Delivered by Publishing Technology to: The Natural History Museum, LIS, IP: 157.140.122.156 on: Tue, 17 Sep 2013 14:03:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. (8 otclclsad1mdlaycl Bndx15;Zhu 1953; (Benedix cell medullary of 1 comprising and stem cells of cortical section 5(–8) transverse lobule, and ventral dorsal underleaves, a a of and consisting leaves lobe folication, incubous its on based =repylu](rdti ta.2003). water al. running et in (Gradstein or rheophyllous] leaves ramicolous], [= living [= of twigs surface prefera- on the epiphyllous], on grow [= as such liverworts habitats, sized extreme in small bly al. usually et (Wilson These Lejeuneaceae 2007a). in lineage liverworts, derived of a genera representing species-rich most the among arguably is http://www.early-land-plants-today.org/Home), report: (ELPT O 10.1600/036364413X670304 DOI © Botany Systematic etdtehptei frmathmpayi h species- we the study, in present homoplasy the rampant genus In rich of 2012a). Dong hypothesis 2011; al. the al. et et tested Heinrichs Devos 2012; 2010; al. al. et et Feldberg 2009; 2007, al. Hentschel et 2007; Brown and Heslewood 2006; al. Heinrichs et 2005a; (Heinrichs preferences niche of com- conservatism morphological and plexity limited with in contrasted which in richness homoplasy lineages liverwort species derived frequent of prob- classification major the for in a lem as Evidence documented was 1997). 1997; morphology Graham gametophyte and al. and 2004) et (Barrett al. plants Tanaka et aquatic of (Ranker predomi- groups ferns the various as been grammitid lineages such epiphytic has in strategies, nantly especially homoplasy life plants, unique of land with of evidence associated groups that many times Thus, in surprising 2011). found al. numerous not et Wake is 1991; recovered Wake it (e.g. life been of tree the has across develop- constraints and/or conditions mental environmental similar to tation oyih 03b h mrcnSceyo ln Taxonomists Plant of Society American the by 2013 Copyright h urnl cetdgnrccnetof concept generic accepted currently The ools asdb ovreteouindet adap- to due evolution convergent by caused Homoplasy Cololejeunea and on ob itrt h ld opiigteremaining the comprising clade the to sister be to found aebe nlecdb rqethmpayo opooia hrcesadd o cuaeyrfetseisrltosis new A relationships. continents and species clades reflect major where accurately events expansion. vicariance not range ancient species do for intercontinental evidence of recent and provide for classifications not characters recovered current was did The morphological evidence data taxa. some Our natural of instead outlined. as associated, homoplasy is supported are genus not frequent this were for accessions by classification multiple influenced with species been 30 have of out Six polyphyletic. or egahcrnea ela h opooia iest fti rdmnnl ppylu eu.Temlclrdt upr he major three support data molecular The derived genus. the epiphyllous of predominantly phylogeny this molecular of diversity comprehensive morphological first the the as lineages: present well we as Here range mosses. geographic and liverworts genus as liverwort such diversity, ecological and/or Keywords— Abstract— 5 7 lrctvnHle-ntttfu Albrecht-von-Haller-Institut tt e aoaoyo ytmtcadEouinr oay nttt fBtn,CieeAaeyo Sciences, of Academy Chinese Botany, of Institute Botany, Evolutionary and Systematic of Laboratory Key State Chondriolejeunea 8 uhr o orsodne(u-in h,[email protected];Hrl cnie,[email protected]) Schneider, Harald [email protected]; Zhu, (Rui-Liang correspondence for Authors 6 ytmtcBtn n yooy aut fBooy nvriyo uih(M) ezne tas 