HikobiaHikobial4:143-147.2004 14: 143-147.2004

PhylogeneticPhylO窪eneticandmorphOlO=icalmtesⅢIノルCD〃"剛〃昭肌oiKiguchi and morphological notes on Uleobryum naganoi Kiguchi eteraL(,Musci)’ ale (Pottiaceae, Musci) 1

HIROYUKIHIRoYuKISATQHⅡRoMITsuBoTA,ToMIoYAMAGucHIANDHIRoNoRIDEGucH1 SATO, HIROMI TSUBOTA, TOMIO YAMAGUCHI AND HIRONORI DEGUCHI

SATO,SATO,H、,TsuBoTA,H、,YAMAGucHI,T、&DEGucHI,H2004Phylogeneticandmor- H., TSUBOTA, H., YAMAGUCHI, T. & DEGUCHI, H. 2004. Phylogenetic and mor­ phologicalphologicalnotesonU/eo6Mイノ'z〃αgα"ojKiguchietα/、(Pottiaceae,Musci)Hikobia notes on Uleobryum naganoi Kiguchi et al. (Pottiaceae, Musci). Hikobia 14:l4:143-147. 143-147.

UleobryumU/eo6/Wm〃αgα"ojKiguchiejα/、,endemictoJapanwithalimitednumberofknown naganoi Kiguchi et aI., endemic to Japan with a limited number of known locations,locations,isnewlyreportedffomShikoku,westernJapanThroughcarefUlexamina- is newly reported from Shikoku, western Japan. Through careful examina­ tionoffTeshmaterial,rhizoidalmberfbnnationisconfinnedfbrthefirsttime・The , tion of fresh material, rhizoidal tuber formation is confirmed for the first time. The phylogeneticpositionofthiscleistocalpousmossisalsoassessedonthebasisofmaxi-phylogenetic position of this cleistocarpous is also assessed on the basis of maxi­ mummumlikelihoodanalysisof′bcLgenesequences、ThecuITentpositioninthePot- likelihood analysis of rbcL gene sequences. The current position in the Pot­ tiaceaetiaceaeissUpportedandacloserelationshiptoEpheme'wmslpj""/OS"川ssuggested is supported and a close relationship to Ephemerum spinulosum is suggested.

H1'わⅦkjStJro,H7ro〃ZM6ota,Tb〃oYZJ〃αg"chj&Hi7o"orjDeg"cAj,Dep”""e"'Hiroyuki Sato, Hiromi Tsubota, Tomio Yamaguchi & Hironori Deguchi, Department of0/Bjo/ogicα/Sbje"Ce,orα`"α/eSbAoo/q/肱e"Ce,肋Cs〃〃α〔ノノ、ノe'Mリノ,KtJgα〃‐ Biological Science, Graduate School of Science, Hiroshima University, Kagami­ yamαノー3-/,Higashj-/iかCsノi〃α-s/ij,価'0s/i伽α734852.ノヒリpα〃yama 1-3-/, Higashi-hiroshima-shi, Hiroshima 739-8526, Japan.

