheHypnalesbasedphylogenetic relationship of the Hypnales based eredunderdifferenttreesearchprocedures・Theered under different tree search procedures. The on76cLdataset，especiallyrevealingthepoly-on rbcL data set, especially revealing the poly­ combinedHypnales（s・str.）andLeucodontalescombined Hypnales (s. str.) and Leucodontales phylyoftheHypnaceae，ｗｉｔｈ３４０ＴＵｓｆｂｒｔｈｅphyly of the Hypnaceae, with 34 OTUs for the cladewassupportedwith８７％ＬＢＰｉｎｔｈｅＭＬclade was supported with 87% LBP in the ML ａｎａｌｙｓｉｓ、ＤｅＬｕｎａｅｔａＬ（1999）suggestedthatanalysis. De Luna et al. (1999) suggested that ｔｒｅｅａｎｄ８２％supportintheconsensustree,root-tree and 82% support in the consensus tree, root­ monophylyofpleurocarpousmossesincludingmonophyly of pleurocarpous mosses including edbythecladeoftheFontinalaceacThiscom-ed by the clade of the Fontinalaceae. This com- 658 HikobiaＨｉｋｏｂｉａＶｏＬｌ３,No.４，２００２ Vol. 13, No.4, 2002 binedbinedcladecon･espondstotheHypnales(s」at.） clade corresponds to the Hypnales (s. lat.) HDP"αceaeHypnaceae asasrevealedbyDeLunaetal.(2000)andnamed revealed by De Luna et a!. (2000) and named TheThefiamiliesHypnaceae（withca40genera： families Hypnaceae (with ca 40 genera: asastheHypniaebyBucketal.(2000)．Members the Hypniae by Buck et a!. (2000). Members Vittl984)areamongthemostdiversifiedtaxon-Vitt 1984) are among the most diversified taxon­ previouslypreviouslytreatedastheLeucodontalesappeared treated as the Leucodontales appeared omicomicgroupsinthepleurocarpousmosses、The groups in the pleurocarpous mosses. The asasseveralcladesinthecombinedclade・Ourre- several clades in the combined clade. Our re­ familialftlmilialcircumscriptionwithintheHypnales，es- circumscription within the Hypnales, es­ sultssultsofthepresentstudybasedonthe76cLgene of the present study based on the rbcL gene peciallypeciallybetweentheHypnaceaeandotherfami- between the Hypnaceae and other fami­ dodonotconcurwiththeclassificationandphylog- not concur with the classification and phylog­ lies,lies，suchastheSematophyllaceae,Plagiothecia- such as the Sematophyllaceae, Plagiothecia­ enyenyoftheorderHypnalesproposedbｙＶｉｔｔ of the order Hypnales proposed by Vitt ceae,ceae，andAmblystegiaceae，ｈａｓｌａｔelybecomea and Amblystegiaceae, has lately become a (1984),(1984),andBuckandVitt(1986）Theseresults and Buck and Vitt (1986). These results subjectsubjectofspecialinterestandbeenrepeatedly of special interest and been repeatedly alsoalsosuggestthattheirsporophyticmorphological suggest that their sporophytic morphological discusseddiscussed(e9.,NishimuraetaL1984)． (e.g., Nishimura et a!. 1984). similarities,similarities，suchasdevelopmentofperistomal such as development of peri stomal InＩ、theobtainedMLandconsensustrees，the the obtained ML and consensus trees, the teethandexothecialcells,aremultipletransitionsteeth and exothecial cells, are multiple transitions familyfamilyHypnaceae,aswellasitstypegenusHDlp- Hypnaceae, as well as its type genus Hyp­ @ totosimilarsolutionstoepiphytismintheHypnales similar solutions to epiphytism in the Hypnales num,""肌provedtobenon-monophyleticdistributing proved to be non-monophyletic distributing (s.(slat.)． lat.). ininvariouspositionsinthetrees,regardlessofthe various positions in the trees, regardless of the inclusioninclusionofD/α伽α火!〃/1α(inc.〃ｏｒ/ieM/α)and of Pylaisiadelpha (inc. Brotherella) and HookerialesHDCﾉ(ｅｒｊａにs Wijkia〃/hiainthefamily，althoughtwospeciesof in the family, although two species of MonophylyMonophylyoftheHookeriales［inc・ｔｈｅＰｔｙ‐ of the Hookeriales [inc. the Pty­ fbip"z川ａｎｄｔｈｅｇｅneraGoﾉﾉα"jαａｎｄ〃/αjsjaHypnum, and the genera Gollania and Pylaisia chomniaceae,chomniaceae,representedbyPltycAom"jo〃αcjc"－ represented by Ptychomnion acicu­ formfbnnasinglecladewith71％LBPsupportThis a single clade with 71 % LBP support. This lare;，７e;sensuBucketaL2000;sensuBuck＆Goffi- sensu Buck et a!. 2000; sensu Buck & Goffi­ fact,fact，togetherwiththepresentanalysiss叩ple- together with the present analysis supple­ netnet2000]wassupportedwith55％ＬＢＰｉｎｔｈｅ 2000] was supported with 55% LBP in the mentedmentedbyadditionaltaxa,corroboratesthereport by additional taxa, corroborates the report MLＭＬｔｒｅｅａｎｄ７３％supportinconsensustree，be- tree and 73% support in consensus tree, be­ ofofTsubotaetal(1999)，suggestingthatthefiam‐ Tsubota et a!. (1999), suggesting that the fam­ ingingsistertothecombinedHypnales-Leucodon- sister to the combined Hypnales-Leucodon­ ilyilyHypnaceae,aswellasthegenusjL卯"""',is Hypnaceae, as well as the genus Hypnum, is talestalesinourphylogeneticanalyses;althoughsome in our phylogenetic analyses; although some polyphyleticalthoughthegenushasbeencare-polyphyletic although the genus has been care­ ofoftheobtainedtreesshowedthatthePtychomni- the obtained trees showed that the Ptychomni­ fullyfilllyrevisedbymonographers(e9.,Ａ､ｄｏ1986, revised by monographers (e.g., Ando 1986, aceaeaceaeappearedasasistertothelargecladｅｉｎ‐ appeared as a sister to the large clade in­ 1995).1995)．Theresultsofthepresentstudybasｅｄｏｎ The results of the present study based on cludingcludingothermemberoftheHookerialesandthe other member of the Hookeriales and the theｔｈｅ７６ｃＬｇｅｎｅｄｏｎｏｔｃｏｎｃｕｒｗｉｔｈthecircum- rbcL gene do not concur with the circum­ combinedcombinedHypnales-Leucodonta1es・Moredata Hypnales-Leucodontales. More data scriptionscriptionoftheHypnaceaebyNishimuraetal． of the Hypnaceae by Nishimura et a!. areareinneedtoclarifythephylogeneticpositionof in need to clarify the phylogenetic position of (1984).(1984)．