67, Strasse Menzinger (LMU), Munich of University Biology, of Faculty Mycology, and Botany Systematic Cololejeunea 21) 83:p.553–563 pp. 38(3): (2013), Colura ihmr hn40pbihdbinominals published 400 than more with , vdnefrRmatHmpayi h hlgn fteEpiphyllous the of Phylogeny the in Homoplasy Rampant for Evidence apn ools a eamjrcalnei h lsiiaino adpat hthv iie opooia differences morphological limited have that plants land of classification the in challenge major a be can homoplasy Rampant Aphanolejeunea , Myriocoleopsis Cololejeunea igYu, Ying eebt eovda uaiemnpyei rus n etdi the in nested and groups, monophyletic putative as resolved both were Src)Schiffn. (Spruce) 3 oayDprmn fEszterha of Department Botany Lejeunea igorpy hools eunedata, sequence chloroplast biogeography, , n t ls eaie.W sdtremres( markers three used We relatives. close its and ,and 1,2 iewr Genus Liverwort 2 oayDprmn,NtrlHsoyMsu,Lno,U K. U. London, Museum, History Natural Department, Botany 4 iteeLte 1 83 Herdwangen-Scho 88634 11, Letten Mittlere Tama Cololejeunea 1 atCiaNra nvriy hnhi204,China. 200241, Shanghai University, Normal China East tp rnhn,lc of lack branching, -type ¨ u-in Zhu, Rui-Liang faznisncatn er-uutUniversita Georg-August Pflanzenwissenschaften, r ´ sPo omnctn dtr ao neWilson Anne Carol Editor: Communicating . ´ Myriocoleopsis cs, Cololejeunea 3 losScha Alfons 03 uih Germany. Munich, 80638 Cololejeunea ejn 003 China. 100093, Beijing Cololejeunea 1,8 pce eersle nacaewith clade a in resolved were species ´ yClee gr f 3 -31 Hungary. H-3301, 43, Pf. Eger, College, zy n aadSchneider Harald and is 553 pce.Cretyacpe ugnr of subgenera accepted Currently species. ¨ fer-Verwimp, Chondriolejeunea rae ssprt eeabas genera e.g. tiveness, separate as treated hlodseiegmtpyein gametophyte sterile thalloid oeua hlgntcsuisrcvrdeiec osup- to of evidence inclusion recovered port studies with phylogenetic their of molecular similarity context in morphological rejected was shared genera these of status generic in underleaves bu h nlso fsvrlmrhlgcldelimitated morphological several of within inclusion groups Po the 2001; raised So been about have and arguments controversial Zhu Some 2012). 1961; Piippo Mizutani 1953; (Benedix Po ehpteie hthmpayotnocr ntemor- the in occurs of often characters homoplasy 1991). phological that (Wake hypothesized relationships the species We to of related problems interpretation for factors current main the of one arguably is relation- natural alone. of morphology interpretation on at we based the be ships by Thus, may puta- caused classification of partly studies. infrageneric status least the the these and about segregates debates in tive ongoing included the that were assumed though even 2007a), samples al. et few Wilson 2006; al. et Gradstein 2005b; of concept wide and eei xMizutani, ex Benedix subgenera: eight Schiffner, recognized genus, studies this Aphanolejeunea for recent proposed most been although have subgenera more twenty Historically, 2003). than Srivastava and Asthana 2001; So and ´ h bnac fhmpayi opooia characters morphological in homoplasy of abundance The cs, Chondriolejeunea trnL–F Chlorolejeunea (Lejeuneaceae) , ¨ Taeniolejeunea rbcL ah Germany. nach, oeua phylogeny, molecular , Aphanolejeunea Cololejeunea 4 n rT)ad16acsin ersnigthe representing accessions 116 and nrITS) and , ohnHeinrichs, Jochen Chondriolejeunea AEas Benedix, (A.Evans) Cololejeunea Cololejeunea Aphanolejeunea 2,7,8 Bndx i Po & Kis (Benedix) eei,and Benedix, Cololejunea ooeenaminutissima Cololejeunea ¨ ,303Go 37073 t, Cololejeunea uhas such Zik)Benedix, (Zwick.) clade. tp rnhsin branches -type Aoe 04 enih