arearemissingMolecularphylogeneticsmdieshave mlssmg. Molecular phylogenetic studies have Introduction beenmadetoclarifythephylogeneticpositionofbeen made to clarify the phylogenetic position of Thegenusueo6'W腕isasmallgenuscom-The genus Uleobryum is a small genus com­ somesomemosseslackingperistomes(Goffinet&Cox lacking peristomes (Goffinet & Cox prisingonlythreespecies(Kiguchiejα/,1996;prising only three species (Kiguchi et al. 1996; 2000;Goffinete/α/、1998,2002;Heddersonem/、2000; Goffinet et al. 1998, 2002; Hedderson et al. Zanderl993,1994;Zander&Dixonl999LwithZander 1993, 1994; Zander & Dixon 1999), with 2004;Stechl999).Theseresearchesdemonstrate2004; Stech 1999). These researches demonstrate adistributionrangeintheWesthdies,Mexicqa distribution range in the West Indies, , thatmoleculardataprovideuswithinfb、nationthat molecular data provide us with infonnation Peru,Brazil,Australia,andJapanThegenusisPeru, Brazil, Australia, and Japan. The genus is usefUltobetterunderstandsuchenigmaticcleisto-useful to better understand such enigmatic cleisto­ characterizedbyitssmallsize,acostawithtwocharacterized by its small size, a costa with two carpousmosses・hthispaper,rhizoidaltubercarpous mosses. In this paper, rhizoidal tuber stereidbandsandelongateadaxialepidennalstereid bands and elongate adaxial epidennal fbnnationfbrU/eo6MJm〃αgα"ojisnewlyre-fonnation for Uleobryum naganoi is newly re­ cells,spheroidalcleistocalpouscapsuleswithacells, spheroidal cleistocarpous capsules with a portedandthephylogeneticpositionofthisspe-ported and the phylogenetic position of this spe­ hyalineseta,transparentcapsulewallsofwhichhyaline seta, transparent capsule walls of which ciesisalsodiscussedonthebasisofribulosecies is also discussed on the basis of ribulose cellsaremamillose,withslightlythickenedfreecells are mamillose, with slightly thickened free 1,5-bisphosphateL5-bisphosphatecarboxylase/oxygenaselarge carboxylase/oxygenase large walls,andconic-campanulatecalyptraethatarewalls, and conic-campanulate calyptrae that are subunit('6cL)genesequenceswithmaximumsubunit (rbcL) gene sequences with maximum roughenedapically(Zanderl993).Ithasbeenroughened apically (Zander 1993). It has been likelihoodanalysis,togetherwithareportofnewlikelihood analysis, together with a report of new placedinthePottiaceaesinceitsestablishmentasplaced in the Pottiaceae since its establishment as habitatinJapan.habitat in Japan. anindependentgenusbyBrotherus(1906).meo-an independent genus by Brotherus (1906). Uleo­ 6M/m〃αgzJ"ojKiguchierα/、hasbeendescribedbryum naganoi Kiguchi et al. has been described MaterialsandmethodsMaterials and methods asendemictoJapanbyKiguchiem/、(1996),andas endemic to Japan by Kiguchi et al. (1996), and ischaracterizedby(1)linear-1anceolateleaves,is characterized by (1) linear-lanceolate leaves, TotalDNAwasextractedffom廿eshmaterialTotal DNA was extracted from fresh material (2)stomataonthemiddleoftheupperhalfofthe(2) stomata on the middle of the upper half of the usingthemodificationsofaphenol-chlorofbImusing the modifications of a phenol-chlorofonn capsule,and(3)unlobedcalyptrae,Thesystem-capsule, and (3) unlobed calyptrae. The system­ method(TsubotaeM/、1999).PCRamplifica-method (Tsubota et al. 1999). PCR amplifica­ aticpositionofcleistocalpousmossesisequivo-atic position of cleistocarpous mosses is equivo­ tionsanddirectsequenceanalyseswerecarriedtions and direct sequence analyses were carried calwhenconclusionsarelimitedtomolphologi-cal when conclusions are limited to morphologi­ outusingstandardtechniquesDetailsandfUr-out using standard techniques. Details and fur­ calcharacters,particularlywhenmajorcharacterscal characters, particularly when major characters therinfbnnationfbrDNAextractingprotocols,ther infonnation for DNA extracting protocols, amplificationprimers,PCRprotocolsandthoseamplification primers, PCR protocols and those lContributionffomtheLaboratoryofPlantTaxonomyI Contribution from the Laboratory of ofsequencingreactionsaredescribedinTsubotaof sequencing reactions are described in Tsubota andEcology,HiroshimaUniversity.N・Ser、No.538.and Ecology, Hiroshima University. N. Ser. No. 538. em/、(1999,2000,2002).et al. (1999, 2000, 2002). 144144 HikobiaVoLl4,No.2,2004Hikobia Vol. 14, No.2, 2004