Thisalsomeansthatthetaxonomiccon‐ This also means that the taxonomic con­ thePtychomniaceae，the Ptychomniaceae. fusionfUsionintheHypnalesmightbeascribedtodiver- in the Hypnales might be ascribed to diver­ TwoＴｗomajorcladesoftheHypnales（s」at.［＝ major clades of the Hypnales (s. lat. [= sifysifyoftheHypnaceaecurrentlyunderstoodasa of the Hypnaceae currently understood as a sensuｓｅｎｓｕＤｅＬｕｎａｅｔａＬ２０００；sensuBucketal De Luna et a!. 2000; sensu Buck et a!. largelargetaxonomicgroupcomprisingalargenumber taxonomic group comprising a large number 2000D:thecombinedHypnales(s・str.)andLeuc-2000]): the combined Hypnales (s. str.) and Leuc­ ofofspecies、 species. odontalesodontaleswiththeexceptionoftheCyrtopo‐ with the exception of the Cyrtopo­ NishimuraNishimuraetal.（1984)placedCZJﾉﾉﾉe堰o"eﾉﾉZJ et a!. (1984) placed Calliergonella daceae,daceae,andHookerialesfbnnasinglelargeclade and Hookeriales form a single large clade ininthesubfIamilyHypnoideaeofthefhmilyHyp- the subfamily Hypnoideae of the family Hyp­ withwithstrongsupport(96％LBPintheMLtree， strong support (96% LBP in the ML tree, naceae．Hedeniis（1990）transferredH)lp""籾naceae. Hedeniis (1990) transferred Hypnum 100%１００％supportinthcconsensustreeLasrepre‐ support in the consensus tree), as repre­ lindberghﾉﾉ"肋eZgjjMittintothegenusCMjeZgo"eﾉﾉα Mitt. into the genus Calliergonella sentedsentedinpreviousworks(DeLuna＆coworkers in previous works (De Luna & coworkers (Amblystegiaceae)(Amblystegiaceae)basedoncharactersincluding based on characters including 1999,2000;l999,2000;BucketaL2000)． Buck et a!. 2000). capsulecapsulestructure，alarceUsandpseudoparaphyl- structure, alar cells and pseudoparaphyl­ lia.lia、Ａｎｄｏ（1995)，however，disagreedwith Ando (1995), however, disagreed with 2.2．Familialrelationships，especiallytheHyp- Familial relationships, especially the Hyp­ Hedeniis（1990）andretainedH)lp""碗//"cIl6ergjjHedeniis (1990) and retained Hypnum lindbergii naceaenaceaeandpossiblyrelatedfamilies and possibly related families ininthegenuM卯""mTheobtainedtreeshows the genus Hypnum. The obtained tree shows Inhthepresentinvestigation,somefiamilies,such the present investigation, some families, such themonophylyfbrtheH1lp""腕ノノ"cIlbergjj-CtJﾉｰthe monophyly for the Hypnum lindbergii-Cal­ asastheSematophyllaceae（s」at.)，Plagiothecia‐ the Sematophyllaceae (s. lat.), Plagiothecia­ liergonellaﾉje堰o"eﾉﾉαｃ"Spjc/、αcladesupportingbyhigh cuspidata clade supporting by high ceaeceae（ｓｓｔｒ.）andFontinalaceae,areresolvedas (s. str.) and Fontinalaceae, are resolved as bootstrapbootstrapvalues（100％ＬＢＰｉｎＭＬｔｒｅｅ)，ａｌ‐ values (100% LBP in ML tree), al­ monophyleticmonophyleticgroups,whereasnoparticularclade groups, whereas no particular clade thoughthoughthecladechangeditspositionaccording the clade changed its position according isisrecognizedtoaccommodatemembersofthe recognized to accommodate members of the totopologiesThisresultwassuggestedbyto topologies. This result was suggested by familiesfamiliesHypnaceaeandLeucodontaceaeThe Hypnaceae and Leucodontaceae. The HedeniisHedeniis(1990)basedonthecharactersincluding (1990) based on the characters including FontinalaceaeFontinalaceaeareplacedatthebasalpartofthe are placed at the basal part of the capsulecapsulestructure，alarceUsandpseudoparaphyl- structure, alar cells and pseudoparaphyl­ combinedcombinedHypnalesandLeucodontalesclade． Hypnales and Leucodontales clade. liaIiaandTsubotaetal.（1999)basedonlbcLdata； and Tsubota et a!. (1999) based on rbcL data; andandconfinnedagaininthepresentstudy． confirmed again in the present study. H.HTsuBoTA,Ｔ・ARIKAwA,Ｈ・AKIYAMA,ＥＤＥＬｕＮＡ,Ｄ・GoNzALEz,MHIGucHIANDH､DEGucHI TSUBOTA, T. ARIKAWA, H. AKIYAMA, E. DE LUNA, D. GONZALEZ, M. HIGUCHI AND H. DEGUCHI 659６５９

Sematophyllaceae比marqp/1〕ﾉﾉﾉZJceae significantsignificantfbrtaxonomicreCognitionofthefam- for taxonomic recognition of the fam­ TheTheSematophyllaceaeisalsoalargetaxon- Sematophyllaceae is also a large taxon­ ilyilySematophyllaceae,asconcludedbyTsubotaet Sematophyllaceae, as concluded by Tsubota et omicomicgroupincludingnumerousspecies(withca group including numerous species (with ca al.aL(200ｌｂ)．Moreover,morerapidrateofnucle- (200 1b). Moreover, more rapid rate of nucle­ 5050genera:Vittl984),andthelimitsbetweenthe genera: Vitt 1984), and the limits between the otideotidesubstitutionsoftheSematophyllaceaｅ（S， substitutions of the Sematophyllaceae (s. HypnaceaeHypnaceaeandtheSematophyllaceaevaryamong and the Sematophyllaceae vary among lat.)lat.)wasdifferentfiomthatoftheotherlineage was different from that of the other lineage taxonomists.taxonomists，ＴａｎａｎｄＹｕ（1998,1999)madea Tan and Yu (1998, 1999) made a and,and,therefbre,characteristictothislineage・This therefore, characteristic to this lineage. This cladisticcladisticanalysisfbrthemorphologicaldata,and analysis for the morphological data, and isolationisolationmighthavebeencoITelatedtotheeco- might have been correlated to the eco­ showedshowedtheparaphylyoftheSematophyllaceae， the paraphyly of the Sematophyllaceae. logicallogicalrequirements，asthesetaxainthelineage requirements, as these taxa in the lineage TsubotaTsubotaandcoworkers（1999,2000,2001a,ｂ） and coworkers (1999, 2000, 2001a, b) withwithhabitatpreferencefbrgrowingontree habitat preference for growing on tree clarifiedclarifiedtheirphylogeneticrelationshipsinthe their phylogenetic relationships in the trunks.trunks． hypnobryaleanmosses，andshowedthemono-hypnobryalean mosses, and showed the mono­ iＰ phylyphylyoftheSematophyllaceae（s」at.；sensu of the Sematophyllaceae (s. lat.; sensu EntodontaceaeＥ'zroqlomaceae TsubotaTsubotaetaL2000，2001a,ｂ)．