tal. et Heinrichs 2004; (Ahonen Cololejeunea Metzgeriopsis do opooia distinc- morphological on ed oee,teindependent the However, . Myriocoleopsis , ooeenaangustiflora Cololejeunea paoeena Metzgeriopsis Aphanolejeunea, Metzgeriopsis and ¨ tne,Germany. ttingen, 5,6 ´ ie h ml ieof size small the given Cryptolejeunea s hs aahdbeen had taxa These cs. Metzgeriopsis Leptocolea Cololejeunea eeparaphyletic were . . n rsneof presence and , Aphanolejeunea Aphanolejeunea Pedinolejeunea Cololejeunea K .Goebel) L. (K. (Spruce) ihna within Recent . Benedix was ´ sand cs , , Delivered by Publishing Technology to: The Natural History Museum, LIS, IP: 157.140.122.156 on: Tue, 17 Sep 2013 14:03:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 5 YTMTCBTN Vlm 38 [Volume BOTANY SYSTEMATIC 554 cife swl stepttv itrgenus sister putative the as well In as genus. Schiffner the e.g. segre- of putative gates, all range of representatives geographic the sampled we the addition, also informative but taxonomic characters, putatively of morpholog- especially the variation, only not ical represent to designed was sampling of ufc ftesm ef hs atclrcr a ae oseparate to taken analyses. DNA was after and care before spec- particular of checked Identification was the Thus, specimens. imens on from co-occurred leaf. extraction DNA taxa same for several gametophytes often the that of fact small the surface rather and the were specimens project of this size to of challenges species Major Four species. (2007b). outgroup as al. et Wilson by work ools nteeouino opooia characters morphological rampant of for evolution evidence Lejeuneaceae. derived the 2) among subgenera; in and homoplasy seg- as genera such pre- respectively, regated of units paraphyly taxonomic or proposed monophyly viously 1) address questions: to following employed the was phylogeny recovered The Dumort. 02,ti td a eindt eosrc h phylogeny ( the reconstruct al. et to Dong designed of 2009a, 2011; was al. al. study et et Sukkharak this 2011; Heinrichs 2012), al. 2006; et 2003; al. Renner c; et al. al. 2012b, et Hartmann previous Wilson et b; 2004; of al. 2007a, et (Ahonen 2004, light Groth-Malonek 2003; Lejeuneaceae In al. et 2009). on Gradstein al. studies et molecular (e.g. Schneider only 2002; morphology on Endress based which efforts in lineages misleads of homoplasy classification natural been the has reveal of data to sequence evolution shown DNA parallel habitats. of extreme in frequency phenotype recovered and group this osrie yteaalblt fmtra utbefrDAsuis One studies. DNA for suitable material of species of availability the by constrained species of sampling Ingroup The of study. classifi- distribution proposed geographic this variation, and morphological cations, for the represent species to selected 80 were representing accessions 128 a-N-oyeae(iln,Lno,U .,2.5 1.5 K.), 25 buffer, U. of reaction total London, a (Bioline, in Taq-DNA-polymerase performed was PCR The 2006), al. amplified et were primers: region, following nrITS the ribosomal using the separately genome, nuclear the chloro- of the region of Hilden, regions (Qiagen, the Two kits genome, specimens. few plast mini a plant for employed DNeasy was while protocol Germany) CTAB 1987), of modified DNA Doyle a The and using applied. extracted (Doyle were was methods specimens of extraction majority Two the yr. collected 20 were last that the specimens within herbarium or gametophytes dried gel silica ape ihoeo w ein eeicue nteiiilanalyses, initial introduced the ambiguity avoid in to datasets. analyses included incomplete Additional final by were the 1. from regions Appendix excluded two