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Hypnaceaeandpossiblyrelatedfn 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,ノブIノ・Csノi〃α739-852.ノヒW/Lshi. Hiroshima 739-8526. Japan. 7bmojiFz`9MノWmwq,Depαノブ'"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-/ゴイ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
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