Arikawaand et al. 2000, 2001a, b). Arikawa and SeveralSeveraldiscussionshavebeenmadeonthere- discussions have been made on the re­ HiguchiHiguchi（2002),however,showedtheinclusion (2002), however, showed the inclusion lationshiplationshipoftheEntodontaceaeandThuidiaceae of the Entodontaceae and Thuidiaceae ofoftheEntodontaceaewithinthecladeoftheSe- the Entodontaceae within the clade of the Se­ tototheHypnaceaeandtheSematophyllaceaefiom the Hypnaceae and the Sematophyllaceae from matophyllaceaematophyllaceae(slat.),althoughthesupportwas (s. lat.), although the support was molecularviewpoints・Mizushima（1960）ａｎｄmolecular viewpoints. Mizushima (1960) and notsostrongnot so strong. BuckBuck（1980）publishedmonographicworkson (1980) published monographic works on DistinctionDistinctionbetweentheHypnaceａｅａｎｄＳｅ‐ between the Hypnaceae and Se­ theEntodontaceaaandtheyhavedifferentviewsthe Entodontaceae, and they have different views matophyllaceaehasbeencontroversiaLespeciallymatophyllaceae has been controversial, especially withregardtothesystematicpositiｏｎｏｆｔｈｅｇｅ－with regard to the systematic position of the ge­ concerningconcerningthephylogeneticpositionofsome the phylogenetic position of some nusｎｕｓＯ７ｒｈｏ/hecj"腕．Mizushima（1960)placedit Orthothecium. Mizushima (1960) placed it genera,genera,suchas〃/α〃α火ＩＩＰＡα(incB7oj/je7eﾉﾉα)， such as Pylaisiadelpha (inc. Brotherella), ininEntodontaceae,whileＢｕｃｋ(1980)intheHyp- Entodontaceae, while Buck (1980) in the Hyp­ Wijkia〃伽ａｎｄＨｂｊｅｍｐＭ/i"腕．ＤｅＬｕｎａｅｔａｌ． and Heterophyllium. De Luna et al. naceae.naceae，Ｖｉｔｔ（1984)，andBuckandGofTmet Vitt (1984), and Buck and Goffinet (2000)(2000)ａｌｓｏshowedtheroughconceptoftheSe‐ also showed the rough concept of the Se­ (2000)(2000)treatedthefamilyinanarrowsense,in‐ treated the family in a narrow sense, in­ matophyllaceae，althoughAQprer〕/ｇｊ"ｍｒｅ"e'wmmatophyllaceae, although Isopterygium tenerum cludingcludingonlythefburgenera:Ｅ"加巾"，ErWAro- only the four genera: Entodon, Erythro­ wouldwouldneedtobetransfelTedfiomtheHypnaceae、 need to be transferred from the Hypnaceae. dontium,c/o"〃腕，MCso"oc/O〃ａｎｄ〃/伽job'Ｗｍ．The Mesonodon and Pylaisiobryum. The ArikawaandHiguchi（1999）suggestedD/伽jaArikawa and Higuchi (1999) suggested Pylaisia presentsmdyshowedthattheEntodontaceae（spresent study showed that the Entodontaceae (s. polb′α"伽reported，ａｓ〃/α伽eﾉﾉａｐｏﾉ!〕ﾉα"伽，polyantha reported, as Pylaisiella polyantha, str.)str.)confinnstheexclusionofO7tAo/hecj"ｍａｎｄ confirms the exclusion of Orthothecium and shouldshouldbeconsideredintheSematophyllaceae be considered in the Sematophyllaceae. ＲＤｅ"血Sc/empocﾉﾉ"ｍｆｆｏｍｔｈｅｆａｍｉｌｙ,supportingPseudoscleropodium from the family, supporting OurOurresultshowedthemonophylyoftheSemato- result showed the monophyly of the Semato­ thethetreatmentofVitt(1984),andBuckandGoffi- treatment ofVitt (1984), and Buck and Goffi­ phyllaceae(slat.)proposedbyTsubotaandco-phyllaceae (s. lat.) proposed by Tsubota and co­ netnet（2000）ｏｒ/hoZhecj"腕ノW/bsce"sfbrmeda (2000). Orthothecium rufescens formed a workersworkers（2000,2001a,ｂ),includingthegenera (2000, 2001a, b), including the genera cladecladewithhOprerygmp川加"eﾙｧﾉα"α,and with Isopterygiopsis muelleriana, and D伽jα火ＩＩＰﾉｶα，附加ａｎｄＨｅ〃qp/Ｍｊ川andPylaisiadelpha, Wijkia and Heterophyllium, and 囚ye"dmc/empodj"ｍｐ"7"mappearedinthecladePseudoscleropodium purum appeared in the clade somesomespeciespreviouslytreatedasmembersof species previously treated as members of withwithmembersoftheBrachytheciaceae、Thetree members of the Brachytheciaceae. The tree theHypnaceae：Ｈｊｌｐ""加伽ro-v"枕,ZFOpjer〕ノーthe Hypnaceae: Hypnum tristo-viride, Isoptery­ alsoalsoshowsthatthecladeco､sistingoftheSe- shows that the clade consisting of the Se­ giumｇｊ"腕／e"e'ｗｍａｎｄ〃/αｊｓｊａｐｏﾉ!〕ﾉα"ﾉﾉ､,ｗａｓｒｅ－ tenerum and Pylaisia polyantha, was re­ matophyllaceae（s」at．［＝sensuTsubotaetaLmatophyllaceae (s. lat. [= sensu Tsubota et al. ｖｅａｌｅｄｂｙｂｏｔｈｔｈｅｍｏｓｔＭＬｔreewithstrongsup-vealed by both the most ML tree with strong sup­ 2000,200q2001a,ｂ]),EntodontaceaeandThuidiaceae 2001 a, b]), Entodontaceae and Thuidiaceae port(96％LBP)andtheconsensustreeWecan-port (96% LBP) and the consensus tree. We can­ wassupportedwithhighLBP(97％LBPinMLwas supported with high LBP (97% LBP in ML Ｐ notsupportthephylogeneticpositionofP､pobノーnot support the phylogenetic position of P. poly­ tree)．Vitt(1984)placedtheEntodontaceaeclosetree). Vitt (1984) placed the Entodontaceae close anthaα"rAaonthebasisofthedatapresentlyavailable on the basis of the data presently available. totheSematophyllaceaeonthebasisofmorphol-to the Sematophyllaceae on the basis of morphol­ Further,Further,itisnecessarytoverifytheidentification it is necessary to verify the identification ogicalogicalandecologicalcharactersOurresultsup- and ecological characters. Our result sup­ ofofthematerialofthespecies・ the material of the species. portsthetreatmentoftheEntodontaceae,butnotports the treatment of the Entodontaceae, but not AspreviouslyindicatedinmorphologicaldataAs previously indicated in morphological data thetheThuidiaceae． Thuidiaceae. analysesanalysesbyHedeniis（1995,1996),phylogenetic by Hedenas (1995, 1996), phylogenetic relationshipsrelationshipsbasedonmoleculardatashouldbe based on molecular data should be Ｐﾉagjoj/iecmceaePlagiotheciaceae surveyedsurveyedbetweentheSematophyllaceaeandthe between the Sematophyllaceae and the TheThetreeshowedthecladeconsistingofonly tree showed the clade consisting of only familiesfamiliesintheorderHookeriales・Themono- in the order Hookeriales. The mono­ thegenusP/αgjorhecj"mnearthecladesinclud-the genus Plagiothecium near the clades includ­ phylyoftheSematophyllaceaeandtheHookeri-phyly of the Sematophyllaceae and the Hookeri­ ingingtheBrachytheciaceaeandsomemembersof the Brachytheciaceae and some members of alesaleshadnophylogenicevidenceinouranalyses had no phylogenic evidence in our analyses. theHylocomiaceae・Recently，Pedersenandthe Hylocomiaceae. Recently, Pedersen and TheThepresentsmdyrevealsatleastthatthechar- present study reveals at least that the char­ Hedeniis(2002)showedthephylogenyofthePla-Hedenas (2002) showed the phylogeny of the Pla­ actersactersofwell-developedalarcellsandhabitatare of well-developed alar cells and habitat are giotheciaceaegiotheciaceaebasedonmorphologicaｌｄａｔａａｎｄ based on morphological data and 660660 HikobiaＨｉｋｏｂｉａＶｏＬｌ３,No.