Copyright (c) American Society for Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 DOI 10.1600/036364411X553081 © Copyright2011 bythe American SocietyofPlantTaxonomists Systematic Botany Mitt., ter clades:thecore sematophyllaceoustaxa (e.g. 2001a , b ) show thattheSematophyllaceaes.l.includestwosis- might notbelongthere. 1998 ; Hedenäs and Buck 1999 ) alsosuggested that morphological cladisticanalyses ( Hedenäs 1996 ; Tan andJia suggested theexclusionof genusfrom thefamily, and otherwise diagnosticfortheSematophyllaceae. Seki (1969) cells, longrostrate operculum, andinflatedalarcellsthatare However, and But1997 ; Buck andGoffinet 2000 ; followed thistreatment ( Ramsay etal.2002 Vitt 1984 ). ; Sematophyllaceae, andmanytaxonomistshavesubsequently Buck andVitt 1986 ; Tan a briefdiagnosisandincludedonly by Mitten (1869) inthetribeSematophylleae.Mittenprovided Spruce (1867) , butthenamewasonlylatervalidlypublished all forthe African or Asian species. ment publishedforthegenusandnosystematictreatment at within thegroup. To datethere hasbeennoworldwidetreat- and lesscommonspeciesof inthetropics, there are anumberofpoorlyknown (Brid.) Mitt.and differentiated asinmostSematophyllaceaes.l. with leaveshavinganalarregion withcellsnotnearlyaswell useful inrecognizing the genus are thecomplanatebranches only twiceintheSematophyllaceae.Othercharactersthatare is arare characterinthe Hypnalesandhasbeendescribed cells (hence,Tax- taso=arrangedandthelionnipple).This of multiplepapillaedisposedinseriesontheluminaleaf ily Pylaisiadelphaceaeasasegregate from Sematophyllaceae. until 2004,when Goffinet andBuck(2004) describedanewfam- Sematophyllaceae (),from itsoriginaldescription and Longton1996 ; Ramsay etal.2002 ). Southeast Asia, especiallytheMalesianregion ( Damanhuri between 30°Nand20°S,withmostspeciesoccurringin generainthetropics. Thegenusisbestrepresented Farge-England 1996 ) isprobably oneofthemostwidespread Recent molecular studies( Buck etal.2000 ; Tsubota etal. Brotherus (1925) placedthegenusin thefamily History— Taxonomic Even thoughthemostcommonspecies(e.g. The maincharacterthatdefines The genushasbeentraditionallyassociated withthe Taxithelium and theremaining from Southeast Asia andOceania.Keys,illustrations,descriptionsare provided. pseudoparaphyllia inthelatter. The subgenus ghished from subgenus Keywords— Abstract— 2

Current adress: UniversidadedeBrasília,Deptartmento deBotânica,CampusUniversitárioDarcy Ribeiro, Taxithelium (2011), 36(1):pp.7–21 A ecrusrpin f h Ms Gns Genus Moss the of Re-circumscription A , agenusofpleurocarpous mosses(sensu La The mossgenus T. nepalense roht lsiiain, Hpae eaohlaee, Pliidlhca otes Ai mrc fia, t Africa, America, Asia, Southeast Pylaisiadelphaceae, Sematophyllaceae, Hypnales, classification, Bryophyta, Mitt.,and 1 CAPES Fellow, BrazilianGovernment.MissouriBotanicalGarden, P.O. Box299,St.Louis, lacks thecollenchymatousexothecial Taxithelium Taxithelium (Schwägr.) Broth.) are abundant Trichosteleum with aTaxonomic RevisionofSubgenus Taxithelium Taxithelium bythelanceolateleaves andfilamentouspseudoparaphylliaintheformerovateleavesplusfoliose Taxithelium wasfirstrecognized by T. planum isreclassified intotwosubgenera:

3 Author forcorrespondence ( [email protected] ) Mitt.),andaclade andmanynames Vernieri inthegenus. isthepresence omnctn Eio: ea Struwe Lena Editor: Communicating Sematophyllum Missouri 63166-0299,U.S. A. al E A S Câmara S. A. E. Paulo isrevised here andcompriseselevenspecies;onefrom Africa, twofrom the Americas, Taxithelium T. planum Brasília, DF, Brazil

7 names andisin great needofrevision. simultaneously grown tomore than 230acceptedspecies unplaced overtheyears.Not surprisingly, free” treatments of et al.2002 ) haveexcludedspecieswithoutpapillaefrom local without papillaeandsomeauthors (e.g. Tan etal.1996 ; Ramsay taxonomists havelargely rejected theconceptofa W. R.Buck&B.C.Tan, phologically from abroad rangeofgenerasuchas Taxithelium opercula. Thisfurtherbroadened thegenericboundariesof of collenchymatousexothecialcellsandlacklongrostrate cell organization, andsporophyte features suchasthelack within (an autonymicsubgenus). are synonymsbecausetheyfailedtoidentifysubg. some ofthesubgenericnamescreated byRenauldandCardot Pilotrichaceae withinHookeriales( Buck etal.2005 ). However, new familyPylaisiadelphaceaeforthe“ Based ontheseresults, Goffinet andBuck(2004) describedthe et al.(2001a) called thelattergroup “the Schimp., genus Phyllodon section cells), and lose alarcells), into three sections: one papillapercell).Thesubgenus lae percell,notseriallydisposed),and posed seriallypercell), into three subgenera: morphologies. Renault andCardot (1901) divided Taxithelium the SematophyllaceaeplusPylaisiadelphaceae. still remain unclear, Irefer here toSematophyllaceaes.l.for tionships withinSematophyllaceaes.andPylaisiadelphaceae Taxithelium pomorphy, Goffinet andBuck(2004) are followedhere and Although thisgroup lacks anobviousmorphologicalsyna- that includes Later, Brotherus (1925) included plantswithoutpapillae Since itwasfirstdescribed,thegenericboundariesof Taxithelium Taxithelium Taxithelium Pseudohypnella Taxithelium Anastigma 1 ( Buck 1987 ) and Brotherus (1909) created thesub- , Isopterygium 2 , 3 andmadeitmore difficult todifferentiate itmor- havebeenstretched tofitagreat varietyofplant isincludedinthePylaisiadelphaceae. As therela- Limnobiella

and Taxithelium Aptera Taxithelium species have remained mostlyignored and Vernieri forasinglespeciesthatisnowplacedin basedonsimilaritiesinleafshape,alar (Pylaisiadelphaceae) Vernieri Vera forasinglespeciesnowplacedin (aquaticplants). Cardot (1905) created Mitt.,and (nonaquaticplantswithquadratealar Polystigma Isopterygium, . Thesubgenus Oligostigma , (nonaquaticplantswithvesicu- . Consequently these“papillae- Pylasiadelpha

(withseveralpapillaedis- Brotherella (withoneorfewpapil- and Polystigma Vernieri Monostigma Cardot, Brotherella Brotherella Trichosteleum. axonomy. Loeske. Tsubota canbedistin- Taxithelium wasdivided Platygyrium Taxithelium Taxithelium Taxithelium (withonly lineage.” lineage.” Radulina Most has

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 from type material. under aNikonLabophot-2 lightmicroscope. All illustrations were made from leaves takenfrom themiddle ofthestemorbranchandviewed Gradstein etal.(2001) and Magill (1990) . All measurements were made and facilitateitslocation. logues are presented whenneededtosecurely identifythetypematerial decisions. logical datapresented here are sufficient asthebasisforthesetaxonomic by usingmultivariatemethods(availableuponrequest), themorpho- by molecularstudies(unpublishedresults, availableuponrequest) and Even thoughthetaxonomicresults presented here are alsosupported were madefrom mountedmaterial. solution( Hoyer’s Anderson 1954 ). All observationsandmeasurements W). Specimenswere rehydrated inboilingwaterandthenmounted MO, NICH,NY, NSW, PC,PHS,S,SING,SINU,SP, TSN,UB,UPS,US, herbaria (B,BM,BR,CANB,DUKE,E,FH,G,H,JE,L,M,MG,MICH, [Volume 36 SYSTEMATIC BOTANY only three species( Buck (1985) reviewed 8 be publishedlater. the subgenus circumscription for here. tion andinfragenericclassificationof polyphyletic andprovides supportforthenewcircumscrip- (unpublished) showedthatRenauldandCardot’s groups are not supported.ThemolecularstudyofCâmaraandShaw the subgenericclassificationof Renault andCardot (1901) is either ignored orwere notknownatthattime.Inaddition, such aspseudoparaphylliaandperichaetialleaves,were acters considered important ingenericplacementtoday, fit atallintoRenauldandCardot’s concepts,andmanychar- Many speciesof Acanthorrhynchium genera suchas of thespeciestheyincludedin which wasproposed by Renault andCardot (1901) . Most tion ofthecurrently accepted circumscription of characters ofspeciesincludedin veloped papillae,onlyvisibleundertheSEM. showed thatsuchsmoothleafcellsmighthavesmall,unde- of micromorphology ofpapillae( Câmara andKellogg2010 ) species withsmoothleafcellsinthegenus.Detailedstudy clusion supportstheviewsof Brotherus (1925) , whoplaced species of lose leafcellsare synapomorphicfor recognized byaparticularpapillamorphology, 2)pluripapil- posed oftwostrongly supportedclades,eachofwhichcanbe data) showthat,1) and Africa. Southeast Asia (1998) . were includedby Sharp etal.(1994) andthree speciesby Buck vided alocalrevision ofsix Australian species,twospecies 1996 ) andtheeffort washalted. Ramsay etal.(2002) preliminary results were published( pro- Damanhuri andLongton the genuswasunderworldwiderevision inthe1990s,only Cardot) W. R.Buck,and Even detailedstudiesonsmaller partsofthegenusare few. Objectives— Circumscription— Molecular phylogeneticdata(CâmaraandShaw, unpubl. The morphologicaltermsusedwere definedandillustratedin Typifications are provided forallnamesandinformationfrom proto- Species were recognized onthebasisofmorphologicaldiferences. — Taxithelium Taxithelium Loans totaling6,200specimenswere obtainedfrom 29 Vernieri The goalsofthispaperare toprovide: 1)anew Phyllodon Taxithelium hasnotbeentreated inrecent florasfor M.Fleisch.,and Taxithelium T. planum Taxithelium Detailed studiesofthemorphological . A revision ofsubgenus havelostthepapillae.Thelattercon- Taxithelium T. juruense Methods Bruch &Schimp., describedafter1901wouldnot , , and2)ataxonomicrevision of ismonophyleticandcom- T. pluripunctatum Taxithelium Taxithelium forBrazil,andrecognized (Broth.) Broth.). Although Taxithelium Taxiphyllum Taxithelium belong toother ledtotherejec- Taxithelium Trichosteleum , and3)some Rnud & (Renauld M.Fleisch. Taxithelium presented will , , papillae onlyoccurin and undergo changesinshapeduringdevelopment.Baggy this group are ofthe“baggy”type( Câmara andKellogg2010 ) the presence offoliosepseudoparaphyllia.Papillaewithin branches andleavesalternatelydisposedalongthestem can berecognized bytheiroblong,dorsiventrallycomplanate specimens depositedinherbaria.Speciesofthissubgenus and and widespread mossspeciesinthetropics (e.g. were alsorecognized usingmolecularevidence). groups canberecognized within careful studyofdiscrete morphologicalcharacters, twomain the morphologyofpapillae( Câmara andKellogg2010 ) and reported here represents onespecimenperlocality. Abbreviation ofjournalsfollowsBPH.Therepresentative materials section). nately, thesamplingfor this specieswaspoor;seetaxonomy appears toalwayshaveadistinct doublecosta(unfortu- may varywithinthesameindividual, although when present theyare alwaysdoubleandshort.Thisfeature closely related to Câmara andShaw(unpubl. data) showthat which are lackingin tous exothecialcells,andinflatedcolored alarcells,allof from papillose leafcellsis be fewanddifficult tosee(e.g. times maketheleafappeardark(e.g. Taxithelium no leaflackspapillae(asopposedtosometaxainsubgenus laminal cells.Eventhoughthenumberofpapillaecanvary, presence ofseriatelyarrangedpapillaeoverthelumina diagnostic feature ofallspeciessubgenus diagnostic feature ofthesubgenus. presence offilamentous pseudoparaphyllia isconstantand Branching patternsvaryfrom irregular tosubpinnate.The Branches are mostlycreeping, butsometimeslong-ascending. 1F). ( Fig. absence ofacentralstrandistheonlyfeature ofinterest omy showsnovariationwithinthegenus.Thecomplete development ( Câmara andKellogg2010 ). group, andthepapillaehave aconicalshapethroughout their seriately disposedoverthecellluminaalwaysoccurinthis filamentous pseudoparaphyllia.Numerous orfewpapillae presence ofspiral,patently disposed,lanceolateleavesand imens inherbaria.Itsmemberscanberecognized bythe variation inmorphology, yetitisrepresented byfewerspec- ical variationwithinthisgroup. or reversal ofacharacter. Ingeneral,there islittlemorpholog- this subgenusmaylackpapillae,asalaterevolutionaryloss constitute asynapomorphyforthegroup. A fewspeciesin Sbeu Subgenus Subgenera— of Circumscription Costae are highlyvariable.They are usuallyabsent,but The onlyotherSematophyllaceous(s.l.)genuswithpluri- Leaves are highlyvariablewithinsubgenus Branching patternsshowlittleornointraspecificvariation. Species— Vernieri Subgenus of Morphology The secondsubgenus Abbreviation ofauthorsfollows Brummitt andPowell(1992) . T. nepalense Taxithelium ). Thepapillaemaybeveryobviousandsome- Taxithelium ) andincludesabout85%ofall byits distallyverrucose seta,collenchyma- Taxithelium Radulina Taxithelium comprisessomeofthemostcommon Taxithelium Vernieri Results . , butthisgenusisdistinguished T. ramivagum . Thephylogeneticstudiesof comprisesplantswithmore Based ondetailedstudyof ugn subgen. Taxithelium T. levieri ). Taxithelium Radulina h se anat- stem The ), ortheymay tee clades (these Vernieri T. muscicola Taxithelium Vernieri T. planum s the is isnot and A .