were in but or listed one are with numbers details samples Voucher accession study. this GenBank in investigated and were accessions markers 128 three the All for protocols. obtained biochem- suggested BigDye manufacturer’s the the analyzer using and DNA istry California) 3730xl City, ABI Foster an Biosystems, on (Applied generated were sequences Bidirectional :0mn lnaina 72 at elongation mins 1:30 0cce f10 i eauaina 94 at denaturation min 1:00 of cycles 30 Boie,1 (Bioline), eubodtbodthm)adMcld . aalbefo http:// from http://www.mbio.ncsu (available from 4.0 manually (available MacClade and 7.1.3.0 Michigan), and Arbor, Bioedit .edu/bioedit/bioedit.html) Ann using (GeneCodes, aligned 4.8 Sequencher in olwn rga:40 isiiildntrto t94 at denaturation initial mins 4:00 program: following (10–25 DNA plate trnL–F ao apigadOtru aaSelection— Taxa Outgroup and Sampling Taxon N xrcinadPCR— and Extraction DNA hlgntcAnalysis— Phylogenetic Myriocoleopsis Cololejeunea , Drepanolejeunea rbcL trnL–F m paoeena Chondriolejeunea Aphanolejeunea, aho 10 of each l n n ula eoergo nIS.Our (nrITS). region genome nuclear one and ) Cololejeunea eeicue sigopseisbsdo previous on based species ingroup as included were rbcL Tbre ta.19) n rT Hrmn ta.2006). al. et (Hartmann nrITS and 1991), al. et (Taberlet m yuigtoclrpatgnm regions genome chloroplast two using by m ) h C mlfcto a are u sn the using out carried was amplification PCR The g). aeil n Methods and Materials gl 5 M Boie,0.5 (Bioline), mM) (50 MgCl2 l eeadtenon-coding the and gene ,twoof m  ;adafnletninse t72 at step extension final a and C; owr n ees rmr n 1 and primer, reverse and forward m l eunedt eeeie n assembled and edited were data sequence All 16acsin ersnig6 pce)was species) 68 representing accessions (116 eoi N a xrce ihrfrom either extracted was DNA Genomic Diplasiolejeunea Cololejeunea rbcL  m ,5 e neln t50–56 at annealing sec 50 C, ecin otiigoeunit one containing reaction, l Wlo ta.20;Gradstein 2004; al. et (Wilson Lejeunea he of three , trnL–F ,and esmldattlof total a sampled We n t ls relatives. close its and Colura m m a Polymerase Taq l NP 1 mM) (10 dNTPs l i.wr selected were Lib.  Myriocoleopsis ein n one and region, Colura ;floe by followed C;  o mins. 7 for C (Dumort.) n two and , m tem- l  C, 2 eeprioyifraie h Paayi eutdin resulted analysis MP The informative. parsimony were 922 ncnieaino h hlgntcrslsa ela icsino these 2003). of al. discussion et as Gradstein (e.g. well studies as previous in results characters phylogenetic the of consideration evaluated critically in were assessments character homology each preliminary for The (RI) 1). calcu- index (Table retention by and inferred (HI) was index characters homoplasy eight the lating these character of each Homoplasy of trees. states 100 ancestral reconstructing over by uncertainty Phylogenetic account states. into character taken of states was order multi-character the or without binary or with either articu- and schemes: were scoring that character assessments in homology preliminary lated obtain to Po used 2003; Srivastava Zhu were 1985; and 1979; Asthana Tixier 2001; consider- 1961; So Mizutani and and 1953; specimens (Benedix herbarium literature of the of study ation careful the characters by size, these associ- obtained about stem Information cells liverworts). was margin, secretory in cells thin-walled, leaf initial free (tiny, leaf with papillae of ated hyaline cells sigmoid of margin, position hya- leaf mammillae), and liverwort free and the (papillae of cells in cells lobe cells line leaf walled of thick prostruction often dorsal elongate, leaf), of longitudi- composed (a stripe vitta type, nal branching underleaves, are: completeness. characters and eight classifications These in usage previous on characters based morphological selected (available were The 2.74 1). Mesquite Table in http://mesquiteproject.org; implemented from morphological as using eight hypotheses reconstruction phylogenetic addition, character obtained MP In paraphyly the onto 1998). and plotted Holms monophyly were characters and of (Page criteria applied the 1953; were particular, (Benedix In classifications 1961). infrageneric Mizutani and inter- published evaluate ape ihntebr-npae Pi ie sP-Pfrtenon- the for PP-UP as given analyses. is partitioned the PP for trees PP-PA phase. the and analyses discarding burn-in partitioned after the trees sampled within all sampled of tree consensus majority http://beast.bio burn-in the from for of (available calculated were v1.4.1 estimation (PP) probabilities posterior Tracer Bayesian and .ed.ac.uk/Tracer.). using runs checked every of were sampling phase convergence with generations The million generation. ten 1,000th simul- for four tree and chains rates, Markov with unlinked tree, taneous random simultaneously a with inferred starting model runs GTR searcher, values the with parameter out carried implemented, was search Bayesian Each (nrITS). region n ihasnl oe o h obndainetadascn one second ( a regions and alignment chloroplast combined into the partitioning for a model performed: with were single BI a of kinds with Two one 2001). Ronquist and (Huelsenbeck 3.1.2 parameters and search. model tree optimal same the the in using as PhyML in via obtained replicates were bootstrap analyses 200 invariable into ML for + (ML-BS) the plugin values with gamma Bootstrap simultaneously searches. inferred + tree a values parameter (GTR with parameters via but selected of sites) implemented number sub- and as the model with http://www.geneious.com) stitution 2003) from (available Gascuel 5.5.6 Geneious and in ratio implemented (Guindon PhyML likelihood as in 3.0 performed (AIC) hierarchical were (ML) Criterion analyses the likelihood Information Maximum Modeltest. Akaike using and selected (hLRT) model and test were substitution (Posada DNA parameters appropriate 3.7 The Modeltest and 4.0. for PAUP* interface and 1998) cross-platform http://www.genedrift.org/mtgui Crandall a from is which (available .php) 1.01 MrMtgui using RAS. ten with each swapping, branch gener- TBR replicates were addition if and stepwise (MP-BS) simple values tree 1,000 Bootstrap with consensus searches found. heuristic using was strict ated as tree a treated MP as one were characters than summarized Gaps more All were unordered. step. Trees and each characters. weighted at missing equally held as trees treated ten were with swapping, bisection-reconnection branch tree (TBR) (RAS), 1,000 mode, replicates search random-addition-sequence heuristic methods: following the using 2002) (Swofford recovered. investigations was phylogenetic heterogeneity topological for for employed evidence was no as accessions 128 of dataset chloroplast of analyses ( were parsimony regions that maximum trees independent consensus by bootstrap generated two comparing visually by explored analyses. subsequent in used alignments from excluded and visually identified were positions macclade.org.). Ambiguous neec fMrhlgclCharacters— Morphological of Inference fattl262DAbs ar,157wr osatand constant were 1,547 pairs, base DNA 2,682 total a Of aeinifrneo hlgn B)wspromduigMrBayes using performed was (BI) phylogeny of inference Bayesian selected were analyses phylogenetic based parameters for Models 4.0 