４，２００２ Vol. 13, No.4, 2002 twotwoDNAregions:ノアｓ４ａｎｄ伽Ｌ－伽F,suggesting DNA regions: rps4 and trnL-trnF, suggesting ogyogyandtreatedtheAmblystegiaceaewithanar- and treated the Amblystegiaceae with a nar­ thethefamilywithawidecircumscriptionArikawa family with a wide circumscription. Arikawa rowerrowersenseThepresenttreeconfinnsthisnar- sense. The present tree confinns this nar­ andandHiguchi(2002)clarifiedthephylogeneticpo‐ Higuchi (2002) clarified the phylogenetic po­ rowerowefamilialcircumscription，placing4"αcam- familial circumscription, placing Anacam­ sitionsitionofthegeneraA0prerygj叩巾andHblzo- of the genera Isopterygiopsis and Herzo­ ptodonptoqlO〃intheAmblystegiaceae． in the Amblystegiaceae. giella,gjeﾉﾉα，andproposedtheinclusionofAOpZer)Igj- and proposed the inclusion of Isopterygi­ opsisqpsなwithinthePlagiotheciaceaeThepresent within the Plagiotheciaceae. The present Brachytheciaceae〃αc/iﾘﾉrAecjaceae investigationinvestigationdidnotsupportthesepreviousanal- did not support these previous anal­ TheThepresentsmdyshowedthemonophylyof present study showed the monophyly of yses,yses,discoveringthemonophylyofthePlagiothe- discovering the monophyly of the Plagiothe­ thetheBrachytheciaceae，includingOAzJ〃"ｍ１ｅａａｎｄ Brachytheciaceae, including Okamuraea and ciaceaeciaceaeincludingonlythegenusP/αgjorAecj"腕． including only the genus Plagiothecium. HelicodontiumＨｂ/jco血""z"'ｚａｓｗｅｌｌａｓＲｓｅ"ﾛﾉＤＳＣ/empoい""Z as well as Pseudoscleropodium andandexcluding7bme"卯""腕.Ｈセノjcoc/Wz伽mwas excluding Tomentypnum. Helicodontium was StereophyllaceaeS〃e叩Ｍ/αceae previouslypreviouslytreatedintheMyriniaceaebyVitt treated in the Myriniaceae by Vitt ArikawaArikawaandHiguchi（1999),andthefbllow‐ and Higuchi (1999), and the follow­ (1984)(1984)andBuckandGoffinet(2000)．Ourresult and Buck and Goffinet (2000). Our result ingingstudies（e9.,Pedersen＆Hedeniis2002； studies (e.g., Pedersen & Hedeniis 2002; supportssupportstheplacementoｆＡｅ"cﾉＤＳＣ/empo`j"m the placement of Pseudoscleropodium ArikawaArikawa＆Higuchi2002)clarifiedtheexclusion & Higuchi 2002) clarified the exclusion bybyBuckandGoffinet(2000),ｂｕｔｎｏｔＨｂ"ＣＯＣ/b"‐ Buck and Goffinet (2000), but not Helicodon­ ofoftheStereophyllaceaeffomthePlagiotheci- the Stereophyllaceae from the Plagiotheci­ tium.〃'７０．ＴｈｅｇｅｎｅｒａＰ/αgjor/iecj"腕,Ｐ/e"rozj"mand The genera Plagiothecium, Pleurozium and aceae,aceae,andthemonophylyoftheStereophyllaceae and the monophyly of the Stereophyllaceae fMoco〃ａｓﾉﾉwmwereplacedinacommoncladeHylocomiastrum were placed in a common clade basedonmolecularphylogeneticanalysesThebased on molecular phylogenetic analyses. The withthetaxaoftheBrachytheciaceae(excludedwith the taxa of the Brachytheciaceae (excluded presentresultcorroboratedthepreviousworkspresent result corroborated the previous works TomenthypnumTb"Te"/ﾉ!〕lp""〃〃"e"s）ｗｉｔｈ６６％ＬＢＰｉｎＭＬ nitens) with 66% LBP in ML showingshowingtheStereophyllaceａｅａｓａｓｉｓｔｅｒｔｏｔｈｅ the Stereophyllaceae as a sister to the ｔｒｅｅ．tree. Tomentypnum-７ｂｍｅ"卯""加一 ﾉVbo巾ﾉﾉc/ＩＣ加加clade，althoughtheirphyloge-Neodolichomitra clade, although their phyloge­ MCjeo7mceaeMeteoriaceae neticpositionisnotsostableintheHypnales(s、netic position is not so stable in the Hypnales (s. HuttunenHuttunenandIgnatov(perscomm.)suggested and Ignatov (pers. comm.) suggested lat.lat.[＝sensuDeLunaetaL2000;sensuBucket [= sensu De Luna et al. 2000; sensu Buck et thethecloserelationshipbetweentheBrachythecia- close relationship between the Brachythecia­ al.aL2000])clade． 2000]) clade. ceaeceaeandMeteoriaceaeintheirpreliminalyanaly- and Meteoriaceae in their preliminary analy­ ses.ses・Ourpresentanalysisincludesonlyonespe- Our present analysis includes only one spe­ HMocomaceaeHylocomiaceae ciesoftheMeteoriaceae・Furthersequencesofcies of the Meteoriaceae. Further sequences of lntheMLtree,theHylocomiaceaeappearedasIn the ML tree, the Hylocomiaceae appeared as theMeteoriaceaeareneededtoshowtherelation-the Meteoriaceae are needed to show the relation­ twodistinctclades:Ｒ/Uﾉ肋α火llpAMD-fMoco加川two distinct clades: Rhytidiadelphus-Hylocomium shipsshipsbetweenthetwofamilies． between the two families. 一LoalAeo6'Ｗｍ－ｄｅ"jcﾉﾉ"腕cladeatthebasalpo--Loeskeobryum-Ctenidium clade at the basal po­ sitionsitionofthecombinedseveralclades,suchasthe of the combined several clades, such as the Thuidiaceae、"伽aceαｅα"ＣＭ"o"zodo"raceae and Anomodontaceae BrachytheciaceaecladeａｎｄｔｈｅＰ/CJgjorAecj"mBrachytheciaceae clade and the Plagiothecium TheTheThuidiaceae(s・str.[＝sensuTsubotaetaL Thuidiaceae (s. str. [= sensu Tsubota et al. clade;ａｎｄＰ/e"'℃ｚｊ"ｍ－Ｈｙ/ＯＣＣ〃ａｓ〃ｍｃｌａｄｅａｓclade; and Pleurozium-Hylocomiastrum clade as 2000]),consistingofonlythegenusT1h"ｊ`ｊ"川，2000]), consisting of only the genus Thuidium, ｔｈｅｓｉstercladetothePmgjoj/iecj"mcladcAre-the sister clade to the Plagiothecium clade. A re­ fbnnedasinglecladebeingsistertotheEntodon-fonned a single clade being sister to the Entodon­ viewofthcirfamilialcircumscriptionisneededview of their familial circumscription is needed taceae(s・str.