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 present they are shortanddouble ( late onlynear the apex.Costaeare mostly absent,butwhen margins canbeentire orserrulate; sometimestheyare serru- ally distinct,beinglongtolong-acuminate orsetaceous.The 0.40–1.85 mmlongand0.15–0.8 mmacross; theapexis usu- be triangular, lanceolate, orovate,withsizerangingfrom the margins, andthisisanexception. wide). 30–85 ally slightlylarger thanthebranchones. not differ insize,butwhentheydo,thestemleavesare usu- T. ramivagum T. damanhurii ( there islessvariation,withsomespecieshavingsmaller mm longand0.08–0.40wide,butwithinaspecies mostly acuminateoracute.Leafsizerangesfrom 0.30–2.00 oblong. Themargins canbeentire orserrulate; theapexis are usuallyconcaveandlanceolate,ormore rarely ovateto alar cellsare nottraditionally associatedwith (s. l.)members,althoughalwaysmuchsmaller. Suchdeveloped sometimes resembling those typicalofSematophyllaceae T. muscicola, in afewspecies( rows ofcellsthatare neithercolored norinflated.However, ( Fig. 1A , B ), andusuallyconsistofonlyoneortwo oped. Theyresemble the alarcellsofsomemembersthe in 9 CAMARA: A RE-CIRCUMSCRIPTIONOFTAXITHELIUM 2011] T. damanhurianum Taxithelium The variationinperichaetial leavesisextensive.Theycan Leaf Cellsare usuallylinear(orlong-linear),varyingfrom The leavesvaryconsiderablyinshapeandsize.They Alar cellsare alwayspresent, butusuallynotwelldevel- Fig. . 1. A. Poorlydevelopedalarcellsin µ Taxithelium kaernbachii m inlengthbutwithlittlevariationwidth(ca.2µm (400×)D.Well-developed alarcellsin (400×). thealarcellsare welldeveloped(Fig. 1C , D ), ) leaves. Thebranchandstemleavesusuallydo T. damanhurianum,T. levieri,T. lindbergii n and T . . kaernbachii hasmore rhombic cellscloseto ) orlarger ( T. isocladum T. planissimum T. lindbergii T. muscicola Taxithelium , whichhas (400×).B.Poorlydevelopedalarcellsin (400×).E.Collenchymatousexothecial cellsin and . and Taxithelium spores varyfrom 10–20µmindiameter. lenchymatous cellsare seeninsomeNewWorld taxa.The from matous exothecialcellsare generallyabsent,butare known Vernieri rostrate operculum, thisfeature ispresent inthesubgenus and although character thatdoesvarywithinthesubgenusissetalength, of thestem. either clustered orevenly distributedontheventralsurface between species. but perichaetialleavescanbeveryusefulindistinguishing is littleinfraspecificvariationintheseperichaetialfeatures nal andapicalcellsare either pluripapilloseorsmooth.There single costa(whenpresent), isanexception).Boththelami- Most sporophyte structures showlittleornovariation.One Rhizoids are usuallyyellowishtoreddish. Theycanbe 263. 1851; Hedw. subsect. Taxithelium num (Brid.) Mitt.,J. Linn. Soc.,Bot.12:496.1869. Müll. Hal.,Bot.JahrbSyst.3:328. 1896. Naturwiss. Ges.1876–77:411. 1878; (Müll. Hal.) A. Jaeger, Ber. Thätigk.St.Gallischen- Mus. Godeffroy 3(6):86. 1874; T. damanhurianum n in Brid. Musc. Recent.Suppl.2:97. 1812. T. planissimum Spruce exMitt.,J.Linn.Soc.,Bot.12:496.1869. Hypnum Taxithelium Spruce, Cat.Musc.14.1867, nom.nud. T. ramivagum aooi Treatment Taxonomic Sigmatella sect. alone(Fig. 1E ), althoughweaklycol- n and hasbeenthoughttolackalong- T. damanhurianum. Sigmatella (400×).C.Well-developed alarcellsin Müll.Hal.,Syn.Musc.Frond 2: T. levieri Trichosteleum (Müll.Hal.)Müll.Hal.,J. Smlry collenchy- Similarly, . Sigmatella F. Cross sectionofstem — TP: TYPE: sect. º Hypnumpla- (Müll.Hal.) Sigmatella T. planum Hypnum

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 1 1. 1. Exothecialcellsnotorstrongly collenchymatous,plantsfrom theOldWorld . 1. Exothecialcellsslightlycollenchymatous,plantsfrom the Americas . Taxithelum [Volume 36 1. Plantswithspreading andlanceolateleaves;pseudoparaphylliafilamentous . 1. Plantscomplanatewithovateleaves;pseudoparaphylliafoliose . . lenchymatous ornot;annulusnotdifferentiated. Operculum long; exothecialcellssubquadrate,thick-walled,slightlycol- dric, constrictedbelowmouthwhendeoperculate, 0.5–1.2mm long. Capsulesinclinedorerect, asymmetric,ovoidorcylin- differentiated. Setaeelongate,slender, smooth,4.8–25.0mm ca. 2µm,lax,pluripapilloseorsmooth;alarcellsnotrarely absent, singleorshortanddouble;laminalcellslinear, 24–95× 0.4–1.8 ×0.15–0.8mm;apexacuminateoraristate;costae ent; archegonia 3–5;perichaetialleaveslanceolateorovate, SYSTEMATIC BOTANY cells notdifferentiated. Perichaetialateral;paraphyses pres- absent; laminalcellslinear, lax,usuallypluripapillose;alar 3–5; perigonialleaveslanceolatetooblong,concave;costae Autoicous. Perigonialateral;paraphysespresent; antheridia sometimes inflatedandcolored. Asexual propagula absent. walled; differentiated alarcellsfew, quadrateinbasalangles, lumina, sometimessmooth,neverunipapillose,thin-orthick- nal cellslinear, 30–85×ca.2µm,seriatelypapilloseoverthe obtuse toacuminate;costadoubleandshortorabsent;lami- late, 0.3–2×0.08–0.70mm;margins entire orserrulate; apex erect towide-spreading, broadly oblong-ovatetolanceo- leaves usuallysimilar, straight,sometimesfalcate-secund, amentous; branchescomplanateorterete. Stemandbranch or not;centralstrandabsent;pseudoparaphylliafoliosefil- ing, irregularly branchedtosubpinnate,shootslong-ascending 10 Taxithelum smalltomedium-sized,forming mats.Stemscreep- . 2. Leavessymmetric;perichaetialleafcellssmooth;margin entire 2. Leavesfalcate;perichaetialleafcellspluripapillose;margin serrulate . Taxithelium damanhurianum (holotype: HYO!; isotype:NY!). Seram. Manusela NationalPark,Sawai, Bryologist 112(3): 589–592.2009.—TYPE:INDONESIA. —TYPE: foliose, leavesovatetoorbicular. num 3. Alar cellsnotpresent orpoorlydeveloped . 3. Alar cellswelldeveloped . . , Hispaniola. Axes complanate,pseudoparaphyllia 4. Leaveswithentire margins . . . 4. Leaveswithmargins serrulate at apex subgenus subgenus 5. Exothecialcellsnotcollenchymatous;perichaetialleavesmore than1mmlong . 5. Exothecialcellscollenchymatous;perichaetialleaveslessthan1 mm long . T. vernieri 6. Leaveslinear-lanceolate; perichaetialleavesnotserrulate atapex . . . . 6. Leavesoblong-lanceolate;perichaetialleavesstrongly serrulate atapex . 7. Leavesmore than0.6 mmlong,oblongtolanceolate 7. Leaveslessthan0.6mmlong,oblongtoelliptical . 8. Perichaetialleaveswithserratemargins . 8. Perichaetialleaveswithentire margins . Db) Besch. (Duby) Taxithelium— Vernieri 9. Leavesstrongly papillose,0.6–1.2mmlong;Pacificislands . 9. Leavesscarcely papillose, 1–1.6mmlong; Africa . 10. Operculum shortlyrostrate; perichaetialleaveswithpluripapillosecellsonlyatapex 2. . 7. 10. Operculum longrostrate; perichaetialleaveswithmostcellspluripapillose . . P. S.Câmarasubg.nov. P. S.Câmara.The TYPE: Key totheSpeciesofSubgenus Key totheSubgeneraof H. Akiyama9329 Hypnum pla-

naked, smooth. finely papillose,7–20µmindiameter. Calyptraecucullate, cilia single,narrow, nodulose.Spores spherical,smoothor papillose, broad, keeled,perforate,aslongtheteeth; endostome withahighbasalmembrane,segmentskeeled, cross-striolate below, papilloseabove,trabeculateatback; teeth narrowly triangular, withzig-zagdorsalmedianline, mm long.Peristomedouble,well-developed,exostome short, rarely long,conicorobliquelyconic-rostrate, 0.3–0.8 lanceolate; margins serrulate; apexacuminate;laminal cells erect-spreading, concave,0.52–0.88 × 0.10–0.20mm,linear- long-ascending, branched.Stem andbranchleavessimilar, cells forSematophyllaceaeby Tan andJia(1999) . in Hypnaceaeanddoesnotfitintotheclassificationofalar Taxithelium tophyte, thesporophyte characters beingconstant. Also in late. Mostvariationinmorphologyisfoundthegame- long-rostrate) operculum andthecalyptraisusually cucul- phyte iswelldeveloped;ithasaconicorapiculate(rarely inflated andoftennotcolored. Thediplolepideoussporo- in othersematophyllaceous(s.l.)genera,thecellsare rarely tiated alarregion ispresent, butisnotaswelldeveloped as tiple papillaearrangedinlinesoverthelumina. A differen- are linearorrhomboid; inmanyspecieseachcell bears mul- disposed andovatetolanceolate,sometimesfalcate.Thecells branches. Theleavesvaryfrom complanate-foliatetospirally creeping plantswithsporophytes onlateralperichaetial subgenus, themissionaryVernier. tose; leaveslanceolatetooblong-lanceolate. (haud foliosis)differt. complanatis) dispositisetpseudoparaphyllisfilamentosis Species of Plants small,forminggolden-yellow mats.Stemcreeping, Etymology— Axes withspreading leaves;pseudoparaphylliafilamen- A subgenus T AXITHELIUM 4 . V , thealargroup appearanceismore similar tothat ERNIERI ...... 5 . Taxithelium Taxithelium ...... 7 . Vernieri ...... subg. . 2 .