PAUP* with conducted were (MP) analyses parsimony Maximum was regions marker three the among signals congruent for Evidence rLF rbcL trnL–F, n h ula ein(rT) ial,acombined a Finally, (nrITS). region nuclear the and ) Results h hlgn a sdto used was phylogeny The ´ s21) hs observations These 2012). cs trnL–F , rbcL n nuclear and ) Delivered by Publishing Technology to: The Natural History Museum, LIS, IP: 157.140.122.156 on: Tue, 17 Sep 2013 14:03:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. xiie ihlvl fhmpay(I=097 I=0.662; = RI 0.917, = (HI homoplasy each of cells levels hyaline by high bordered of margin exhibited leaf protrusions and dorsal cells, papillae, lobe char- hyaline leaf eight of these position across Among the 1). characters acters, (Table morphological phylogeny selected recovered eight the of were RI accessions and HI multiple with Twenty- 1). species (Fig. thirty monophyletic. D5 as and supported of D4, out D2, above D1, support four bootstrap for analyses, obtained (MP-BS ML was supported in In 95% strongly while 2). were 1), clades (Fig. D5 Fig. six and D4 95%; the D4, of of D3, exception D1, each the MP, BI, with In support good sub-clades D1–D6. received clades and Six analyses as phylogenetic labeled all D. in were recovered clade were D within in defined monophyletic as resolved o ul eovdi PadM nlss(i.1.Tetwo The 1). A (Fig. to analyses were ML sister D and clade, and MP segregates C, robust in B, a resolved clades fully formed not among D relationships and the C, although B, clades the groups, other osdrdt eo aooi infcnefrthe for significance (2001). taxonomic So of be to considered caeA,adteohroecnitn fteremaining the of consisting one other the of species comprising and one A), clades: of split (clade sister of first two The accessions in clades. D three resulted and clade C, MP CO B, in the A, the MY in as clades and defined smaller CO clade, several both CO MY recognized to comprising We sister clade 1). or the (Fig. analysis, analysis to ML sister in or CO 1.0), and = PP-PA 1.0, = UP ( CU was ML-BS with 1). where ana- together clade (MY) (Fig. BI CO 95% and the than ML, of less MP, exception in the supported with lyses well were the clades as analyses These defined phylogenetic all clades, (CU), clade, three this identified Within consistently 1.0). = PP-PA Myriocoleopsis it 0 akn,()=peet0670.714 0.757 1.0 0.00 0.808 0.667 0.900 0.00 0.750 0.837 BI and 2). ML, 1, MP, (Figs. The both topologies 2). similar (Fig. largely for shared shown topologies analyses is them identical of one and Only model trees. 32,685.455 partitioned cells, = cortical -InL the and 5–8 trees un-partitioned (0) for 9,001 the of for on 33,070.732 likelihood based = mean analyses a -InL with BI present hypotheses = 1). (1) phylogenetic (Fig. lacking, tooth, obtained (0) analysis apical found the was ML steps 36,701.166) of = peak 5,693 the (-InL at tree of in or likely distal length most (0) single a A 1). of (Fig. trees parsimonious equally 1,212 present = (1) lacking, (0) present present = = (1) (1) (0) lacking, lacking, (0) present (0) = (1) lacking, (0) papillae hyaline 555 of Position size margin Stem leaf free of cells margin Sigmoid leaf free of cells Hyaline prostructions Dorsal Vitta type Branching (COLOLEJEUNEA) LIVERWORTS EPIPHYLLOUS IN Underleaves HOMOPLASY AL: ET YU 2013] h eeso ools eedtce ycluaigthe calculating by detected were homoplasy of levels The Cololejeunea Table Colura Myriocoleopsis .Caatrsae,hmpay n eeto ne fec hrce acltdars h eoee retplg.Teecaatr were characters These topology. tree recovered the across calculated character each of index retention and