[＝sensuVittl984;sensuBuck＆taceae (s. str. [= sensu Vitt 1984; sensu Buck & １ fbrthegeneraP/e"７ozJ"mandfMoco〃ａｓ加川．for the genera Pleurozium and Hylocomiastrum. Goffinet2000])，ａｓｓｈｏｗｎｂｙＴｓｕｂｏｔａｅｔａＬGoffinet 2000]), as shown by Tsubota et al. (2001a)．肋ｊｅ""eﾉﾉZza6jerj"αａｎｄＢｏ"/ZlO′α〃ﾉｰ(2001 a). Abietinella abietina and Boulaya mit­ 伽6(WegjaceaeAmblystegiaceae /emappearedoutofbutcloseto,thecladeofthetenii appeared out of, but close to, the clade of the Thephylogenetictreesinthepresentstudydis-The phylogenetic trees in the present study dis­ ThuidiaceacThuidiaceae. coveredpolyphylyoftheAmblystegiaceae（scovered polyphyly of the Amblystegiaceae (s. Thegenera比p/o/j〕ﾉ腕e"伽ａｎｄ』"omo巾"，The genera Haplohymenium and Anomodon, lat.）aspreviouslysuggestedbytraditionaltax-lat.) as previously suggested by traditional tax­ ｅｘｃｅｐｔｆｂｒＡｇｉ７ａｒＺﾉﾉﾉ，appearedinthecladein-except for A. girardii, appeared in the clade in­ onomists（e､９，Hedeniisl997)，showingthreeonomists (e.g., Hedeniis 1997), showing three cludingmembersoftheNeckeraceae，fbrmingacluding members of the Neckeraceae, fonning a clades:ＣＭＯ"e"ro"-DreYﾌα"oc/αc/"s-Hyg7oα"'‐clades: Cratoneuron-Drepanocladus-Hygroam­ resolvedcladeplacingthesecondalybasalposi-resolved clade placing the secondary basal posi­ 611〕Mgj"〃一A"αcα叩/o血〃clade,ＣａﾉﾉjeZgo"eﾉﾉablystegium-Anacamptodon clade, Calliergonella tioninthecombinedHypnales-Leucodontalestion in the combined Hypnales-Leucodontales. clade,ａｎｄ７１ｏｍｅ"卯"""Zclade，BuckandGoffi‐clade, and Tomentypnum clade. Buck and Goffi­ MaedaetａＬ（2000）alreadyclarifiedthepoly‐Maeda et al. (2000) already clarified the poly­ net(2000)treatedthefiamilywithanarrowSense，net (2000) treated the family with a narrow sense, phylyoftheAnomodontaceae，showingacladephyly of the Anomodontaceae, showing a clade excludingsomegeneratreatedinthepresentexcluding some genera treated in the present withHZZp/o/i)ﾉｍｅ"j"耐,比Ｚｐａｊ"e"ro〃ａｎｄｓｏｍｅwith Haplohymenium, Herpetineuron and some analysis,ｅ,9.,CMjeHgo"eﾉﾉα(inc.ｆ卯""腕ﾉﾉ"‘/Lanalysis, e.g., Calliergonella (inc. Hypnum lind­ Ａ"o"zo吻刀species・ThesefactsindicatedthatAnomodon species. These facts indicated that 6e卿/）ａｎｄｎｍｅ"卯""腕，ｆｉ･omthefiamily・bergii) and Tomentypnum, from the family. somegenera,suchasHZJpM〕ﾉ腕e"伽ａｎＭ"o-some genera, such as Haplohymenium and Ano­ Vanderpoortenetal.（2002）showedaphyloge-Vanderpoorten et al. (2002) showed a phyloge­ mo血〃whicharesometimestreatedwithinthemodon which are sometimes treated within the netictreebasedonDNAsequencesandmolphol-netic tree based on DNA sequences and morpho 1- Thuidiaceae,shouldbedividedintotwofamilies：Thuidiaceae, should be divided into two families: H.HTsuBoTA,Ｔ・ARIKAwA,Ｈ・ＡＫⅣAMA,ＥＤＥＬｕＮＡ,Ｄ､GoNzALEz,MHIGucHIANDH・DEGucHI６６１ TSUBOTA, T. ARIKAWA, H. AKIYAMA, E. DE LUNA, D. GONZALEZ, M. HIGUCHI AND H. DEGUCHI 661

thetheThuidiaceaeandAnomodontaceae，support- Thuidiaceae and Anomodontaceae, support­ tiontionoftheFontinalaceae． of the Fontinalaceae. ingingthesystemofMaedaeｔａＬ(2000),andBuck the system of Maeda et al. (2000), and Buck andandGoffinet(2000)． Goffinet (2000). CyrtopodaceaeqWQpodnceae TheTheCyrtopodaceaeweretreatedintheLeuco‐ Cyrtopodaceae were treated in the Leuco­ Leucodontaceae,比"ＣＯ伽"mceae，CrXpAaeaceae，α"‘Pjero6r〕ﾉa- Cryphaeaceae, and Pterobrya­ dontalesdontalesbyVitt（1984)，ａｎｄＢｕｃｋａｎｄＶｉｔｔ by Vitt (1984), and Buck and Vitt ceaeCeae (1986).(1986)Jnthepresentinvestigation,theCyrtopo- In the present investigation, the Cyrtopo­ MaedaMaedaetaL(2000)alsodiscussedthefamilial et al. (2000) also discussed the familial daceae,daceae,representedbyB“Ｃｈｅ'eﾉﾉﾉａ６７ｅｖｉ/bﾉﾉα,Ｂ、 represented by Bescherellia brevi/olia, B. affinityaffinityofFolTsrroe〃α,showingrelationshipbe- of Forsstroemia, showing relationship be­ elegantissima,e/egzJ"伽加α,ａｎｄＣＷｍｐ"sMos"s,appearedin and Cyrtopus setosus, appeared in tweentweentheCryphaeaceaeandLeucodontaceae・As the Cryphaeaceae and Leucodontaceae. As aapositionoutsidetheHypnales(s」at.［＝sensu position outside the Hypnales (s. lat. [= sensu theytheytreatedonlytheleucodontaleanmossesfbr treated only the leucodontalean mosses for DeDeLunaetaL2000;BucketaL2000])-Hookeri- Luna et al. 2000; Buck et al. 2000])-Hookeri­ theirtheiranalysis,thephylogeneticpositionfbrmem- analysis, the phylogenetic position for mem­ alesalesclade,fionninganotableclade(100％ＬＢＰin clade, forming a notable clade (100% LBP in ’ bersbersoftheorderinthepleurocarpswasnotso of the order in the pleurocarps was not so thetheMLtree）sistertotheHypnodendraceae・ ML tree) sister to the Hypnodendraceae. clear.clear・hthepresenttreesbasedonthelarge-scale In the present trees based on the large-scale BuckBuckandGofY1net（2000）treatedtheCyrtopo- and Goffinet (2000) treated the Cyrtopo­ datadataset，thetraditionalleucodontaleanmosses set, the traditional leucodontalean mosses daceaedaceaeintheRhizogoniales,andthepresenttree in the Rhizogoniales, and the present tree formfbrmfiourma]ｏｒａｎｄｓｏｍｅｍｉｎｏｒｃｌａｄｅｓｉｎｔｈｅ four major and some minor clades in the supportssupportstheirplacementofthefamily． their placement of the family. combinedｃｏｍｂｉnedHypnales-Leucodontales，supporting Hypnales-Leucodontales, supporting thethethreecladespresentedbyMaedaetaL(2000)． three clades presented by Maeda et al. (2000). ﾉVCcAe'αceaeα'１．