5. .

...... 9. . 8. . 3 ...... subg. . refers tothecollectoroftype are yellowish-green todullgreen infoliislanceolatis, spiralis(haud 9 ...... 10 ...... 3. . 6. . 1...... 8 ...... 4. . 6 ...... 10. 11.. T. damanhurianum T. pluripunctatum T. planissimum T. portoricense T. ramivagum T. kaernbachii T. isocladum T. muscicola T. lindbergii Taxithelium T. vernieri T. levieri Vernieri

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 Kurzs.n.( I. Perichaetialleaf J.Branchleaf,HandJscalea; and F, G,I,scaleb. isocladum B andEscalea; A, C,D,scaleb. leaf, C.Leafmargin cells,D.Perichaetialalarregion, E.Perichaetialleaf. but well-developed alarcells,makeitsimilartogenus tous exothecialcells(Fig. 1E ). These,alongwiththerelatively than thoseof smooth. Although theplantsof 11µm across, whilein ent; in leaves are smallerandnarrower. Thespores are alsodiffer- 100–650 m. ( Fig. 3 is restricted totheforests oftheIslandSeram, Indonesia ), where itisepiphyllousandoccursataltitudesof 2. Figure Operculum notseen.Spores smooth,16–20µm diameter. mouth; exothecialcellsquadrate,strongly collenchymatous. CAMARA: asymmetric, ovoid,0.5–0.8mmlong,notconstrictedbelow A RE-CIRCUMSCRIPTIONOFTAXITHELIUM oped inasinglerow. Setae7–10mmlong.Capsulesinclined, pluripapillose atapex;costaeabsent;alarcellswelldevel- setaceous; laminalcellslinearatmidleaf,40–46×ca.2µm, leaves 0.4–0.6×0.14–0.25mm,ovate;margins entire; apex cells. Rhizoidsevenlydistributedalongthestem.Perichaetial ing of1–2rows, 1row ofinflated,vesiculoseandnot colored smooth; costaeabsent;alarcellswelldifferentiated, consist- linear, 70–74×ca.2µm,thick-walled,basalcellssometimes 2011] Fig. Unique to Discussion— Habitat— and Distribution T. damanhurianum . AE A–E: 2. T. muscicola . F. Alar cells,G.Leafmargin cells,H.Perichaetialalarregion, L!) T. damanhurianum T. muscicola Taxithelium damanhurianum. This speciesresembles theyare finelypapilloseandonly12–16 T. damanhurianum isamuchsmallerplant. theyhavelarger spores. Akiyama 9329 are thestrongly collenchyma- T. damanhurianum Taxithelium damanhurianum theyare 16–20µmand A. Alar cells,B.Branch T. muscicola (HYO).F–G: r smaller are Taxithelium bt the but , Taxithelium Radulina ,

Taxithelium magnum Taxithelium epiphyllum mm, narrow-lanceolate; margins entire atbase,serrulate at tributed alongthestem.Perichaetialleaves1.5–2.5×0.30–0.40 ated, consistingof2rows, notinflated.Rhizoidsevenlydis- costae shortanddoubleorabsent;alarcellspoorlydifferenti- 40–50 ×ca.2µm,thick-walled,basalcellssometimessmooth; entire; apexentire, slightlyacuminate;laminalcellslinear, concave, 1.0–1.5×0.28–0.45mm,oblong-lanceolate;margins differentiated; stemleaveslarger andlonger, erect-spreading, long-ascending branched.Stemandbranchleavesslightly Taxithelium isocladioides ther inflatednorcolored liketheonesfoundin prorate setae),andthealarcellsare lessdeveloped andnei- damanhurianum Taxithelium deningeri Trichosteleum werneri 2 2. species. are knownandmoleculardataare sofarunavailable forthis in placement ofthisspeciesandtherefore itishere retained it phylogenetic dataisavailableatthemomenttoverify cladum Plants large, forminggolden-yellowmats.Stemscreeping, Fig. Taxithelium isocladum Taxithelium PAPUA NEWGUINEA,Mt. Antares, Guinea, Bot.10(16):343,plate.31, Fig. 4 . 1964.—TYPE: Winkler 3145 7(2): 136.1928.—TYPE:INDONESIA.Kalimantan. type: BM!). type: L!;isotypes:BM!,NICH!). 1924. —TYPE:MALAYSIA: Malacca. isotypes: BM!,S!). MALAYSIA: Malacca. Range. 11: 443.1925.—TYPE:PAPUA NEWGUINEA.Finisterre Taxithelium werneri isotype: H!). & Cardot, Rev. Bryol.28:111. 1901; INDONESIA. Sumatra,Bangka. Bosch &SandeLac.,Bryol.Jav. 2:173.272.1867.—TYPE: (circles). . Dsrbto mp o for map Distribution 3. E. Werner s.n. . Unfortunately, fewcollectionsofthisplant alsohasasmoothseta( (holotype:H!). M.Fleisch.var. Herzog,Hedwigia61:298.1919.—TYPE: Herzog,Hedwigia49:126.1909. Broth., Mitt.Inst. Allg. Bot.Hamburg. (Herzog)Broth., Nat.Pflanzenfam.(II) Dixon,Bull.Torrey Bot.Club51:243. (holotype:JE!). E. Stresemann 89 (Bosch&SandeLac.)Renauld T. damanhurianum Kurz s.n. laticuspis Radulina Hypnum isocladum (triangle)and Zanten 440 Ridley 737 (holotype:JE!; Zanten,Nova (holotype:L!;. Radulina hasdistally (holo- (holo- T. iso- . No . H.

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 at BM. Luzon, 974 Highlands, Brass13634 Pontianak, ium isatNY, thetypeof T. ramivagum Some specimensof subpercurrent costa,auniquefeature withinthesubgenus. ally longandoftenecostatebutsometimestheyhaveasingle cies insubgenus poorly differentiated alarcellsdistinguishitfrom otherspe- twigs, andasanepiphyll,from sealevelto1,500m. Southeast Asia, inMalesia(Fig. 3 ). Itgrows ontree trunks, lose, 15–20µmindiameter. Figure 2 . obliquely conic-rostrate, ca.0.3mmlong.Spores finelypapil- quadrate, notcollenchymatous.Operculum short,conicor 0.8 mmlong,constrictedbelowmouth;exothecialcellssub- Setae 5–10mmlong.Capsuleserect, asymmetric,ovoid,0.6– gle; alarcellspoorlydifferentiated with2rows, notinflated. thick-walled, pluripapilloseonlyatapex;costaeabsentorsin- apex; apexsetaceous;laminalcellslinear, 50–65×ca.2µm, [Volume 36 SYSTEMATIC BOTANY 12 Beccari147 leaf, J.Perichaetialalarregion, IandGscalea1;F, H,J,scaleb1. levieri and Escalea; A, C,D,scaleb. C. Leafmargin cells,D.Perichaetialalarregion, E. Perichaetialleaf.B Nmnltrl Notes— Nomenclatural Discussion— Habitat— and Distribution Fig. (MO).PAPUA NEW GUINEA.Morobe, Representative SpecimensExamined— F. Alar cells,G.Perichaetialleaf,H.Leaf margin cells,I.Branch Ramos 22166 . AE A–E: 4. (H). Ledrui 2332 (MICH);Java,Tijibodas, Câmara 870 (seenotesunder Taxithelium kaernbachii Its large leaveswithentire margins and (NY). SINGAPORE.sineloco, Vernieri (G);Kalimantan, Serawei, (MO);Sarawak, Taxithelium isocladum . Theperichaetialleaves are unusu- Taxithelium isocladioides Even thoughtheMittenherbar- Kaernbach s.n. Fleisher s.n. T. ramivagum This speciesoccursonlyin Everett s. n. . A. Alar cells,B.Branchleaf, Werner s.n. INDONESIA. Kalimantan, Kalimantan, INDONESIA. Winkler 3145 (JE).MALAYSIA. Genting Ridley 37 (BM).F–G: (M);Selangor, ). mayresemble (JE).PHILIPPINES. (H). (H). (H);IrianJaya, wasfound Taxithelium Câmara Câmara Taxithelium bakeri Taxithelium petrophilum Taxithelium perminutum 3 3. (triangles). likely undercollected due toitssmallsize(leavesare lessthan Malesia, Cameroon, and theSeychelles( Fig. 5 only from afewcollections butfrom awidearea, incluiding ). Itismost 4. Figure rostrate, ca.0.3mmlong. Spores smooth,7–8µmdiameter. rate, notcollenchymatous.Operculum long,obliquelyconic- mm long,constrictedbelowmouth;exothecialcellssubquad- mm long.Capsulesinclined,asymmetric,long-ovoid,0.6–0.8 smooth; costaeabsent;alarcellsnotdifferentiated. Setae4–7 aristate; laminalcellslinear, 24–30×ca.2µm,thick-walled, 0.8 ×0.16–0.30mm,triangular;margins entire; apexlong- Rhizoids clustered beneath thestem.Perichaetialleaves0.4– basal cellssmooth;costaeabsent;alarnotdifferentiated. apex acute;laminalcellslinear, 28–36×ca.2µm,thick-walled; 0.30–0.55 ×0.12–0.26mm,lanceolate-ovate;margins entire; ated, stemleavesslightlybigger, erect-spreading, complanate, freely branched.Stemand branchleavesslightlydifferenti- Taxithelium archboldii Taxithelium meiothecioides Fig. Habitat— and Distribution Plants small,formingdark-green mats.Stemscreeping, PHILIPPINES, Luzon,LagunaProvince, isotypes: FH!,H!,US!). Lamao River, Gard. 8(31):370.1914.—TYPE:PHILIPPINES.Batan, (holotype: H!;isotypes:BR!,PC!,S!) 1897. (not seen);isotypes:BM!,FH!). NEW GUINEA.Sineloco. Broth., Bot.Jahrb.Syst.17:480.1893.—TYPE:PAPUA Pflanzenfam. I(3):1091.1908; (holotype: FH!;isotypes:H!,NICH!). TYPE: PAPUA NEWGUINEA. Baiawa, 3009 7(2): 136.1928.—TYPE:INDONESIA.Serawei, totype H!,designatedhere). Taxithelium kaernbachii . Dsrbto mp o for map Distribution 5. (holotype:H!;isotype:BM!) — TYPE: CAMEROON.Mokundange, Broth., Philipp.J.Sci.13:218.1918. R. Williams 865 E.B.Bartram,Brittonia9:53.1957.— R.S.Williams, Bull.New York Bot. Broth., Bot.Jahrb.Syst.24:267. Broth., Mitt.Inst. Allg. Bot.Hamb. T. kaernbachii Taxithelium kaernbachii L. Kaernbachs.n. (holotype:NY?(notseen); (Broth.) Broth., Nat. Trichosteleum kaernbachii (circles) and Baker 2379, (holotype:H? Brass 22175 Dusén 1030 T. ramivagum H. Winkler isknown — TYPE: (lec-