FiftysixmossspecieswereincludedintheFifty six moss species were included in the haplolepideousmossesincludingsomediplolepi-haplolepideous mosses including some diplolepi­ analysesandthesequencesweremanuallyanalyses and the sequences were manually deousmosses・Fortheanalyses,atotalofL210deous mosses. For the analyses, a total of 1,210 alignedOutgrouptaxawereselectedbasedonaligned. Outgroup taxa were selected based on sites(84.73%in1,428bpoftheentiresequence)sites (84.73% in 1,428 bp of the entire sequence) thephylogenetichypothesisproposedbythe phylogenetic hypothesis proposed by of5676cLsequenceswereusedof 56 rbcL sequences were used. Magombo(2003)andTsubotaejaL(2003).Magombo (2003) and Tsubota et at. (2003). lnfbnnationonthespecimensfTomwhichInfonnation on the specimens from which AphylogenetictreewasconstructedusingA phylogenetic tree was constructed using DNAwasextractedwiththeaccessionnumbersDNA was extracted with the accession numbers maximumlikelihoodcriteria(ML;Felsensteinmaximum likelihood criteria (ML; Felsenstein isgivenbelow:is given below: l981)AnNJtreewasobtainedusingNjdistin1981). An NJ tree was obtained using Njdist in Bノビ)′o抑ノカノ"m〃o'WgJc""zsubspノ叩o"Jc""’Bryoxiphium Ilorvegicum subsp. japonicum ’ MOLPHY23b3(Adachi&Hasegawal996)MOLPHY 2.3b3 (Adachi & Hasegawa 1996) (Berggr.)A・L6ve&DL6ve,AB194720,Japan,(Berggr.) A.Love & D.Love, ABI94720, Japan, withHKY85model(Hasegawaetα/、1985),andwith HKY85 model (Hasegawa et at. 1985), and Honshu,Tottori-ken,Hino-gun,KoufU-chqMt・Honshu, Tottori-ken, Hino-gun, Koufu-cho, Mt. thebestMLtreewassearchedusi、gNucMLinthe best ML tree was searched using NucML in Karasugasen,1,200malt.,June18,2000,LegS・Karasugasen, 1,200 m alt., June 18, 2000, Leg. S. MOLPHYwiththeNJtreeasastarttopologyfbrMOLPHY with the NJ tree as a start topology for NogawainhbHDeguchi36501(HIRO).Nogawa in hb. H. Deguchi 36501 (HIRO). alocalrearrangementalgorithma local rearrangement algorithm. UleobryumU/eo6ノW"z〃αgzJ"ojKiguchie/αL,AB194717, Ilaganoi Kiguchi et al., AB 194717, New76cLsequencesweredetenninedfbrfburNew rbcL sequences were detennined for four Japan,Shikoku,Kagawa-ken,Ayauta-guLAyakami-Japan, Shikoku, Kagawa-ken, Ayauta-gun, Ayakami­ ● species:*Br〕ノoxjphj""z〃orvegzczJ"1subsp,ノ叩o"ノーspecies: *Bryoxiphium norvegicum subsp. japoni­ cho,NagaraLake,Ca」OOmalt.[associatedwithcho, Nagara Lake, ca. 100 malt. [associated with c"腕(ABl94720),*meo6'Wm"αgzJ"oj(AB1947cum (AB 194720), * Uleobryum naganoi (AB 1947 印/jemelw"Tspj""/Cs""7Bruch&Schimp],Dec・Ephemerum spinulosum Bruch & Schimp.], Dec. 17),肌maco""ove耐α(ABl94718)and*Elphe-17), Weissia controversa (AB 194718) and *Ephe­ 13,2003,'3,2003,HSato377(HIRO). H. Sato 377 (HIRO). "'e'wmgpj"[イノCs"〃(AB194719).Threespeciesmerum spinulosum (ABI94719). Three species Weissiaリリノi2jlssjaco"〃ove'FaHedw.,AB194718,Japan, controversa Hedw., AB194718, Japan, withanasteriskwereshownasanentiresequencewith an asterisk were shown as an entire sequence Honshu,Hiroshima-ken,Higashi-hiroshima-shLHonshu, Hiroshima-ken, Higashi-hiroshima-shi, 1,428l,428bplongAdatamatrixwaspreparedfbr bp long. A data matrix was prepared for ca200ma1t.,July3,2003,HSatol61(HIRO)ca. 200 m alt., July 3,2003, H. Sato 161 (HIRO). analysesanalysesofphylogeneticrelationshipsoftlle of phylogenetic relationships of the Eドフハeme'w"zSpj""/Cs""ZBruch&Schimp,Ephemerum spinulosum Bruch & Schimp.,