ｋ肋QpﾉＭ/αceaeNeckeraceae and Lembophyllaceae TheThemembersoftheLeucodontaceae，exceptfbr members of the Leucodontaceae, except for TheTheotherrnembersofthetraditionalleucodon‐ other members of the traditional leucodon­ Fo'Fsj7oe〃α，CryphaeaceaeandPterobryaceaeForsstroemia, Cryphaeaceae and Pterobryaceae taleantaleanmosses,previouslytreatedastheNeckera- mosses, previously treated as the Neckera­ formfbrmacladewithinthecladeoftheHypnales(s・ a clade within the clade of the Hypnales (s. ceae,ceae,appearedasamajorcladewith86％LBP appeared as a major clade with 86% LBP lat.lat.[＝sensuDeLunaetaL2000;sensuBucket [= sensu De Luna et al. 2000; sensu Buck et support,support,asasistertotheLembophyllaceaeclade・ as a sister to the Lembophyllaceae clade. al.aL2000])．Fol《Ｗｏｅｍｚａ,amemberofthetradi- 2000]). Forsstroemia, a member of the tradi­ InIntheclade，ＥｃＡｊ"o血"腕〃m6ros"ｍａｓｗｅｌｌａｓ the clade, Echinodium umbrosum as well as tionaltionalLeucodontales，isplacedinthecladenear Leucodontales, is placed in the clade near Ａ"o"TOC/O〃ｇ〃αﾉﾋI/ｊｊａｎｄＦｏ応sｊ'Ce"zjaspp、wereAnomodon giraldii and Forsstroemia spp. were theNeckeraceaeTheseresultsdidnotsupportthe Neckeraceae. These results did not support alsoalsoincluded included. thetheappearanceoftheleucodontaleanfamiliesin appearance of the leucodontalean families in thecombinedHypnales（s・ｓｔｒｌ＝sensuVittthe combined Hypnales (s. str. [= sensu Vitt C/〃αcjaceaeα"c/P/e"rozmpMﾉﾋJceaeClimaciaceae and Pleuroziopsidaceae 1984])l984])andLeucodontalesbyBucketaL(2000)． and Leucodontales by Buck et al. (2000). TheClimaciaceaeandPleuroziopsidaceaeThe Climaciaceae and Pleuroziopsidaceae fbnnedaweakclade(57％LBP,９１％)neartheformed a weak clade (57% LBP, 91%) near the Ｆａ６７ｏ"mceaeFabroniaceae Stereophyllaceae.Stereophyllaceaelreland（1968）segregated Ireland (1968) segregated TheTheFabroniaceaefbrmnonotablecladeinthe Fabroniaceae form no notable clade in the P/e"γozjqpsLsfiomtheClimaciaceaeintothemon-Pleuroziopsis from the Climaciaceae into the mon­ MLandconsensustrees，asshowntbeaboveML and consensus trees, as shown the above. otypicfamilyPleuroziopsidaceaeNorrisandlgna-otypic family Pleuroziopsidaceae. Norris and Igna­ Furtherinvestigationswithadditionaltaxafi･omFurther investigations with additional taxa from tov(2000)observedthestemsurfaceanatomyintov (2000) observed the stem surface anatomy in otherothergenera，ｓｕｃｈａｓＦｃＪ６ｒｏ"ｊα，ＳｂＡｗ伽cﾊﾙeα， genera, such as Fabronia, Schwetschkea, ClimaciumC/〃αcj"mandP/e"mzmp川,andsuggestedthat and Pleuroziopsis, and suggested that

, andａｎｄＳｂｈｗｅなcAAcOpsjSareneeded． Schwetschkeopsis are needed. P/e"'ozmpsjSshouldbereturnedtotheClimaci-Pleuroziopsis should be returned to the Climaci­ aceacOurtreesdonotconfIictwiththeirsug-aceae. Our trees do not conflict with their sug­ Ｆｏ"""α/αceaeF ontinalaceae gestiononP/e"rozjOpsjS，althoughthevalueofgestion on Pleuroziopsis, although the value of AlthoughonlyFo"/ｊ"α/jJspecieswerein- Ｐ Although only Fontinalis species were in­ thesupportisｎｏｔｓｏｈｉｇｈthe support is not so high. cludedintheanalysis，theFontinalaceaeappearcluded in the analysis, the Fontinalaceae appear asasingleclade,atthemostbasalpositionintheas a single clade, at the most basal position in the 3.Singlecostaanddoublecosta3. Single costa and double costa Hypnales-Leucodontalesclade，supportingtheHypnales-Leucodontales clade, supporting the Vitt1ssystem（1984）fbrtheclassiflcationofVitt's system (1984) for the classification of conclusionofVitt（1984）ａｎｄＢｕｃｋ＆Ｖｉｔｔconclusion of Vitt (1984) and Buck & Vitt hypnaleanmossessuggeststhatthegroupswithahypnalean mosses suggests that the groups with a (1986）Ｂｕｃｋ＆Vitt（1986）constructedanew(1986). Buck & Vitt (1986) constructed a new singlecosta，asrepresentedbytheBrachythecia-single costa, as represented by the Brachythecia­ classificationfbrpleurocaIpousmosses，recir-classification for pleurocarpous mosses, recir­ ceaeandAmblystegiaceae，wouldhavedivergedceae and Amblystegiaceae, would have diverged cumscribingtheHypnales（s・str.）includingthecumscribing the Hypnales (s. str.) including the fi･omthehypnaleanancestoratanearlyevolu-from the hypnalean ancestor at an early evolu­ HypnodendrineaeandFontinalineaeOuranaly-Hypnodendrineae and Fontinalineae. Our analy­ tionarystage,fiollowedbythegroupswithadou‐tionary stage, followed by the groups with a dou­ sessupporttheinclusionoftheFontinalineaeses support the inclusion of the Fontinalineae blecosta，ａｓrepresentedbytheHypnaceae，Se-ble costa, as represented by the Hypnaceae, Se­ withintheHypnales，butnottheinclusionofthewithin the Hypnales, but not the inclusion of the matophyllaceaeandEntodontaceae,whichhere-matophyllaceae and Entodontaceae, which he re­ Hypnodendrineae、FurtheranalysisincludingHypnodendrineae. Further analysis including gardedasthemostrecentevolutionarystageofgarded as the most recent evolutionary stage of somespeciesoftheFontinalaceae,especiallyａｎ‐some species of the Fontinalaceae, especially an­ theorderHypnales（ｓｓｔｒ.［＝sensuVittl984]）the order Hypnales (s. str. [= sensu Vitt 1984]) othergenusDjchel【Wlm,wouldclarifytheresolu‐other genus Dichelyma, would clarity the resolu- (seealsoTsubotaetaLl999;BucketaL2000)．(see also Tsubota et al. 1999; Buck et al. 2000). 662 ＨｉｋｏｂｉａＶｏｌｌ３,No.４，２００２Hikobia Vo!. 13, No.4, 2002