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 type isnotlostandmaystillbeinH. loan, sointhisparticularcaseIamassumingthattheholo- stay atH,Iwasinformedthatthespecimenprobably on was foundintheBrotherus herbariuminH,butduringmy (PNH). SEYCHELLES.Vallè deMai, geographical range. alar cellsandlarger leaves.Thetwospeciesdonotoverlapin 5 5. (L); IrianJaya, mm long.Spores finelypapillose, 15–20µmdiameter. Figure 4 . chymatous. Operculum long, obliquelyconic-rostrate, ca.0.8 stricted belowmouth;exothecialcellssubquadrate,notcollen- Capsules inclined,asymmetric,ovoid,0.6–0.8mmlong,con- tae absent;alarcellsnotdifferentiated. Setaeca.5.0mmlong. laminal cellslinear, 60–86×ca.2µm,thick-walled,smooth;cos- 1.2–1.6 ×0.2–0.3mm,lanceolate;margins entire; apexaristate; ored. Rhizoidsclustered beneaththestem.Perichaetialleaves cells welldifferentiated, consistingof1–2rows, sometimescol- ca. 2µm,thick-walled;apicalcellspapillose;costaeabsent;alar ceolate; margins entire; apexacute;laminalcellslinear, 60–65× ilar, wide-spreading, concave,0.74–0.90×0.16–0.25mm,lan- Stem creeping, freely branched.Stemandbranchleavessim- Taxithelium horridulum 4. 13 GUINEA. Simbong, (NY). MALAYSIA. Selangor, Gombak, Dusén1030 tous pseudoparaphylliaplacesthisspeciesinsubg. CAMARA: more rhombicA leafcells.However, RE-CIRCUMSCRIPTIONOFTAXITHELIUM thepresence offilamen- resembles plantsinsubgenus bachii false impression ofaserrateleafmargin. Sometimes thestrong papillosityoftheleafcellscangive small sizeincomparisonwithallother low elevations. 0.5 mmlong).Itgrows onrocks andrarely ontree trunks at 2011] loco, 9409 papillose cells,whereas well-developed alarcells,smallerleavesandmore strongly species resemble thoseof stumps andonvolcanicblocks,between2701,170m. (Fiji, Tonga, andNiue).Itgrows ontree trunks, rotten tree to Southeast Asia ( Fig. 6 ), Malesia,andsomePacificislands loco, Levu, Dsrbto ad Habitat— and Distribution Plants medium-sized,forminglax,yellow-opaquemats. Notes— Nomenclatural Discussion— Discussion— Taxithelium lindbergii Representative Specimens Examined Representative SpecimensExamined— Taxithelium levieri (NY).PHILIPPINES. Mindanao, Yuncker 16175 Yuncker 10251 1878; Ber. Thätigk.St.GallischenNaturwiss. Ges.1876–77:412. Rev. Bryol. 28:111. 1901; BM!, FH!,NY!). Banajao, TYPE: PHILIPPINES.Luzon,Province ofLaguna,Mount isotypes: JE!,FH!). [INDONESIA]: Kalimantan: Broth. &Geh.,Biblioth. Bot. 44:23.19.1898. Pflanzenfam. ed.2,11: 443.1925; Buck 7338 alsohasthesmallestspores observedinthegroup. It (H,PC).FIJI.Viti Levu, Hypnum lindbergii Brass 13764 (NY). SAMOA.sineloco, Bur. Sci.Robinson9773 This speciescanbeeasilyidentifiedbyits The convolute margins oftheleavesin this (NY). (NY, MICH).FIJI.Taveuni, Nyman 73 (MICH);Java, Broth., Philipp.J.Sci.8:90.1913.— T. portoricense (NY).PHILIPPINES.Luzon, Taxithelium portoricense (Broth. &Geh.)Broth., Nat. No typeof (A.Jaeger)Renauld&Cardot, Trichosteleum lindbergii SandeLac., Bryol.Jav. 2:172. Taxithelium Onraedt 157 Taxithelium levieri Buck 7108 Bartlett 15933 Câmara 963 IDNSA Bl, Bali, — INDONESIA. Nyman 8752 Beccari 147 Vaupel 152 ooye H; isotypes: H!; ( holotype: haspoorlydeveloped Taxithelium kaernbachii A. C.Smith793 AEON N’Dian, CAMEROON. (NY);Koro, (BR). initsleafshapeand Trichosteleum levieri Taxithelium (MO).PAPUA NEW (MICH).NIUE. sine Taxithelium kaern- (W);Seram, (JE). TONGA. sine (holotype:H!; . . isrestricted Penecilla 10347 T. levieri A Jaeger, A. Vernieri Touw 24745 Smith 1024 (NY);Viti species. — Akyiama TYPE: has .

Taxithelium falcatulum Trichosteleum parvulum Taxithelium argyrophyllum Taxithelium nossianum Trichosteleum capillarisetum Taxithelium robinsonii Taxithelium benguetiae Taxithelium ludovicae Taxithelium alare Taxithelium voeltzkowii (triangles). Fig. CALEDONIA. MontKoghi, Finska Vetensk.-Soc. 48(15):22.1906.—TYPE:NEW Tsurugi-zan, Nat. Pflanzenfam.I(3):1092.1908.—TYPE:JAPAN. 2, 2:988.1902. BR!, H!,FH!,S!). Antsiranana, Bot. Belgique.33(2):131.1895.—TYPE:MADAGASCAR. (holotype: BM?(notseen);isotypes:PC!,BR!). 310. 1880.—TYPE:MADAGASCAR.Nossi-Bé, nated here). Teysmann s.n. pl. 271.1867,nom.illegit.—TYPE:INDONESIA.Java, Kalimantan. Pflanzenfam. (II) 11: 443.1925.—TYPE: INDONESIA. 1922. FH!, NY!,US!,designatedhere). RobinsonBurSci.9864 TYPE: PHILIPPINES.Laguna province, MountBanahao, 10 PHILIPPINES. Luzon,Province ofBenguet, H!; isotypes:M!,PC!). CALEDONIA. Negropo forest, Vetensk.-Soc. 51A (17):28.1909.—TYPE:NEW H!; isotypes:FH!,PC!). PHILIPPINES. Mindoro, s. n. f. 9:14.1908. types: FH!,M!,PC!,S!). (holotype:H!;isotypes:BM!E!,FH!,NY!,US!,S!). . Dsrbto mp o for map Distribution 6. (holotype:H!;isotypes:BR!,PC!,S!). Taxithelium capillarisetum Broth., Philipp.J.Sci.3:28.1908.—TYPE: C. B.Kloss34 Faurie 1400 — Chenagon s.n. Taxithelium parvulum (lectotypeH!;isolectotype:BM!,desig- TYPE: MAURITIUS,sineloco, Broth. &Paris,Öefvers. Förh. Finska Broth., Philipp.J.Sci.8:90.1913.—TYPE: Broth., Philipp.J.Sci.13:218.1918.— Besch., Ann. Sci.Nat.,Bot.sér. 6,10: Broth., ReiseOstafr., Syst. Arbeit. 3:63, Broth. &Paris,Bull.Herb. Boissier, sér. Broth. &Paris, Oefvers.Förh. Renauld&Cardot, Bull.Soc.Roy. (lectotype:H!; isolectotypes: BM!, Dixon,J.Linn.Soc.,Bot.45: 494. (holotype:H!;isotype:PC!) For. Bur. 4476Merritt. , (holotype:BM!). T. levieri. (holotype:PC!;isotypes:BM!, Le Rats.n. A. LeRats.n (Dixon)Broth., Nat. (Broth. &Paris)Broth., (circles) and (holotype:H!;iso- Sanchez B.S. (holotype: Perville s.n. (holotype: Voeltzkow T. vernieri

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 A, B,D,Escaleaand remaining scaleb. Perichaetial alarregion. J. tial leaf,C. 11 leaf, F. Leafmargin cells,G.Perichaetialalarregion, I. Alar cells.. [Volume 36 long. Spores finelypapillose, 15–20µmdiameter. Figure 7 . lenchymatous. Operculum short,conic-rostrate, ca.0.5mm stricted belowmouth;exothecialcellssubquadrate,notcol- Capsules erect, asymmetric, ovoid,0.8–1.1mmlong,con- with 3–4rows ofusually inflated cells.Setae20–22mmlong. thick-walled, smooth;costaeabsent;alarcellsdifferentiated, apex; apexlong-aristate;laminalcellslinear, 60–90×ca.2µm, 0.25–0.30 mm,lanceolate;margins entire atbase,serrulate at evenly distributedalongthestem.Perichaetialleaves1.0–1.8× inflated colored cellsandtheuppernotinflated.Rhizoids cells welldifferentiated, consistingof2rows, thelowerwith sometimes smooth;costaeshortanddoubleorabsent;alar SYSTEMATIC BOTANY laminal cellslinear, 50–55×ca.2µm,thick-walled,basalcells margins entire atbase,serrulate atapex;apexacuminate; spreading, concave,0.95–1.5×0.18–0.30mm,lanceolate; slightly differentiated; stemleaveslarger andlonger;erect- creeping, long-ascendingbranches.Stemandbranchleaves Taxithelium brassii Taxithelium convolutum Taxithelium clastobryoides 14 (H).B,C,E,H,J: Fig. Plants mediumtolarge, forminggolden-yellowmats.Stem 10977 TYPE: INDONESIA.IrianJaya,LakeHabbema, type: BM!). 1935. —TYPE:MALAYSIA. Sarawak, (holotype: BM!). 10(1): 26.1935.—TYPE:THAILAND.Puket, . A, D,F, G,I: 7. T. linbergii (holotype:FH!;isotypes:L!,MICH!). Taxithelium lindbergii E.B.Bartram,Lloydia5:288.56.1942.— alarcells,E. Taxithelium muscicola T. linbergii Dixon,J.Linn.Soc.,Bot.50:130.41. Dixon,J.SiamSoc.,Nat.Hist.Suppl. T. linbergii leafmargin cells. B. Taxithelium lindbergii A. Branchleaf,D.Perichaetial branchleaf.H. Everett s.n. Teysmann s.n. Kerr 512b T. linbergii Whitelegge perichae- (holo- Brass (H).