L A iill

500~mⅡ、 、

L魁

、の●◆ ~m40Ⅱ、

Fig.Fig・LRhizoidaltubersofU/eo6ひ'z"〃〃αgα'MKiguchieZα/、 1. Rhizoidal tubers of UteoblJ'ulI1 nagalloi Kiguchi et at. A.APlantwithacapsuleandrhizoids・B-FRhizoidalmbers(BcorrespondingtobinA;Ctoc). Plant with a capsule and rhizoids. B-F. Rhizoidal tubers (B corresponding to b in A; C to c). PhotographedPhotographedanddrawnfiPomHSato377. and drawn from H. Sato 377. H.H・SATO,H・TsuBoTA,TYAMAGucHIANDH、DEGucHI SATO, H. TSUBOTA, T. YAMAGUCHI AND H. DEGUCHI 145145

ABABl94719,Japan,Shikoku,Kagawa-keL 194719 , Japan, Shikoku, Kagawa-ken, bersbersofOTUswereshowninparenthesesinFig. of OTUs were shown in parentheses In Fig. Ayauta-gun,Ayauta-gun,Ayakami-cho,NagaraLake,cal00 Ayakami-cho, Nagara Lake, ca. 100 2.2. malt.,malt.,Decl3,2003,HSato378(HIRO). Dec. 13,2003, H. Sato 378 (HIRO). TheTheotherflftytwosequenceswereobtained other fifty two sequences were obtained fromfTomtheDNAdatabases,andtheaccessionnum- the DNA databases, and the accession num-

一一議 Hypnum plumaeforme (AB029384) n------Sematophyllum subhumile (AB039675) Entodon challengeri (AB050993) .---- Plagiothecium nemorale (AB029387) -11-______Adelothecium bogotense (ABI03354) 100 Funaria apophysata (AF0055 14) Physcomitrella patens (X74156) 色 Enc.lypta streptocarpa (AF478239) Encalypta rhaptocarpa (A1275 167) 100 Ephemerum spinulosum (ABI94719)

Uleobryum naganoi (ABI94717) の国の。⑤二」。』 Pseudosymblcpharis schimpcriana (AF226805) Weissia controvcrsa (AB 194718) Barbul. convoluta (A1'478225) 100 Pottia Intemledia (ABI25592) ● Tortula obtusissima (A1'226823) muticum (A1'231 078) 55 Streptopogon calymperes (A1'47823I ) Tortula ruralis (TRU275 169) Aulacopilum hodgkinsoniae (AF005545) Venturiell. sinensis (AF005546) "------Uleastrum palmicola (AF005547) Kiaeria blyttii (AF231306) Rhabdoweisia crenulata (AF005544) -l.....___ Dieranoweisia cirrata (AF23 1297 ) Ditrichum pallidum (AF23 1302) 100 Pleuridium acuminatum (AF231312) Hypodontium pomiforme (AF226803)380 pP71 65 Paraleucobryum enerve (AF226827) 575 100 OrthodicranumDrthod i"ulvum (AF231311) Dicranum condensatum (AI'231298) uiニゼDicnemon seriatum (AI'478229) --""""1L___ Eucamptodon muelleri (A1'231319)9) n-----~ Fissidens mooreaea (A1'2268I5810 0) Ⅲ 杵L-j里『'---=L Fissidens osmundoidesロ (A1'231 0(9) Fissidens dubius (AF231303) Amphidium lapponicum (AF005543) IF"'--- Timmiella crassinervis (AF478236) Li二三iili撰驚Scouleria aquatica (AF226822) Drummondia obtusifolia (AF232697) Bryoxiphium norvegicum sUbsp.japonicum (ABI94720)94720) Brothera leana (AB 122033) 81 Dicranodontium denudatum (AB 122034) Leucobryumjuniperoideum (AB 124786) Leucobryum sanctum (AB 124787) 1.-___ Campylopus argyrocaulon (AF231315) '"=--- Archidium stellatum (AF23 1(66) Blindia acuta (A1'226817) Ptychomitrium gardneri (AF005549) Grimmia laevigata (AF478230) 72 Mitthyridium fasciculatum (A1'226780) 94 Syrrhopodon tortilis (AF226782) b.____ Calymperes palisotii (A1'226791) ....------Octoblepharum albidum (AF226795) Diphyscium longifolium (AF4782 I 7) Diphyscium fulvifolium (AF478222) B■L--....I 0.Olsubstitutions/site0.0 t substitutions/site