lnthepresentinvestigation,taxawithadoubleIn the present investigation, taxa with a double Englishsummary）English summary). costaappearinvariouscladesofthetree・Thiscosta appear in various clades of the tree. This Arikawa,Ｔ､＆Higuchi,Ｍ１999．Phylogeneticanaly-Arikawa, T. & Higuchi, M. 1999. Phylogenetic analy­ resultsuggeststhatasinglecostaisanancestralresult suggests that a single costa is an ancestral sisofthePlagiotheciaceae(MuscOanditsrelativessis of the Plagiotheciaceae (Musci) and its relatives character,andthatthereweremultipletransitionscharacter, and that there were multiple transitions basedon/御6cLgenesequencesCryptogamie,Ｂｒｙ－based on rbcL gene sequences. Cryptogamie, Bry­ toadoublecostaTherefbre，ourresultsoftheto a double costa. Therefore, our results of the ｏＬ２０:２３１－２４５．o!. 20: 231-245. presentstudydonotconcurwiththesuggestionpresent study do not concur with the suggestion --- ＆Z002Phylogeneticpositionofthe& --- 2002. Phylogenetic position of the proposeｄｂｙＶｉｔｔ（1984)，ａｎｄＢｕｃｋａｎｄＶｉｔｔproposed by Vitt (1984), and Buck and Vitt generaLsOpte'f〕/gjOpsjFand比ﾉazOgjeﾉﾉα（Musci）genera Isopterygiopsis and Herzogiella (Musci) (1986)．OuranalysesconfirmtheHndingsbyDe(1986). Our analyses confirm the findings by De basedonlbcLgenesequences・BryoLRes．（inbased on rbcL gene sequences. Bryol. Res. (in LunaetaL(2000)andBucketaL(2000)thattheLuna et al. (2000) and Buck et al. (2000) that the press）press). Hypnales，togetherwithmostoftheleucodon-Hypnales, together with most of the leucodon­ Buck,Ｗ,Ｒ」９８０AgenericrevisionoftheEntodon-Buck, W. R. 1980. A generic revision of the Entodon­ taleangroups，fbnnamonophyleticsistergrouptalean groups, form a monophyletic sister group taceae．』・HattoriBotLab48:７１－１５９．taceae. J. Hattori Bot. Lab. 48: 71-159. tothehookerialeanlineage・Resultsalsoshowto the hookerialean lineage. Results also show -- &＆Goffmet,Ｂ、2000．Morphologyandclas‐ Goffinet, B. 2000. Morphology and clas­ 'II､ thatthefamilialcircumscriptionwithintheHyp-that the familial circumscription within the Hyp­ sificationofmosses・Ｉｎ:Ａ､Ｊ・Ｓｈａｗ＆BGofHnetsification of mosses. In: A. J. Shaw & B. Goffinet nalesandLeucodontalesshouldbereconsidered．nales and Leucodontales should be reconsidered. (eds.),BryophyteBiology,ｐｐ７１－１２３・Cambridge(eds.), Bryophyte Biology, pp. 71-123. Cambridge UniversityPress,Cambridge．University Press, Cambridge. ---－＆Vitt,，.Ｈ」９８６SuggestionsfOranewfa-& Vitt, D. H. 1986. Suggestions for a new fa­ Acknowledgment milialclassificationofpleurocarpousmossesTax-milial classification of pleurocarpous mosses. Tax­ on35:２１－６０ Thispaperisthecombinedresultofaninterna-This paper is the combined result of an interna­ on 35: 21-60. tionalcollaborativeeffbrt、Ｗｅｗｏｕｌｄｌｉｋｅｔｏａｃ－tional collaborative effort. We would like to ac­ ---,－，Goffinet，Ｂ＆Ｓｈａｗ，Ａ・I2000a・Testing Goffinet, B. & Shaw, A. J. 2000a. Testing ｋｎｏｗｌｅｄｇｅＲ．Ｄ・SeppeltfbrcheckingtheEng-knowledge R. D. Seppelt for checking the Eng­ morphologicalconceptsofordersofpleurocarpousmorphological concepts of orders of pleurocarpous mosses (Bryophyta) using phylogenetic reconstruc­ lishtextandhelpfnlsuggestions;BGoffinetandlish text and helpful suggestions; B. Goffinet and mosses(Bryophyta)usingphylogeneticreconstruc- tions based on trnL-trnF and rps4 sequences. Mol. W．Ｒ・BuckfbrredressingtheEnglishandfiorW. R. Buck for redressing the English and for tionsbasedon/r"L-ﾉﾉ侭"Ｆａｎ。'ips4sequences・ＭｏＬ ＰｈｙＬＥｖｏＬｌ６：１８０－１９８．Phyl. Evol. 16: 180-198. theircommentsandsuggestionｓｏｎａｎｅａｒｌｉｅｒｖｅr-their comments and suggestions on an earlier ver­ ---, --- &＆Z000hNovelrelation‐ --- 2000b. Novel relation­ sionsionofthispaper;ＭＳ・IgnatovandSHutmnen of this paper; M. S. Ignatov and S. Huttunen shipsinpleurocarpousmossesasrevealedbycp-ships in pleurocarpous mosses as revealed by cp­ forfbrtheircommentsandsuggestionsonthephylo‐ their comments and suggestions on the phylo­ DNADNAsequences・BryologistlO3：774-789． sequences. Bryologist 103: 774-789. geneticgenetictrees;andBEstebanezfbrhergreathelp trees; and B. Est6banez for her great help. Cox,ＣＯＸ,ＣＪ.,Goffinet,Ｂ,Newton,Ａ､Ｅ､，Ｓｈａｗ,ＡＪ＆ C. J., Goffinet, B., Newton, A. E., Shaw, A. J. & WeWeａｌｓｏｔｂａｎｋＳＭａｅｄａａｎｄＫ・ＫＣsugewith also thank S. Maeda and K. Kosuge with Hedderson，Ｔ，Ａ､12000．Phylogeneticrelation-Hedderson, T. A. J. 2000. Phylogenetic relation­ whomwhomwehavecollaborated，MHasebeand，T we have collaborated, M. Hasebe and, T. shipsshipsamongthediplolepideous-altematemosses among the diplolepideous-aitemate mosses NishiyamaNishiyamafbrtechniqueofexperimentandad- for technique of experiment and ad­ (Bryidae)（Bryidae）inferredfiPomnuclearandchloroplast inferred from nuclear and chloroplast viceviceaboutphylogeneticanalysis,ａｎｄＮ・Minaka about phylogenetic analysis, and N. Minaka DNADNAsequences．BryologistlO3:224-241． sequences. Bryologist 103: 224-241. andandMMiyafbrusageofPAUPRat・Researchon M. Miya for usage ofPAUPRat. Research on ＤｅＬｕｎａ,Ｅ,Newton,Ａ､Ｅ，Withey,Ａ､，Gonzalez,，．De Luna, E., Newton, A. E., Withey, A., Gonzalez, D. molecularsystematicsofmosseswasfUndedbymolecular systematics of mosses was funded by &＆Mishler，ＢＤ、１９９９．Thetransitiontopleuro- Mishler, B. D. 1999. The transition to pleuro­ thefbllowinggrants:KAKENHl（Japan）grantsthe following grants: KAKENHI (Japan) grants carpy:carpy：Aphylogeneticanalysisofthemaindiplo- A phylogenetic analysis of the main diplo­ 09839031O9839031toH､Akiyama,Ｏ９０４１１６５ｔｏＪＭｕｒａｔａ， to H. Akiyama, 09041165 to 1. Murata, 、 lepidouslepidouslineagesbasedonr6cLsequencesandmor- lineages based on rbcL sequences and mor­ 11640698ｌｌ６４０６９８ｔｏＨ・Deguchi，l3640707toMHigu- to H. Deguchi, 13640707 to M. Higu­ phology.phology・BryologistlO2:634-650. Bryologist 102: 634-650. chi,chi，ａｎｄｌ３７４０４９０ｔｏＨ・Tsubota；ａｎｄＣＯＮＡ－ and 13740490 to H. Tsubota; and CONA­ ---,,Buck,ＷＲ.,Akiyama,Ｈ,Arikawa,Ｔ､,Tsubo- Buck, W. R., Akiyama, H., Arikawa, T., Tsubo­ CYT (Mexico) grant 27400-N to E. De Luna et ＣＹＴ(M6xico)grant27400-NtoEDeLunaet ta,ｔａ,Ｈ､，Gonzalez,，.,Newton,Ａ､Ｅ＆Ｓｈａｗ,Ａ､Ｊ、 H., Gonzalez, D., Newton, A. E. & Shaw, A. 