THAILAND. Mt.Luang, Ile Mahé, Tremblet, VIETNAM. BaoLoc. Laguna, but isotypeswere foundin PCandBR. found inBM(where Emile Bescherelle’s herbariumishosted), in BM.Thetypespecimenfor barium isatNY, thetypeof matches thatineachprotologue. EventhoughtheMittenher- mens. However, theadditionalinformationonspecimens collection numbers,eitherintheprotologue oronthespeci- Mindanao, Onraedt272 Holtman1931 Genting Highlands, Sumatra, 756 gii smooth perichaetialleafcells. by itsdifferentiated alarcells,serrateleaves,longseta,and T. lindbergii acters, therefore Idecidedonthewidercircumscription of in thiscomplex,norare there anyqualitative(discrete) char- morphological traitseparatespreviously recognized species ric data(availableuponrequest) showthatnoquantitative being proposed thatare nowsynonymized.Themorphomet- smaller butwider. collections canbegathered; however, suchplantsare slightly T. parvulum have leavesthatcanbeslightlyfalcate.Thetypespecimenof larger plants.Lowlandspecimensare usuallysmaller and (Thailand), Mt.Konghis(NewCaledonia)andelsewhere, are Mt. Kinabalu(Borneo),Halcon(Philippines),Luang tion. Plantsthatoccurathighelevation(1,000–2,000m)on identification of Seychelles, Madagascar, Vietnam, andJapan. Samoa, andPacificIslands.Itisalsopresent intheMascarenes, uted ( Fig. 8 ), from SriLankatoMalesia,NewCaledonia,Fiji, 29980 Nmnltrl Notes— Nomenclatural The variabilityofthistaxonhasresulted inmanynames Discussion— Habitat— and Distribution Fig. , , (NY).INDONESIA.Bangka, Representative SpecimensExamined— T. falcatulum,T. nossianum, (DUKE);Ovalau(Mt.Tana), . Dsrbto mp o for map Distribution 8. Robinson B.S.17077 Touw &Snoek25306 Arts 92/18, Decorié s.n. Ramos B.S.14894 (BR).PHILIPPINES.Mindoro, (NY);Sarawak, from Japanisincludedin proposed here. Thisspeciescanberecognized There hasbeenconfusionoverthecorrect Câmara 878 T. lindbergii (BR),St.Phillippe, Tixier s.n. (BR).SRILANKA.SineLoco, Touw 11800 (NY);Silipan, (L).IrianJaya,sinelegit(BM).MALAYSIA. Richards 2563 (NY);Luzon, (PC) The typespecimensof (MO);Selangor, T. lindbergii. Taxithelium convolutum duetoitsconsiderablevaria- Kurz s.n. (MICH,NY); Puket, and Smith 7719 This speciesiswidelydistrib- T. nossianum Arts 11/59 Phillips 16 (BM).MAURITIUS.LePouce, T. argyrophyllum FIJI. Viti Levu,

MacGregor B.S.19919 (L);Java, T. lindbergii Salgado Edw, 12360 (DUKE);Taveuni, HBR 4022 (BR).SEYCHELLES. (MICH).REUNION. Besch.wasnot Thwaites 217 Schif. 12105 Kerr 512b (NY);Sabah, untilmore wasfound T. lindber- haveno Whitehouse (BR); (NY); (BM). Smith (G). (S); Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 (circles). Island, in loanedmaterial. syntype rect nameis cicolum leaves withentire margins, are adistinctivecombination. oped alarcells,alongwithlarge andnarrowly lanceolate larger (seenotesunder Taxithelium muscicola Tan etal.(1996) studiedthisspeciesforthe mens from mainland Australia but Ramsay etal.(2002) and Island; however, itwaslaterfoundinQueensland( Fig. 9 ). plant wasonceconsidered tobeendemicLord Howe phytic orepiphyllousinmontanerainforest. Thisdistinct papillose, 12–16µmdiameter. Figure 7 . rate, notcollenchymatous.Operculum notseen.Spores finely mm long,constrictedbelowmouth;exothecialcellssubquad- 10–25 mmlong.Capsuleserect, asymmetric,ovoid,0.6–0.8 15 at apex;costaeabsent;alarcellspoorlydifferentiated. Setae cells linear, 72–74×ca.2µm,thick-walled,pluripapilloseonly gins entire atbase,serrulate atapex;apexfiliform;laminal stem. Perichaetialleaves1.5–2.0×0.18–0.3mm,ovate;mar- yellowish orhyaline.Rhizoidsevenlydistributedalongthe alar cellsdifferentiated, consistingof1–4rows, vesiculose, cells sometimessmooth;costaeshortanddoubleorabsent; nate; laminalcellslinear, 60–70×ca.2µm,thick-walled,basal slightly falcate-cuspidate;margins entire; apexlong acumi- spreading, concave,1–2×0.15–0.18mm,narrow-lanceolate, slightly differentiated; stemleaveslarger andlonger, erect- CAMARA: A RE-CIRCUMSCRIPTIONOFTAXITHELIUM creeping, pinnatelybranched.Stemandbranchleaves 6. 2011] Fig. Notes— Nomenclatural Discussion— Habitat— and Distribution Plants medium-sized,formingpale-yellowmats.Stems Taxithelium muscicola Representative SpecimensExamined— designated here). Howe Island,Mt.Gower, Vetensk.–Soc. 42:117. 1900.—TYPE: AUSTRALIA. Lord Trichosteleum muscicola & W. B.Schofield. Austral. Syst.Bot.9:324.1996; . Dsrbto mp o for map Distribution 9. Watts 419 ; however, becauseitisanouninapposition,thecor- Maiden 218 muscicola (NSW), Unfortunately, Iwasunabletostudyspeci- wasnotfoundintheherbariavisitedor resembles Watts 403 (seeICBN Art. 23.5).Unfortunatelythe T. damanhurianum The epithetwasfirstspelled T. muscicola (Broth.) B.C.Tan, H.P. Ramsay Broth., Öefvers.Förh. Finska (PC). T. Whitelegge11 Taxithelium muscicola T. damanhurianum (triangles)and UTAI. od Howe Lord AUSTRALIA. . h well-devel- The ). Flora ofAustralia (lectotypeH!, T. planissimum , butitis isepi- mus- . apex acuminate toaristate;laminalcellslinear, 72–85×ca.2µm, falcate, lanceolate-ovate; margins subentire, serrulate atbase; strongly papillose,wide-spreading, 0.70–1.2×0.20–0.40mm, entiated; stemleaveslarger andlonger, branch leavesmore ing, freely branched.Stemandbranchleavesslightly differ- Taxithelium patulifolium Taxithelium thelidiellum 1106 7 7. 8 8. loco, Palawan, (JE); Perak, (Brastagi), 15–20 µmdiameter. Figure 10 . long, conic-rostrate, ca.0.8 mmlong.Spores finelypapillose, ecial cellssubquadrate,notcollenchymatous.Operculum ric, ovoid,0.6–0.8mmlong,constrictedbelowmouth;exoth- inflated. Setae5–7mmlong.Capsulesinclined,asymmet- costae absent;alarcellspoorlydifferentiated in2rows, not cells linear, 35–40×ca.2µm,thick-walled,pluripapillose; mm, lanceolate;margins serrulate; apexacuminate;laminal tributed alongthestem.Perichaetialleaves1.2–2.0×0.30–0.45 tiated, consistingof1row, notinflated.Rhizoidsevenlydis- costae shortanddoubleorabsent;alarcellspoorlydifferen- 60–65 ×ca.2µm,thick-walled,basalcellssometimessmooth; olate; margins entire; apexacuminate;laminalcells linear, spreading, concave,0.75–1.5×0.20–0.40mm,oblong-lance- slightly differentiated; stemleaveslarger andlonger, erect- creeping, long-ascending,branched.Stemandbranchleaves Taxithelium wewakense Taxithelium ramicola on bark. grows almostexclusively ontwigsorasanepiphyll,rarely Binohan (Palawan,Philippines)atabout1,000m( Fig. 9 ). It between sealeveland300m,butitwasalsocollectedonMt. in lowlandsfrom SriLanka toSE Asia (Malesia),andVietnam, the perichaetialleavesof long-rostrate opercula andmuchlongerseta.Furthermore, oped alarcells,butitcanbedistinguishedbythedistinct in itsleafshapeandsizetheabsenceofwell-devel- mum cells attheapex,whileperichaetialleavesof Plants medium-sized, forminglax,goldenmats.Stemcreep- Dsrbto ad Habitat— and Distribution Plants medium-sized,formingpale-yellowmats.Stems Discussion— Taxithelium pluripunctatum Representative SpecimensExamined— Taxithelium planissimum (S).MALAYSIA. Selangor, Thwaites 217 (holotype: PC!;isotypes:H!,NY!) FRENCH GUIANA.Saint-Jean-du-Maroni SineLegit (Holotype: BM?(notseen);isotypes:H!,NY!). —TYPE: GUADELOUPE.Trois Diables, (holotype: PC!;isotype:NY!). 184. 1890.—TYPE:MARTINIQUE. Ste-Marie, Renauld &Cardot, Bull.Soc.Roy. Bot.Belgique29(1): Buck, Moscosoa2:60.1983; 1910.—TYPE: SRILANKA.Hiniduma, Angoram Area, Angoram Area, TYPE: PAPUA NEWGUINEA.SepikDistrict,Wewak- type: H!;isotypes:BM!,FH!,NY!,S!,US!). PHILLIPINES. Polillo, type: H!;isotypes:BR!,JE!,S!). havesmoothcells. Ebalo 391 Holtmann 25327 Ridley 213 (NY).VIETNAM.LaoCai. (MICH). SRILANKA.Hiniduma, Taxithelium planissimum (H);Sabah, Broth., Philipp.J.Sci.8:91.1913.—TYPE: Robbins 2026 E.B.Bartram,Brittonia13:378.1961.— (NY);Seram, Thér., Ann. Bryol.7:160.1934.—TYPE: Besch.,J.Bot.(Morot) 6:10.1902. Câmara 960 T. isocladum McGregor Bur. Sci.10509 Holtmann 1931 Taxithelium planissimum Broth., Hedwigia50:141. (holotype:FH!;isotype:L!). Trichosteleum pluripunctatum Akiyama 9906 (Renauld&Cardot) W. R. (MO);Malacca, Moctier s.n. resembles NOEI. Sumatra INDONESIA. havepluripapillose (NY).PHILIPPINES. Herzog 3979 (NY);Java, Herzog 20 (S). P. Duss1364 T. isocladum T. planissi- Werner s.n. Bordaz 1 (H);sine occurs Zollinger (holo- (holo-