Fig2、PhylogeneticpositionofU/eo6Ml"1〃αgα"oj.ThemaximumlikelihoodtreeofthealignedFig. 2. Phylogenetic position of Uleoblyum naganoi. The maximum likelihood tree of the aligned r6cLgenesequences,1,210bpof561bcLgenesequences(HKY85model[Hasegawaetα/、1985];rbcL gene sequences, 1,210 bp of 56 rbcL gene sequences (HKY85 model [Hasegawa et at. 1985]; 2α/β=4865;InL=-9120.08±392.04byNucML);andthecladeswithinthePottiaceaeclade2 a I [3 = 4.865; In L = - 9120.08 ± 392.04 by NucML); and the clades within the Pottiaceae clade. ThenumbersinparenthesesshowaccessionnumbersofeachOTU、ThehorizontallengthofeachThe numbers in parentheses show accession numbers of each OTU. The horizontal length of each branchcorrespondstotheestimatednumberofnucleotidesubstitutions・Therootisarbitrarilybranch corresponds to the estimated number of nucleotide substitutions. The root is arbitrarily placedonthebranchleadingtoDjp伽cjzl",ノo"g(/M""Z(AF478217)and、ルノWM"〃placed on the branch leading to Diphyscium /ongi[olium (AF4782 17) and D. fu/v(/olium (AF478222)(AF478222). 146146 HikobiaHikobiaVol」4,No.2,2004 Vol. 14, No.2, 2004

those of the genera of the family Funariaceae and Results thoseofthegeneraofthefamilyFunariaceaeand thethesystematicpositionofthegenusintheorder systematic position of the genus in the order RhizoidalRhizoidaltubersconsistofasingle,bifUrcated tubers consist of a single, bifurcated FunarialesFunarialesprovedtobedoubtfUl,judgingfrom proved to be doubtful, judging from ororbranchedcell-row,tingedreddish-brown, branched cell-row, tinged reddish-brown, thethepresentresultandtheirdissimilarmoIpho- present result and their dissimilar morpho­ whichwhichissimilarincolortotherhizoidsThetu- is similar in color to the rhizoids. The tu­ logicallogicalcharacters、HenceEpAeme'wmshouldbe characters. Hence Ephemerum should be berberceUsarebarrel-shaped,moreorlessisodia- cells are barrel-shaped, more or less isodia­ transferredtransfierredfi-omtheEphemeraceaetothePottia- from the Ephemeraceae to the Pottia­ metric,metric,ca20-551LLmthick;eachoftbemisusu- ca. 20-55 Ilm thick; each of them is usu­ ceaeceaeaswasdonebyGofHnet&Buck(2004) as was done by Goffinet & Buck (2004). allyallyaccompaniedbyasmalllenticularinitialcell accompanied by a small lenticular initial cell TheThehomologyvalue98、39%(=L405bp)of homology value 98.39% (= 1,405 bp) of ca.cal4-32lImindiameter、InitialceUsbeganto 14-32 Ilm in diameter. Initial cells began to UUl〃αgα"ojandE・spj""/Cs""zsuggestedmore naganoi and E. spinulosum suggested more developdevelopintoprotonemainafiewweeksafterliq- into protonema in a few weeks after liq­ detaileddetailedinvestigationisrequiredwithmorespe- investigation is required with more spe­ uiduidcultivationinapetridishatroomtemperature cultivation in a petri dish at room temperature ciesciessampledfiDmthePottiaceaetoobtainmore sampled from the Pottiaceae to obtain more (20-25"C).(20-25℃) preciseprecisephylogeneticinfbrmationonthegenera phylogenetic information on the genera TheTheMLtreewasobtainedbyNucMLOnL= ML tree was obtained by NucML (In L = UleobryumU/eo61WmandElD/ie腕e'脚加withinthePottia- and Ephemerum within the Pottia­ -9120.08-9120.08±392.04;Fig.2).Fourlargeclades ± 392.04; Fig. 2). Four large clades ceae.ceae・meo6Mイmwasrelatedtoothercleisto- Uleobryum was related to other cleisto­ て correspondingcoITespondingtotheBryidae,FunariaceaQEn- to the Bryidae, Funariaceae, En­ carpouscarpousgeneraBryoce"tAoSpolaand〃αc/j〕ノcar- genera Bryoceuthospora and Trachycar­ calyptaceae,calyptaceae,andDicranidaewereresolvedwitha and were resolved with a pj`j"脚aspreviouslysuggestedbyZander(1993,pidium as previously suggested by Zander (1993, highhighBPsupport(100%,100%,100%,and95%, BP support (100%, 100%, 100%, and 95%, 1994).1994). respectively).respectively).WithintheDicranidaeclade,the Within the Dicranidae clade, the Finally,Finally,populationsofU/eo6M"〃〃αgzJ"ojre- populations of Uleobryum naganoi re­ ヴ Pottiaceae-EP〃eme'w"ZcladewasresolvedwithPottiaceae-Ephemerum clade was resolved with mainmaininrestrictedlocationswhichincludeKana- in restricted locations which include Kana­ strongstrongsupport(100%).WithinthePottiaceae- support (100%). Within the Pottiaceae­ gawa,gawa,thetypelocality,Shizuoka,andSaitama the type locality, Shizuoka, and Saitama EphmerumE;phmelwmclade,twosubcladeswereresolved: clade, two subclades were resolved: PrefecturesPrefecmresofEastJapa、(Kiguchietα/1996; of East Japan (Kiguchi et at. 1996; thetheU/eo6'Wm-Elphemer""0-Ae"`/DWZ6ノビアノiMs Uleobryum - Ephemerum - Pseudosymblepharis IwatsukiIwatsuki&Suzukil998;Kiguchi2003).Anew & Suzuki 1998; Kiguchi 2003). A new --肌ma-Bα'仇ノノaclade(100%)andthePo"jα-Weissia-Barbula clade (100%) and the Pottia­ locationlocationisfirstreportedhereffomShikoku,west- is first reported here from Shikoku, west­ Tortula-Acaulon-Streptopogon7M"/α-化α"/o'1-s"叩rOpogo〃clade(100%). clade (100%). ernernJapanTheplantsweregrowingonsandysoil Japan. The were growing on sandy soil WithinWithinthefionnerclade,〔ノノeo6Mノノ〃hasaclose the former clade, Uleobryum has a close mixedmixedwithgravelontheshoreofawaterreserve with gravel on the shore of a water reserve relationshiprelationshipwith助hemeノwm(100%)Homolo‐ with Ephemerum (100%). Homolo­ forfbrirrigation,beingassociatedwith印Aemelwm irrigation, being associated with Ephemerum gygyvaluesofthetotalr6cLgenesequencesofU values of the total rbcL gene sequences of U spinulosum.叩j""/Cs""0. naganoi"αgα"ojandE・叩/""/Cs"mprovedtobeidentical and E. spinulosum proved to be identical withwiththevalue98.39%(=L405bp). the value 98.39% (= 1,405 bp). Acknoledgements