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Hattori Bot. Lab. 88: 79-99. chichi39333（TNS),thissmdy;Climaciaceaedj"、c伽１ 39333 (TNS), this study; Climaciaceae Climacium ---,－，Akiyama，Ｈ､，Yamaguchi，Ｔ＆Deguchi，Ｈ、 Akiyama, H., Yamaguchi, T. & Deguchi, H. dendroidesdeノルojm（Hedw.）ＦＷｅｂｅｒ＆DMohr,ABO19442, (Hedw.) F.Weber & D.Mohr, AB019442, 2001aMolecularphylogenyoftheSematophyl-2001a. Molecular phylogeny of the Sematophyl­ Japan,Honshu,MiePref,Nabari-shLNagaki,MarchJapan, Honshu, Mie Pref., Nabari-shi, Nagaki, March laceae(Hypnales,Musci)basedonchloroplastlbcLlaceae (Hypnales, Musci) based on chloroplast rbcL l6，1998,HAkiyamal4150（HYo),thissmdy;Ｃ〃16, 1998, H. Akiyama 14150 (HYO), this study; Cli­ sequences.sequences．Ｊ・HattoriBotLab90:２２１－２４０． J. Hattori Bot. Lab. 90: 221-240. "１αc川zﾉ叩o"jc""ZLindh,ABO19443,Japan,Honshu，maciumjaponicum Lindb., AB019443, Japan, Honshu, --,--,--&--＆２００ｌｂ・Molecular2001b. 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Schuster (ed.), New Manual of Bryology, Japan，Honshu，HyogoPref，Kamigori-chqJapan, Honshu, Hyogo Pref., Kamigori-cho, ＶＯＬ２，ｐｐ､696-759．HattoriBot・Ｌａｂ,NichinanVol. 2, pp. 696-759. Hattori Bot. Lab., Nichinan. Onaru-keikoku,２５０ｍａｌｔ.,Ｊｕｎｅ１７，１９９８，HAkiyamaOnaru-keikoku, 250 malt., June 17, 1998, H. Akiyama l4234（HYo)，ｔｈｉｓstudy;ノVecAcrqp伽〃ｊ/ｊｑＭａ（Mitt.）14234 (HYO), this study; Neckeropsis nitidula (Mitt.) AppendixA・InfbnnationonspecimensfiomwhichAppendix A. Information on specimens from which MFleisch，ＡＢＯ94790,Japan，Honshu，OkayamaM. Fleisch., AB094790, Japan, Honshu, Okayama DNAwasextractedinthissmdy・DNA was extracted in this study. PrefNishiawakura-son,WakasugiPrimaryForest,Pref., Nishiawakura-son, Wakasugi Primary Forest, Hereisalistoftaxawhoser6cLsequenceswereuti-Here is a list of taxa whose rbcL sequences were uti­ ＬＯＯＯｍａｌｔ.,June17,1998,HAkiyamal4210(HYo)，1,000 m alt., June 17, 1998, H. Akiyama 14210 (HYO), lizedinthissmdywiththeiraccessionnumbers,show-lized in this study with their accession numbers, show­ thisstudy;Ｐｊ""areノルα"16秒α(Bosch＆SandeLac.）this study; Pinnatella ambigua (Bosch & Sande Lac.) ingtheirsourcesandvoucherspecimeninfbnnationing their sources and voucher specimen information. MF1eisch,ＡＢＯ94787,Taiwan,Pingung,MtPeitawu，M.Fleisch., AB094787, Taiwan, Pingung, Mt. Peitawu, VoucherspecimensarekeptinHIRqHYo，ｏｒＴＮｓ・Voucher specimens are kept in HIRO, HYO, or TNS. ｌ９５０ｍａｌｔ.,August２４，１９８８，Ｈ・AkiyamaTaiwan-l771950 m alt., August 24, 1988, H. Akiyama Taiwan-I 77 TreatmentoffhmiliesfbllowsVitt（1984）andBuckTreatment of families follows Vitt (1984) and Buck (HYo),thisstudy;Ｔｈａｍ"o6ﾉＷｍｓ"6ｓｅ７ｉａｊ""z(Mittex(BYO), this study; Thamnobryum subseriatum (Mitt. ex andVitt(1986）and Vitt (1986). SandeLac.）ＢＣＴan，ABO94791［ａｓ７１ｈａｍ"o6ﾉＷｍSande Lac.) B.C.Tan, AB094791 [as Thamnob/yum Thesequenceofinibrmationis：ｔｈｅｎａｍｅｏｆｔａｘｏｎ，The sequence of information is: the name of taxon, ｓα"dejlJapan，Honshu,HyogoPref，Kamigori-cho，sandei], Japan, Honshu, Hyogo Pref., Kamigori-cho, H.Ｈ・TsuBoTA,ＴＡＲＩＫＡｗＡ,Ｈ､ＡＫＩＹＡＭＡ,ＥＤＥＬｕＮＡ,Ｄ､GoNzALEz,MHIGucHlANDH､DEGucHI TSUBOTA, T. ARIKAWA, H. AKIYAMA, E. DE LUNA, D. GONZALEZ, M. HIGUCHI AND H. DEGUCHI 665６６５

Onaru-keikoku,Onam-keikoku,250malt.,Junel7,l99LH・Akiyama 250 malt., June 17, 1991, H. Akiyama ScleanScleanandourprogramsdownloadsite and our programs download site 14231l423］（HYo)，ｔｈｉｓｓｍｄｙ；LeucodontaceaeLezJcM)〃 (HYO), this study; Leucodontaceae Leucodon ＜http://homehiroshima-uacjp/chubo/＞ scillroidesscj"Ｍ伽(Hedw）SchwiigF.,ABO95988,Japan,Hon‐ (Hedw.) Schwagr., AB095988, Japan, Hon­ DatabaseＤα/αbQse shu,shu,NiigataPref,Itoigawa-shi,Kotaki,300ｍａｌｔ.,Ｊｕｌｙ Niigata Pref., Itoigawa-shi, Kotaki, 300m alt., July DDBJDDBJ＜http://www・ddbjnigacjp/〉 28,28,1994,HAkiyamal2594(HYo),thisstudyMyu- 1994, H. Akiyama 12594 (HYO), this study Myu­ SequenceSequencedatadownloadsite (For exam­ Japan,Japan，Shikoku，KochiPref，Monobe-mura，BefUkyo， Shikoku, Kochi Pref., Monobe-mura, Befukyo, ple,ple，ａｌｉｎｋｔｏａｎｅｎｔｒｙｗｉｔｈａｃｃｅｓｓｉｏｎｎｕｍber a link to an entry with accession number JulyJuly２７，１９９８，HAkiyamal4376（HYo),thisstudy； 27, 1998, H. Akiyama 14376 (HYO), this study; AB095988ABO95988is〈http://getentryddbjnigacjp/cgi-bin／ is ）entry. pi? AB095988» AB019466,ＡＢＯ19466,Japan，Shikoku，KochiPref，Monobe‐ Japan, Shikoku, Kochi Pref., Monobe­ AcceptedAccepted30・ＶＩＩＬ２００２ 30. VIII. 2002 mura,mura，Befilkyo，Ｊｕｌｙ２７，１９９８，Ｈ・Ａｋｉｙａｍａｌ４３５５ Befukyo, July 27, 1998, H. Akiyama 14355 鰯 (HYO),(HYo),thisstudy;Pterobryaceae伽c/(〕ﾉﾉollTqj"`/ｊｃｚｌ'〃 this study; Pterobryaceae Trachyloma indiclim .llJ'EBfW~坪田博美･有川智己･秋山弘之･ElTainDeLuna･DDlores -1'f }l1~c -f)([lJ%z -Efrain De Luna -lli1ores Mitt.,Mitt.,ＡＢＯ19464,Taiwan,Pingung,MtPeitawu，ｌ９５０ ABO 19464, Taiwan, Pingung, Mt. Peitawu, 1950 GonzalezGonzalez･樋口正信・出口博則：葉緑体ｒ６ｃＬ遺伝子の - tiIli n if {~ -l±l D fWftIJ : 1iH~f* rbcLirH~TO) malt.,ｍａｌｔ.,HAkiyamaTaiwan-50(HYo),thisstudy． H. Akiyama Taiwan-50 (HYO), this study. fi:T-:$7t·y大データセットに基づく腋蘇類とくにハイゴケ科とその H:~"'5indexｈｔｍｌ＞ tI'::k-:r'-:$7t 'Y H:~"'::J~~ -CfWtfiHT'?t.:. fWtfiO)*fi MOLPHY＜fip://fipjsmacjp/pub/ISMLIB/MOLPHY/〉MOLPHY *=,果，ハイゴケ目とイタチゴケ目が－つの大きなクレー /\1' ::f7 Fie l' :$7"f-:J7 EI tJ!-JO)::k,;! t.J. 7 v­ PAUP*ＰＡＵＰ＊＜http://paupcsit・fSu・edu/＞ ~~~pjzl..J.:ドを形成したDeLuna他(2000)およびBuck他(2000） De Luna ff!! (2000) sJ::LF Buck ff!! (2000) PPAUPRat＜http://viceroy・eehuconn・edu/paupratweb／ A UPRat pauprathtm＞ イタチゴケ目それぞれの単系統性は支持されなかっ1':$7"f-~7R..z:.n~nO).*.tt~5ZM~n~~'? TREE-TREE-PUZZLE＜http://www､tree-puzzlede/＞PUZZLE t.:.た．

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