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 by itspluripapillose leafcellsandpoorlydifferentiated alar T. pluripunctatum pluripunctatum lose leaves,andsmoothperichaetial leafcells. T. portoricense one ofthetwospeciesthatoccur intheNewWorld. However and 900m. trunks, limestoneandhumus,atelevationsbetweensealevel I havenotseenspecimensfrom theseareas. Itgrows ontree and Central America ( Buck 1998 ; and theWest Indies( Fig. 12 Sharp etal.1994 ). Itisalsoreported from Mexico ) but restricted totheNewWorld, beingfoundinSouth America µm diameter. Figure 11 . conic-rostrate, ca.0.3mmlong.Spores finelypapillose,15–20 or notcollenchymatous.Operculum short,conicorobliquely stricted belowmouth;exothecialcellssubquadrate,slightly Capsules inclined,asymmetric,ovoid,0.6–0.8mmlong,con- absent; alarcellspoorlydifferentiated. Setae4.9–5.1mmlong. 60–80 ×ca.2µm,thick-walled,poorlypluripapillose;costae rulate atapex;apexacuminatetoaristate;laminalcellslinear, leaves 1.0–1.5×0.18–0.30mm,long-triangular, margins ser- not inflated.Rhizoidsclustered onlowerstem.Perichaetial double orabsent;alarcellspoorlydifferentiated in2rows, thick-walled; apicalcellsusuallysmooth;costaeshortand s. n. gin cells,J.Perichaetialleaf.FandJscalea;remaining scaleb. mum a; and A, C,D,scaleb. margin cells,D.Perichaetialalarregion, E.Perichaetial leaf.BandEscale [Volume 36 SYSTEMATIC BOTANY 16 Some speciesof Discussion— Habitat— and Distribution Fig. (G). . F. Branchleaf,G. Alar cells,H.Perichaetialalarregion, I.Leafmar- 0 –: A–E: 10. differs in itsinvoluteleafmargins, lesspapil- alsohasslightlyfalcateleaves. Taxithelium vernieri This species, togetherwith ( Buck 1998 ), but thelattercanberecognized Mittenothamnium McGregor B.S.10509 . A. Alar cells,B.Branchleaf,C.Leaf Taxithelium pluripunctatum Henn.mayalsoresemble (H).F–G: T. portoricense, Taxithelium planissi- Taxithelium Vernier is is 8390 region, J.Perichaetialleaf.HandJscalea;F, G,I,scaleb. F. Alar cells,G.Leafmargin cells,H.Branchleaf, I.Perichaetialalar D scalea;and A, C,D,scaleb. C. Leafmargin cells,D.Perichaetialleaf,Ealarregion, Band MICH). PUERTO RICO.Luquillo, Djan-Chékar94–510 Sofaia, ium isatBM,notypeof lose leafcellsandbetterdevelopedalarcells. cells, whereas walled; apicalcells usuallysmooth;costaedouble andshortor apex acuminate; laminalcellslinear, 44–50×ca.2µm,thick- oblong-lanceolate; margins subentire, serrulate onlyatbase; strongly papillose,wide-spreading, 0.70–1.2×0.20–0.35mm, entiated; stemleaveslarger andlonger, branch leavesmore ing, freely branched.Stemandbranchleavesslightly differ- 9. (NY) de Saül, 13 Isla Gorgona, & Buck20321A Uamatã, information onthespecimenmatchesthatofprotologue. tor, eitherintheprotologue oronthespecimen,butother type specimensof (NY, MICH).FRENCHGUIANA.Dt.Laurent-du- Maroni: Commune Fig. Nmnltrl Notes— Nomenclatural Plants medium-sized, forminglax,goldenmats.Stemcreep- Taxithelium portoricense Representative Specimens Examined–— (NY). ; ignated here). 8390 37. 1927.—TYPE:PUERTO RICO.Cidra, CommunedeMatoury 1 –: A–E: 11. Allorge s.n. Buck 3148 Buck 18349A (lectotype:NY!;isolectotypes: FH!,PC!.MICH!,des- Rudas &Aguirre 130 (NY); Roraima:BocadaMata, Mittenothamnium Taxithelium pluripunctatum (NY);Bahia:Ilhéus, (NY). (MICH).MARTINIQUE. Absalon, (NY);Communede Appropague-Kaw, T. patulifolium T Bordaz 1 Even thoughBescherelle’s herbar- Buck 32904 . . thelidiellum (NY).DOMINICA.FourHunk, Buck 4192 Boom &Mori870 specieshassmoothorproru- R. S.Williams, Bryologist30: (PC).F–J: havenodataonthecollec- (NY, MO).GUADELOUPE. . A. Alar cells,B.Branchleaf, Buck 1948 (NY).TRINIDAD. Aripo, wasfoundthere. The RZL Aaoa: Rio Amazonas: BRAZIL. Taxithelium portoricense (NY),Uruçuca, (Y. COLOMBIA. (NY). Welch 21336 E. G.Britton E. G Buck 37799 . . Fishlock Britton (NY, Vital .

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 port itsplacementhere. molecular evidence(Câmara andShawunpubl.data)sup- but boththekeycharacteristicsofsubgenus T. pluripunctatum ichaetial leafcellsthatlackpapillae (seealsocommentsunder and more symmetricleaveswithinvolutemargins, andper- entiated from close to more oblongleafithasbeensuggestedthat toricense Pines, & Earle1054 Americas (theotherbeing is of cited. Theirspeciesdescriptionisunclearandtheillustration although itwasnotpossibleformetostudythespecimenthey from theBrazilian Amazon byLisboaandIlkiuBorges (1997), dead logs,betweensea-leveland1,500m.Itwasalsocited islands intheCaribbean( Fig. 12 ), andgrows ontwigsand lose, 15–20µm. Figure 11 . obliquely conic-rostrate, ca. 0.3mmlong.Spores finelypapil- slightly ornotcollenchymatous.Operculum short,conicor long, constrictedbelowmouth;exothecialcellssubquadrate, mm long.Capsulesinclined,asymmetric,ovoid,0.6–0.8 cells poorlydifferentiated in2rows, notinflated.Setae4.9–5.1 ear, 80–90×ca.2µm,thick-walled,smooth;costaeabsent;alar triangular; margins entire; apexacuminate;laminalcellslin- lower stem.Perichaetialleaves0.80–1.2×0.18–0.45mm,long- absent; alarcellspoorlydifferentiated. Rhizoidsclustered in 17 CAMARA: A RE-CIRCUMSCRIPTIONOFTAXITHELIUM 2011] Discussion— Habitat— and Distribution Fig. Representative SpecimensExamined— T. pluripunctatum E. P. Killip43735 2 itiuin a fr for map Distribution 12. (circles). T. planum, (NY),CaletaCocodrilos, T. pluripunctatum This speciesisoneoftwogrowing inthe ). ofthesubgenus (FH,S).DOMINICA. Picard Valley, ; therefore Iamexcludingthisrecord. Taxithelium portoricense T. pluripunctatum T. pluripunctatum This speciesisrestricted to byitslessstrongly papillose Britton etal.15281 UA E Ynu, Yunque, El CUBA. Taxithelium (triangles)and ). Becauseofits T. portoricense canbe differ- uk 1998), (Buck Vernieri (NY),Islede W. R.Elliot Underwood T. por- and is Middle IslandParish, 4101 Repressa Dam, 961c Island, (FH). JAMAICA.PortlandParish, Taxithelium ramivagum 10. Taxithelium falcifolium Taxithelium protensum Taxithelium kuniense Taxithelium francii 11. SIERRA LEONE.Freetown, Mt.Oriel. Mavenga, REPUBLIC OFCONGO.Kivu,Luhoho, 797 bark from sealevelto875 m. ( Fig. 5 ); thelastcollectionwasmadein1920s.Itgrows on (mostly types),thisspeciesisrestricted totropical Africa Capsules unknown. Figure 13 . differentiated in2rows, notinflated.Setaeca.10mmlong. ca. 2µm,thick-walled,smooth;costaeabsent;alarcellspoorly margins entire; apexsetaceous;laminalcellslinear, 32–40× Perichaetial leaves1.0–1.5×0.24–0.30mm,narrow-lanceolate; rows, notinflated.Rhizoids evenlydistributedalongthestem. costae absent;alarcellspoorlydifferentiated, consistingof2 75–80 ×ca.2µm,thick-walled,basalcellssometimessmooth; entire; apexentire orslighltyacuminate;laminalcells linear, concave, 1.0–1.6×0.25–0.40mm,oblong-lanceolate;margins differentiated, stemleaveslarger andlonger, erect-spreading, long-ascending, branched.Stemandbranchleavesslightly Taxithelium theriotii Taxithelium nitidulum smooth ( as longin in itsleafsizeandshapebutthecellsare abouttwice Discussion— Habitat— and Distribution Plants large, forminggolden-yellowmats.Stemscreeping, (PC).CÔTED’IVOIRE.Hourotte, Representative SpecimensExamined— Taxithelium ramivagum (FH!).DOMINICANREPUBLIC.Prov. Samará, (FH,NY);LasCruces, Taxithelium vernieri type: H!;isolectotype:PC!,designatedhere). 1897.—TYPE: CAMEROON.Mbanga, Pauahi Bishop Mus.10(10):14.1933.—TYPE: [FRENCH (holotype: BM!). Wales 55:297,f.24.1930.—TYPE:FIJI.Sine loco, here). 1372 CALEDONIA. IledesPins,forêt deGadge, Vetensk.-Soc. 53A(11): 36.1911.—TYPE: NEW s. n. 1910.—TYPE: NEWCALEDONIA.MontKoghis, M!, PC!). 1875.—TYPE: TAHITI. France 45:123.1898; (holotye: PC!). 364. f.16.1925.—TYPE:GABON.Benzé, mabey, Bryol Lichénol.5:207.1933.—TYPE:GABON.Malongo- CALEDONIA. Thi, Finska Vetensk.-Soc. 48(15):23.1906.—TYPE: NEW types: BM!,NY!,PC!). Steere 2767 (holotype:PC!;isotypes T. isocladum Le Testu s.n. (lectotype: H!;isolectotypes:PC!,S!,designated T. ramivagum Le testu6768 Allard 17275 (NY).PUERTO RICO.CaribbeanNationalForest, Taxithelium ramivagum Thér., Bull. Acad. Int.Géogr. Bot. 20: 103. Buck 29826 P. delaVarde, Bull.Soc.Bot.France72: (M).LIBERIA.SinoeDistrict, Broth. &Paris,Öefvers.Förh. Finska haspluripapillosecellsatapex). Broth. var. Dixon,Proc. Linn.Soc.NewSouth E.B. Bartram,Occas.Pap.Bernice Broth. &Paris,Öefvers.Förh. Steere 6361 (NY).GUADELOUPE.Sofaia, (holotype Le Rats.n. andtheperichaetialleafcellsare Hypnum vernieri (NY). (Duby)Besch.,Bull.Soc.Bot. D. Vernier s.n. Crosby 13743 Broth., Bot.Jahrb.Syst.24:266. Known from fewcollections (FH,MICH).ST. KITTS.St.Thomas Arnell 2306 : elongatum BM!,FH!,H!). : Jolly s.n. PC!). AEON Eud, Ekundu, CAMEROON. (holotype:H!;isotypes: Fzaire s.n. resembles (NY).MEXICO.Cozumel (PC) Duby, Flora58:285. (holotype:G!;iso- P. delaVarde, Rev. (S).DEMOCRATIC Staudt 277 Baldwin 11337 (BR).GABON. Buck 8701 Le testu5324 Louise LeRat T. isocladum Le Gallo444a (lecto- Steel 4 (NY); Franc Dusen (PC). Buck