WeWethankDrT、MatsuiandDr・TArikawafbr thank Dr. T. Matsui and Dr. T. Arikawa for Discussion constructiveconstructivesuggestionsonanearlierversionof suggestions on an earlier version of UloebryumU/oeZW"misacleistocarpousmoss,andthe is a cleistocarpous moss, and the thisthispaper,andMr.T,Suzukifbrcommentsand paper, and Mr. T. Suzuki for comments and gametophytesgametophytesshowfeaturescharacteristicof show features characteristic of providingprovidingecologicalinfbnnationFurtherthanks ecological information. Further thanks manymanyspeciesofthePottiaceae,suchas(1)obo‐ species of the Pottiaceae, such as (1) obo­ areareextendedtoDr.R・Stotlerfbrcheckingthe extended to Dr. R. Stotler for checking the vatevatetolinearlanceolateleaveswithanacutetip, to linear lanceolate leaves with an acute tip, EnglishEnglishtext. text. andand(2)laminacellsthataredenselypapillateon (2) lamina cells that are densely papillate on bothbothsurfacesintheupperandmiddlepartsofthe surfaces in the upper and middle parts of the LiteratureLiteraturecited cited leaves.leavesThepresentresultssuggestacloserela- The present results suggest a close rela­ tionshiptionshipofU/eo6ノW"zwithElpAe"Te'w"Land of Uleoblyum with Ephemerum, and Adachi,Adachi,』.&Hasegawa,Ml996ComputerScience J. & Hasegawa, M. 1996. Computer Science theirtheirinclusioninthePottiaceaewasstronglysup- inclusion in the Pottiaceae was strongly sup­ Monographs,Monographs,Z8MOLPHYversio、2.3.Programs 28. MOLPHY version 2.3. Programs ported,ported,aspreviouslyreportedonthebasisofthe as previously reported on the basis of the forfbrmolecularphylogeneticsbasedonmaximum molecular phylogenetics based on maximum datadataonlps4genesequences(Gofflnet&Cox on rps4 gene sequences (Goffinet & Cox likelihood.likelihoodl50pplnstimteofStatisticalMath- 150 pp. Institute of Statistical Math­ 2000;2000;Gofflnete/α/、2001;Heddersone/α/、 Goffinet et at. 2001; Hedderson et al. ematics,ematics,Tokyo・ Tokyo. 2004).2004).HeddersoneM/、(2004)ftlrtherinfierred Hedderson et at. (2004) further inferred Brotherus,Brotherus,V・F1906.Musciamazonicietsubandini V. F. 1906. Musci amazonici et subandini thatthatsomespeciesof〃IgjssjainthePottiaceae some species of Weissia in the Pottiaceae U1eani.Uleani・Hedwigia45:260-288. Hedwigia45: 260-288. alsoalsoapproachEipノieme'wm,i,e、,specieswithre- approach Ephemerum, i.e., species with re­ Felsenstein,Felsenstein,J198LEvolutiontreesfiPomDNAse- J. 1981. Evolution trees from DNA se­ ducedducedplantsize,immersedsporophytes,andper- plant size, immersed sporophytes, and per­ quences:quences:amaximumlikelihoodapproachJ・Mol a maximum likelihood approach. J. Mol. sistentsistentprotonemata・ protonemata. Evol.Evoll7:368-376 17: 368-376. TheThecloserelationshipofEpAeme'wmwith close relationship of Ephemerum with Goffinet,GofHnet,B&Buck,W、R・Z004Systematicsofthe B. & Buck, W. R. 2004. Systematics of the H.H・SATO,H・TsuBoTA,TYAMAGucHIANDH・DEGucHl SATO, H. TSUBOTA, T. YAMAGUCHI AND H. DEGUCHI 147147

BryophytaBryophyta(mosses):frommoleculestoarevised (mosses): from molecules to a revised weisiaceae,weisiaceaeBryopsida)NovaHedwigia68:291- ). Nova Hedwigia 68: 291- classification.classiHcationlnGofHnet,B,Hollowel1,W.& In Goffinet, B., Hollowell, W. & 300.300. Magill,MagilLR(eds),MolecularSystematicsof R. (eds.), Molecular Systematics of Stone,Stone,LG」984.U/eo6ノWmc"'7Msp・nov.(Pottia- I. G. 1984. Uleobryum curtisii sp. nov. (Pottia­ Bryophytes,Bryophytes,pp205-239・MissouriBot、Garden pp. 205-239. Missouri Bot. Garden ceae)ceae)fiFomQueensland,Australia、JBryoL13: from Queensland, Australia. J. Bryol. 13: Press,Press,StLouis. St. Louis. 19-24.l9-24 ----&COX,C・l2000Phylogeneticrelationships& Cox, C. J. 2000. Phylogenetic relationships Tsubota,Tsubota,H、,Akiyama,H、,Yamaguchi,T&DeguchL H., Akiyama, H., Yamaguchi, T. & Deguchi, amongamongbasal-mostarthrodontousmosseswith basal-most arthrodontous mosses with H.H・Z00LMolecularphylogenyoftheSematophyl- 2001. 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1 nese) nese). 佐藤裕幸・坪田博美・山口富美夫・出口博則:ツチ~~m*·~rnM.·iliD ••*·mDM.:Y~ -,Stone,1.G.&Iwatsuki,Zl996〔ノル06ノW、I. & ---, Stone, G. Iwatsuki, Z. 1996. Uleobryum ノウエノハリゴケの系統および形態についてJ t'JIJ 1\ I) :i70)iMiJC;f3J::a:m~I::-:J~\-C 〃αgα"oJspnov.(Pottiaceae)fbundinJapannaganoi sp. nov. (Pottiaceae) found in Japan. Hikobial2:157-l60Hikobia 12: 157-160. 西日本新産となるツチノウエノハリゴケについて[.q B *m~ c t;;'9 Y~ J t'J I J 1\ I) :1''rl:: -:J ~\-C Magombo,ZLK、2003.ThephylogenyofbasalMagombo, Z. L. K. 2003. The phylogeny of basal 仮根上結節を報告した.さらに葉緑体,bcL遺伝子のf&:fJLI:~~£i1j Gt:::.o ~ G I::~*'1