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 a; and A, C,D,scale b. margin cells,D.Perichaetialalarregion, E.Perichaetial leaf.BandEscale leaves. Itisbyfar themostcommonspeciesof lanceolate leaves withentire margins andsmoothperichaetial Polynesia), Micronesia andMarquesas ( Fig. 6 ). islands ofthePacific,NewCaledonia, SocietyIslands(French trees, betweenthesealeveland700m,isrestricted tothe forests, usuallyondeadlogs,sometimesbarkof living mm long.Spores finelypapillose,15–20µm. Figure 10 . Operculum short,conicorobliquelyconic-rostrate, ca.0.3 ecial cellssubquadrateorrectangular, notcollenchymatous. ovoid, 0.6–0.8mmlong,constrictedbelowmouth;exoth- ated. Setae14–16mmlong.Capsulesinclined,asymmetric, thick-walled, smooth;costaeabsent;alarcellsnotdifferenti- apex; apexlong-aristate;laminalcellslinear, 50–70 ×ca.2µm, 0.40–0.50 mm,lanceolate;margins entire atbase,serrulate at evenly distributedalongthestem.Perichaetialleaves1.2–1.6 × cells poorlydifferentiated, consistingof1–3rows. Rhizoids sometimes smooth;costaeshortanddoubleorabsent;alar nal cellslinear, 58–60×ca.2µm,thick-walled, basalcells lanceolate; margins entire; apexacuminateoracute;lami- erect-spreading, concave,0.6–1.2×0.15–0.30mm,oblong- long ascendingbranched.Stemandbranchleavessimilar, [Volume 36 SYSTEMATIC BOTANY 18 Discussion— Habitat— and Distribution Plants small,forminggolden-yellowmats.Stemscreeping, Fig. type: FH!;isotypes:L!,US!). POLYNESIA]. Eiao,furasforest, 13. 13. Taxithelium ramivagum Taxithelium vernieri Staudt 277 (H). . A. Alar cells,B.Branchleaf,C.Leaf This speciesgrows indense canberecognized byits W. B.Jones1522 Taxithelium (holo- in the the in Taxithelium confusum bilobed leaves,whichare notfoundin Taxithelium bilobatum than in bles thewidespread type. description orillustration thatwouldallow placement ofthis Taxithelium glabrisetum resemble thosefoundin Taxithelium shows unipapillosecellsandconsequentlyisprobably nota Taxithelium decrescens the descriptionof of whichare nowplacedin theunrelated genus Cardot assignedthisspecies tosection gle papillapercell;therefore itisunlikelytobea Taxithelium aureolum protologue isuninformative. known tooccurthatfarsouth.Thedescrptionprovided inthe men, itisprobably nota Taxithelium anderssonii ing thebombingofBerlinherbariumin1943. Müller Hallensisnamesandwere mostlikelydestroyed dur- I lookedinalltheherbariacitedmethods.Someare ing namesitwasimpossibletolocatethetypespecimens. Whittier&Miller797 in Pacific. Itcanresemble specimensof Puluwatt, (NY). VANUATU. Sineloco, shore, et Baumann9152 M); Mé Aoui, Hiva, Moorea, Vanua Mbalavu, (G). FIJI.Vanua Levu, According totheoriginaldescription,specimenhas The descriptionandillustrationprovided byCardot resem- Unfortunately theprotologue does notprovide agood The illustrationprovided intheoriginaldescription According totheoriginaldescriptionthisplanthasasin- Even thoughitwasnotpossibletolocatethetypespeci- Types NotSeeninGenusTaxithelium— T. vernieri Representative SpecimensExamined— Marie, 39: 471.1915.—TYPE:MADAGASCAR.IleaSainte- Burkill13007 Settlem. 4:35.1926.—TYPE:MALAYSIA. Malacca,Perak, type B?,probably destroyed). 329. 1896.—TYPE:SAMOA. Olosina, 1898. Bat. s.n. 2: 168.266.1866.—TYPE:INDONESIA.Sulawesi, I (3):1092.1908. type: PC?). 319. 1913.—TYPE:JAPAN. Sineloco. (holotype S?). TYPE: CHILE.PortFamine,Magalhaens, Förh. Kongl.SvenskaVetensk.-Akad. 29(4):15.1872.— (3): 1237.1909. Jordan s.n. Yuncker 9517 Taxithelium Sloover 20946 Whittier &Miller1108 Sigmatella glabriseta . Theillustrationoftheperichaetiaalsodoesnot Boivin s.n. Guillaumin etBaumann10519 are notwelldevelopedastheyare in (holotype:L?). (PC);Mt.Coughi, (NY).NEWCALEDONIA. Sineloco, Smith 1478 (NY, MICH).TONGA. IslandofEua, . (holotype:BM?). (NY); Atoll Ulul, T. aureolum . (NY);Tahiti, Smith 1618 Plagiothecium anderssonii Cardot, Hist.Phys.Madagascar, Mousses Cardot, Bull.Soc.Bot.Genèvesér. 2,5: var. Taxithelium lindbergii. Hypnum decrescens (SandeLac.)Broth., Nat.Pflanzenfam. (holotype:PC?). (Ångstr.) Broth., Nat.Pflanzenfam.I (Müll.Hal.) Paris,IndexBryol.1261. (NY).FRENCHPOLYNESIA. SocietyIslands, Campbell 3 Taxithelium scabrifolium Taxithelium (NY); Atoll Iruh, (NY);Viti Levu, Balansa 2579b Müll.Hal.,Bot.Jahrb.Syst.23: wouldfitinto Vernier 1316 (BM). Whittier &Miller1075 . (PC);Mé Ammeri, T. levieri Dixon,Gard. Bull.Straits IRNSA Etn Island, Etten MICRONESIA. , sincethegenusisnot Taxithelium. SandeLac.,Bryol.Jav. Anastigma (PC).SAMOA.Southeast (G);Marquesas, Nuku

Whittier &Miller7471 Hirosaki 18. Flechten 91 , butthealarcells Smith 8548 Ågt. Öfvers. Ångstr., o te follow- the For Phyllodon L. LeRat1403 Andersson s.n. Phyllodon T. levieri Yuncker 15392 Taxithelium , allspecies

(Y, Atoll (NY), Guillaumin (DUKE); . (holo- . (holo- rather Ludg. , and , . (H, .

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 with ferring inal descriptionisvague.HoweverBrotherus, whentrans- Taxithelium ivoreanum herbarium. unsuccessfully tolocatethisspecimeninTixier’s Vietnam. IvisitedtheherbariuminParis(PC)twiceandtried Taxithelium isocladum herbarium. men inTixier’s Paris (PC)twiceandtriedunsuccessfullytolocatethisspeci- nonpapillose speciesof leaf cells,buttheillustrationprovided doesnotresemble the Taxithelium inerme Taxithelium specimen havesinglepapillaeandtherefore itisprobably nota from Japan( to 19 CAMARA: A RE-CIRCUMSCRIPTIONOFTAXITHELIUM Taxithelium herpetium 2011] smooth leafcells. As theonlyspeciesof Taxithelium laeve barium wastransferred to NY. listed asnonvide,soitwasalready lostwhenMitten’sher- described byWilliam Mitten”( Thiers 1992 ) thisspecimenis Phyllodon Taxithelium natans locate thisspecimenduringmy visittoBM History Museum(BM)onapermanent loan.Iwasunableto longer, sincethey were senttotheherbarium oftheNatural that thetypeisatK,there are nomosscollectionsatKewany of subgenus the twotobecloselyrelated. Thedescriptionwouldfitspecies Taxithelium liukiuense during mytwovisitstoPC. member ofthegenus.Itwasnotpossibletolocatethistype The original descriptionandillustrationprovided matches This speciesisprobably a The originaldescriptionsaysthatthisplanthassmooth The originaldescriptionstatesthattheleafcellsoftype The originaldescriptionsaysthatthetype specimenhas At the“Indicestospeciesofmossesandlichens Although thetypeinformationinprotologue states Sakurai compares thisspecieswith T. glossoides,T. similans type: PC?). McIvors.n. Bot., Suppl.2:105.1859.—TYPE:NEPAL. Mont.Nilghiri, 1093. 1908. 1966.—TYPE: Vietnam. Quang-Binh, var. Waterfalls, 1971 [1972].—TYPE:THAILAND.Chandhaburi,Plew legit. (holotype:B?,probably destroyed). Godeffroy 3(6):84.1874.—TYPE:SAMOA.Tutuila, sine I(3): 1091.1908. type PC?). 1912.—TYPE: JAPAN: Tosa, Arakusa, Cintra, 70. 1901.—TYPE: BRAZIL.RiodeJaneiro, Morro da 28: 111. 1901. 2170 1932.—TYPE: [JAPAN]. Liukiu,Nishi-Omotejima, T. isocladum Stereodon ivoreanus . (holotypeBM?,notseen). . vietnamense ; unfortunately, noillustrationofthisplantexists. E.Ule 161 Taxithelium T. lindbergii Tixier 965 (holotype:NY?). Cardot, Bull.Soc.Bot.Genèvesér. 2,4:387. Stereodon ivoreanus , butthisspeciesissofarunknownfrom (Müll.Hal.)Renauld &Cardot, Rev. Bryol. Tixier, Rev. Bryol.Lichénol.38:159,f.8. Sigmatella natans (Bosch.&SandeLac.)RenauldCardot Tixier, Rev. Bryol.Lichénol.34:171. (Müll.Hal.)Broth., Nat.Pflanzenfam. . (holotypeB?, probably destroyed). Sakurai,Bot.Mag.(Tokyo) 46:505. (Mitt.)Broth., Nat.Pflanzenfam.I(3): Hypnum herpetium (inwhich Taxithelium ) ispapillose, . (holotype:PC?). and to Phyllodon Taxithelium, T. ligulatum T. nepalense . Ivisitedtheherbariumin Müll.Hal.,Hedwigia40: Mitt.,J.Proc. Linn.Soc., . Unfortunatelytheorig- T. nepalense T. laeve Müll. Hal.,J.Mus. considered itclose Okamura s.n. Maunier s.n. Taxithelium , allnowplacedin belongs). maynotbea , considering , known (holo- (holo- Y. Dor Taxithelium rhizophoreti provided. tinctis sparsimtenuiterpapillosi,”butnoillustrationwas Taxithelium plumularia Taxithelium vescens Taxithelium orthothecium Taxithelium oophyllum herbarium in1943. of bothwere probably lostduringthebombingof theBerlin known tooccurinRiodeJaneiro andMinasGerais.Thetypes lum genus Taxithelium Brotherus, whentranferring tion. Theprotologue isbroad, butitdoesfit this planttothesamegroup as there isnoillustrationavailable.Brotherus howeverassigned Taxithelium spathulifolium genus inSamoais Taxithelium planum Vernieri The mostcommonofthespecies alare variable Taxithelium planum omyms. Inthiscase,unfortunatelythetypeismostlikelylost. varieties havebeendescribedandIbelievethattheyare allsyn- Taxithelium planum Taxithelium The originaldescriptionisvague;itsaysthecellsare “indis- See commentsunder The originaldescriptionsofboth There isnoillustrationaccompanyingtheoriginaldescrip- Unfortunately theoriginaldescriptionismostlyuselessand The originaldescriptionmatches I wasunabletolocatethetype,butdescriptionfits match (= I(3): 1091.1908. s. n. Frond. 2:684.1851.—TYPE:INDONESIA.Java, I(3): 1092.1908. 1878. —TYPE:SAMOA.Tutuila, Thätigk. St.GallischenNaturwiss.Ges.1876–77:414. I(3): 1091.1908. Uberaba, Hedwigia 40:70.1901.—TYPE:BRAZIL.MinasGerais, Bryol. 28:111. 1901. legit. ( holotype B?,destroyed). Godeffroy 3(6):83.1874.—TYPE:SAMOA.Tutuila, sine probably destroyed). 511b 10: 26.1935.—TYPE: THAILAND.Puket, Krabi, Joseph, TYPE: TRINIDADandTOBAGO. Trinidad, nearSt. flavescens Paris, IndexBryol.1262.1898. B?, probably destroyed). TYPE: CUBA.LomaSanJuan, Bull. Mens.Soc.Linn.Bot.Lyon 34:326.1965.— Vernieri ) ). T. lindbergii inSamoais (holotypeB?,probably destroyed). T. planum . (holotypeBM?). Taxithelium planum ), wichisacommonandwidespread species. knownfrom Cuba. considerd itcloseto speciesnowincludedinsub- T. planum Crüger 14 Müll.Hal.,Syn.Musc.Frond. 2:265.1851.— . The mostcommonrepresentative ofthatsub- E.Ule 1598 (seecommentsabovefor isalsoavariablespeciesandtherefore many T. vernieri (Brid.)Mitt.var. ), allnowbelongingtosubgenus (Brid.)Mitt.var. Hypnum plumularia T. vernieri. , whichistheonlyspeciesof Trichosteleum orthothecium Hypnum rhizophoreti (Müll.Hal.)Renauld&Cardot, Rev. . (holotypeB?,probably destroyed). (Müll.Hal.)Broth., Nat.Pflanzenfam. (Müll.Hal.)Broth., Nat.Pflanzenfam. (A.Jaeger)Broth., Nat.Pflanzenfam. T. natans . (holotypeB?,probably destroyed). Dixon,J.SiamSoc.,Nat.Hist.Suppl. istheonlyspeciesofsubgenus . Sigmatella oophylla

T. vernieri . Hypnum rhizophoreti Hypnum planum hookerioides Graeffe s.n.( Hioram 11808 T. natans flavescens Müll.Hal.,Syn. Musc. T. planum Taxithelium Müll.Hal.,J.Mus. , T. isocladum T . . planum A. Jaeger, Ber. Bizot&Thér., and Müll.Hal., (Müll.Hal.) hltp B?, holotype (subgenus (subgenus Taxithelium . (holotype . Taxithelium subgenus Brid.var. T. oophyl- Vernieri var. and into Blume Kerr fla- T. T. . .

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 Bohrs V F. 10 rae I In Bryales . mounting 1909. permanent F rapid . V. a Brotherus, as solution Hoyer’s 1954. E. L. Anderson, anonymous reviewers. improved bythesuggestions ofLenaStruwe, Alan Whittemore, andtwo of Missouri,St.Louisallprovided funding.Themanuscriptwasgreatly Botanical Garden, Withney HarrisWorld EcologyCenterandUniversity tors ofalltheherbariacited.CAPES,Braziliangovernment,Missouri Mohamed, HajiRoyceLongton,JoseHidalgo,andthecura- Benito Tan, Yong KienThai,MonicaSuleiman, Ahmad Damanhuri Stevens, Toby Kellogg,BillBuck,Bruce Allen, StevenChurchill, SiHe, [Volume 36 SYSTEMATIC BOTANY considered ittobeclose cells. Brotherus, whentransferringthisnameto protologue, Taxithelium subretusum my visittoBM. known from Thailand.Iwasunabletolocatethetypeduring being smooth,whichisnottrue ofanyspecies 20 Bohrs V F. 12 uc (abos) 2 Hlt I In Hälfte. 2. (Laubmoose). Musci 1925. F . V. Brotherus, Taxithelium, now in but Brotherus, whenhetransferred Taxithelium ventrifolium agree withthesomewhat vague originaldescription. species belongtosubgenus sidered itcloseto Taxithelium tongense visit toNY. Taxithelium annandii tum Mem New York Bot.Gard. 45:521.1987. 478. 1915.= Physique, Naturelle etPolitiquedeMadagascar, Mousses39: this isthebasisofillegitimatename. the protologue information itisimpossibletoknowwhether Dr. Annand in1903,makingitatypecandidate,butwithout anywhere elseintheliterature. journal (Proc. Linn.Soc. New SouthWales 40:152.1915.)or TROPICOS, butIwasunabletofindthenameincited Acknowledgments. This speciesisprobably a In theoriginaldescriptionleaf cellsare describedas The originaldescriptiondoesnotmention thepapillae, Probably aspeciesinsubgenus Taxithelium annandii Ecue Taxa— Excluded There isonespecimennamedassuchinS,wascollectedby , bothinsubgenus medium forbryophytes . Ege epi . Engelmann. W. Leipzig: Engler . Pflanzenfamilien Ed SRI-LANKA. Thwaites &Mitt.,J.Linn.Soc.,Bot.13:321.1873.—TYPE: Pflanzenfam. I(3):1093.1908. (holotype B?,probably destroyed). Godeffroy 3(6):84.1874.—TYPE:FIJI.Ovalau,sinelegit. I(3): 1090.1908. Ed.Graeffes.n. Godeffroy 3(6):83.1874.—TYPE:TONGA. Tongatabú, I(3): 1090.1908. Phyllodon considered ittobeclose Ectropothecium subretusum Phyllodon truncatus . Iwasunabletolocatethetypeduringmy Thwaites s.n. T. planum Taxithelium latitruncatum . 2. d . nlr . Lizg: W Engelmann . W. Leipzig: Engler . A. ed. 2. . ( holotype B?,probably destroyed). Broth. &Watts, nom.nud. (Müll.Hal.)Broth., Nat.Pflanzenfam. Hypnum ventrifolium I express mygratitudetoBob Magill,Peter Hypnum tongense Taxithelium ieaue Cited Literature (Müll.Hal.)Broth., Nat.Pflanzenfam. iscitedin (Thwaites&Mitt.)Broth., Nat. The Bryologist T. glossoides Phyllodon Die NatürlichenPflanzenfamilien and Taxithelium (holotypeNY?). (Welw. &Duby)W. R.Buck. . Taxithelium T. instratum; Ectropothecium subretusum 7 242–244. 57: Hypnum ventrifolium . According tothevague T. planum Index Muscorum hasunipapilloseleaf and Müll.Hal.,J.Mus. , whichwouldalso Müll.Hal.,J.Mus. Cardot, Histoire T. ligulatum Bohrs con- Brotherus . bothofthese and Die Natürlichen Taxithelium, Taxithelium T. instra- . d A. ed. . , both , and into

Tn B C, . . asy ad . . coil 96. A otiuin to Chinese contribution A of 1996 . Schofield. revision B. W. preliminary and Ramsay , A P. H. 1999. C., B. Jia. Tan, Y. and C. B. Tan, Src, 87. 1867. R. Spruce, 1994. Eckel. M. P. and Crum , A. H. with J., A. Japan Sharp, of Sematophyllaceae family the of revision A 1969. T . Seki, Mte, 89. Msi uto mrcn Americani. Austro Musci 1869. W . Mitten, Tn B C ad . i 98. A ldsi suy f h fml Sematophyllaceae family the of study cladistic A 1998. Hong Jia. Y. of and collections C. post-war B. Tan, of revision A 1997 . But. Ph. P. and C. B. Tan, genre le sur Note 1901. Cardot . J. and F. Renault, genus The 2002. Tan . C. B. and Schofield, B. W. P., H. Ramsay, de ocorrências Novas 1997. Ilkiu-Borges . L. A. and L. C. R. the per- Lisboa, and of pattern, analysis branching phylogenetic form, Growth A 1996. 1999. C. Buck. Farge-England, La R. W. and L. Hedenäs, 2001. Salazar-Allen . N. and Churchill , P. S. R., S. Gradstein, (mosses): Bryophyta the of Systematics 2004. Buck. R. W. and B. Goffinet, Formose. l’íle de Mousses 1905. J. development Cardot, and Morphology 2010. Kellogg. A. E. and S. A. E. P. Câmara, Mgl, . 90. 1990. E. R. Magill, Hdns L. 19 caitc vlain f eainhp bten the between relationships of evaluation cladistic A 1996. L. Hedenäs, moss the of revision a Towards 1996. Longton. E. R. and A. Damanhuri, rela- Ordinal 2005. Goffinet . B. and Shaw J. A. Cox, J. C. R., W. Buck, Bc, . . B Gfie , n A J Sa 00. Nvl eainhp i pleu- in relationships Novel 2000. Shaw . J. A. and Goffinet , B. R., W. Buck, mosses. of classification and Morphology 2000. Goffinet . B. and R. W. Buck, 1998. R. W. Buck, 1987. R. W. Buck, of review A 1985. R. W. Buck, 1992. Powell. E. C. and K. R. Brummitt, Bc, . . n D H Vt 96. Sgetos o a e fmla classifica- familial new a for Suggestions 1986. Vitt . H. D. and R. W. Buck, Botany Australian Sematophyllaceae () . 1–70. Sematophyllaceae . oaia Garden. Botanical Memoirs oftheNewYork BotanicalGarden69 Science ofHiroshima University special reference toastatisticaldemarcation ofthefamily . 81–102. 27: Botany in China . Kong Sematophyllaceae . Newman. et Andinis,perAnnos1849–1860,legitRicardusSpruceus RevueBryologique Botany (Bryopsida, Sematophyllaceae)in Australia . Bryophyta (musgos)paraoEstadodoPará,Brasil . TheBryologist ichaetial positioninmosses:cladocarpyandpleurocarpy redefined . Sematophyllaceae . 49–82. 21: Garden . Botanical Missouri Louis: St. 98 . MonographsinSystematicBotanyfrom theMissouriBotanicalGarden systematics ofbryophytes from moleculestoarevised classification . Pp. 205 – 239 in of leafpapillaeinSematophyllaceae . Botanical Garden sary forbryology. Monographs inSystematicBotanyfrom theMissouri Hookeriales, theSematophyllaceaeandsomeothertaxa . 86 bryophytes oftropical America.MemoirsoftheNewYork BotanicalGarden Universidad AutónomadeMexico,SeriesBotanica genus 85–148. Hookeriales . tionships ofpleurocarpous mosses,withspecialemphasisonthe 774–789. 103: rocarpous mossesasrevealed bycpDNA sequences . Press . University Cambridge Cambridge: in 225–237 Pp. Press. the NewYork BotanicalGarden their names,includingabbreviations authors ofscientificnamesplants,withrecommended standardformsof tion ofpleurocarpous mosses . York BotanicalGarden82 ActaAmazonica . Nw ok: Nw ok oaia Garden. Botanical York New York : New . : 319–327. 9: 2 1–659. 12: 5 583–596. 15: Taxithelium The JournaloftheHattoriBotanicalLaboratory Catalogus Muscorumfere Omnium quosinTerris Amazonicus Pleurocarpous mossesoftheWest Indies.MemoirsoftheNew Systematics andBiodiversity Notes onAsianHypnaceaeandassociatedtaxa.Memoirsof 9 170–186. 99: 15( Glossarium polyglottumbryologiae:amultilingualglos- o. 3. S. oi isui oaia Garden . Botanical Missouri Louis: St. 33. vol. Bryophyte biology : 109–112 . 6: Smtpylca). (Sematophyllaceae). Suppl Lindbergia The JournaloftheHattoriBotanical Laboratory , d . ofnt V Hloel ad . Magill. R. and Hollowell, V. Goffinet , B. ed. , . Nw ok: Nw ok oaia Garden. Botanical York New York : New . . 43–53. .): Journal ofBryology Taxithelium . Nw ok: Nw ok oaia Garden Botanical York New York : New . 4 103–132. 24: 2 1–80. 12: Taxon . Kw: Ryl oai Gardens. Botanic Royal Kew: . , d . . hw n B Goffinet . B. and Shaw J. A. ed. , 5 21–60. 35: The Bryologist Authors ofplantnames,alist (Sematophyllaceae)inBrazil . Journal oftheLinneanSociety : 121–145. 2: Annales InstitutodeBiologia. Botanisches Centralblatt 9 787–798. 19: The mossfloraofMexico. . Nw ok: Nw York New York : New . Taxithelium 7 35–58. 67: Australian Systematic Australian Systematic 1: 22–33. 113: 4 49–55. 84: Acta Amazonica The Bryologist . Lno E. London: . Guide tothe R Spruce . R. , Taxithelium Lindbergia Molecular Journal of vol. 19: 86:

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Thu, 30 Sep 2021 01:21:04 01 CMR:AR-ICMCITO FTXTEIM 21 CAMARA: A RE-CIRCUMSCRIPTIONOFTAXITHELIUM Molecular 2001a. Deguchi. H. and Yamaguchi , T. Akiyama, H. H., Tsubota, described lichens and mosses of species the to Indices 1992. M. B. Thiers, 2011] oratory chloroplast phylogeny oftheSematophyllaceae(Hypnales,Musci)basedon Garden. Botanical York New York : by William Mitten . 0 221–240. 90: rbcL sqecs. sequences. Memoirs oftheNewYork BotanicalGarden68 The JournaloftheHattoriBotanicalLab- . New . Vt, . 94. Casfcto o te rosd P. 66–79 i in 696–759 Pp. Bryopsida. the of Classification 1984. H. D. Vitt, Tuoa H, . kym , . aauh , n H Dgci . 2001b. Deguchi. H. and Yamaguchi , T. Akiyama, H. H., Tsubota, Laboratory . ual ofbryology Hikobia (Sematophyllaceae, Musci)basedonchloroplast Molecular phylogenyofthegenus 3 529–549. 13: o. e. RM Shse ihnn: Htoi Botanical Hattori Nichinan: Schuster . R.M. ed. 2, vol. Trismegistia andrelated genera rbc L sequences. L New man-