homoplasy, states, Character 1. hrce hrce ttsHmpayidxRtninindex Retention index Homoplasy states Character Character Cololejeunea ld a ihrsse oM nB nlss(PP- analysis BI in MY to sister either was clade ) Chondriolejeunea M-S=9% LB 0% PU 1.0, = PP-UP 100%, = ML-BS 99%, = (MP-BS a eovdi ld oehrwt four with together clade a in resolved was Aphanolejeunea ooeue minutissima Cololejunea ooeenaangustiflora Cololejeunea M) and (MY), cae –) nM n Ianalyses, BI and ML In B–D). (clades Myriocoleopsis and , Chondriolejeunea Aphanolejeunea Cololejeunea a etdi clade a in nested was Lejeunea etlo ipae,()poia faia tooth apical of proximal (2) lobule displace), the or of (ental surface interior the on (1) cells cortical 9–20 (1) rpartly or Cololejeunea S. cif.The Schiffn. (Sm.) Seh)Mizut. (Steph.) , C)clades. (CO) tp,()exclusively (1) -type, Colura Aphanolejeunea eeeach were lsiiainacrigt eei 15) iuai(91,Txe 18) n h and Zhu and (1985), Tixier (1961), Mizutani (1953), Benedix to according classification Colura and , ³ -type n.Gvntetedo rqethmpay utpeorigins multiple the homoplasy, frequent of and of trend group the this Given ing. of of divergence evolution genetic developmental the the that into certain insight not e.g. more studies, are to future we assigned in taxa 1), critical several (Fig. of clade Incorporation clade exclusive an CO in Aphanolejeunea the resolved in were study nested this para- in a of sampled introduction the avoid to rejected phyletic is level recognition genus of the hypothesis at alternative our the and in with 2009) as consistent Bernecker 2005b; well are of as studies al. inclusion 2007b) phylogenetic the al. et All et study. Heinrichs Wilson present 2006; 2004; al. et (Ahonen Gradstein studies logenetic of Aphanolejeunea imdaraywt h cainlocrec fti yeof type this of occurrence occasional the with already firmed shn n rvsaa20;Po 1979; 2003; Tixier Srivastava 1953; (Benedix and genus the Asthana of recognition for sup- port insufficient provided forms transitional of occurrence the I=074 I=070 I=020 respectively). 0.200, bor- = RI 0.667, margin 0.750, = = leaf respectively) (HI HI and 0.792, values 0.714; = moderate = vitta RI showed RI of cells 0.837, sigmoid status by = the dered HI whereas 0.771; 1), = (Table RI 0.900, = HI of Aphanolejeunea 98 hes18;ZuadS 01.Hwvr aito of variation However, to 2001). assigned species So across and characters these Zhu 1982; Thiers 1938; fClljuei iewrssc sterjcino the of rejection the as such of status generic liverworts Cololejeuneoid classification of subgeneric and generic the concerning evidence Genera— Segregate rnhn n iopi evsta eeue ysome from by taxon used the were distinguish that to its leaves authors dimorphic of and interpretation branching the Po on and of focused status Kis the about 1982; arguments Thiers Conflicting 2001). 1911; (Evans characters mor- of interpretation phological the as on based genera authors independent some by as suggested treatment their with inconsistent is within nested both are mlctosfrGnsCasfcto n Previously and Classification Genus for Implications Aphanolejeunea Cololejeunea Aphanolejeunea Cololejeunea tp rnhn loocr nsvrlspecies several in occurs also branching -type (Po smnpyei eas flmtdsampling. limited of because monophyletic is Aphanolejeunea Aphanolejeunea htwr o osdrdt eogto belong to considered not were that ´ aebe nlddi eetmlclrphy- molecular recent in included been have sadPip 02.Svrlrepresentatives Several 2012). Piippo and cs lhuhalacsin of accessions all Although . h eut fti td rvd new provide study this of results The tp rnhn r xetd scon- as expected, are branching -type Cololejeunea .diaphana A. Discussion .1 0.677 0.00 0.913 0.500 .0 0.875 0.500 within and Aphanolejeunea h eoee phylogeny recovered The . ´ .Eas ih provide might Evans, A. sadBrekr2009). Bernecker and cs Chondriolejeunea Cololejeunea Aphanolejeunea Cololejeunea Aphanolejeunea Aphanolejeunea Aphanolejeunea Aphanolejeunea tp branch- -type (Po sthese as , ´ (Evans sand cs -type and ´ cs Delivered by Publishing Technology to: The Natural History Museum, LIS, IP: 157.140.122.156 on: Tue, 17 Sep 2013 14:03:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 5 YTMTCBTN Vlm 38 [Volume BOTANY SYSTEMATIC 556 nlss eo rnhs=bosrpvle bandb aiu ieiodaaye.Clmst ih fo ett ih) ldsrcgie withi recognized clades right): to = LE left follows: (from of as classification right abbreviated the to in Columns as analyses. assignments likelihood subgenera maximum by obtained values Cololejeunea bootstrap = branches below analyses; values eflb el;5 yln el ffe efmri;6 imi el ffe efmri;7 tmsz;8 oto fhaieppla.Caatrstat Character papillae. hyaline of postion 8: size; stem 7: margin; leaf free nodes, of on dots cells prostruc black dorsal sigmoid with 4: 6: clades vita; two margin; 3: for branching-type; leaf assignments 2: free Subgeneric underleaves; of 1. 1: Table cells are: in Characters hyaline shown (1961). 5: Mizutani cells; and (1953) lobe Benedix leaf to according assigned were taxon Fig. ³ .Src osnu f122eulyprioiu re bandb aiu asmn nlsso h obnddtst nybootstrap Only dataset. combined the of analysis parsimony maximum a by obtained trees parsimonious equally 1,212 of consensus Strict 1. 5 * 0% 59% r lte ntesrc osnu re bv rnhs=bosrpvle bandb aiu parsimony maximum by obtained values bootstrap = branches above tree: consensus strict the on plotted are 95–99%) = * 100%, = (** 95% brvain sgvni h et hrces18a ecie eo n ntetx,oe qae ,baksurs=1 unknown; = ? 1, = squares black 0, = squares open text, the in and below described as 1–8 characters text; the in given as abbreviations , Lejeunea ,DR= Drepanoeljeunea Cololejeunea ,DI= nmsgvni etbxo h oe etcre ftefgr) urn eu classification genus current figure); the of corner left lower the on box text in given (names Diplasiolejeunea ,CU= Pedinolejeunea Colura ,MY= and Myriocoleopsis Chondriolejeunea ,CO= r niae nphylogeny. in indicated are Cololejuenea ugnr feach of Subgenera . inof tion sare es n Delivered by Publishing Technology to: The Natural History Museum, LIS, IP: 157.140.122.156 on: Tue, 17 Sep 2013 14:03:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 03 UE L OOLS NEIHLOSLVROT CLLJUE)557 (COLOLEJEUNEA) LIVERWORTS EPIPHYLLOUS IN HOMOPLASY AL: ET YU 2013] rN.Tecnesste sbsdo h re eoee n90000gnrtosta eeasmldfo h eut fMBysaaye with analyses MrBayes of results the from assembled were that (PS) generations values Support 9,000,000 Posterior in generations. recovered 1,000,000 as trees determined the phase on burn-in based the of is exclusion tree consensus The nrDNA. Sgnrtdwt eaae oesfrcDAadnDApriin r ie bv rnhsadsnl oe o l ein eo branches. below regions all for model single and branches above given are partitions nrDNA and cpDNA for = models LE Abbreviations: separated with generated PS Fig. .Bysa aoiycnesste acltdfo h eut faBysa nlsso h obnddtstwt atto fcDAand cpDNA of partition a with dataset combined the of analyses Bayesian a of results the from calculated tree consensus majority Bayesian 2. Lejeunea ,DR= Drepanoeljeunea ,DI= Diplasiolejeunea ,CU= Colura ,MY= Myriocoleopsis ³ .5aeidctdb *=10 n y*=0.95–0.99. = * by and 1.00 = ** by indicated are 0.95 ,CO= Cololejuenea . Delivered by Publishing Technology to: The Natural History Museum, LIS, IP: 157.140.122.156 on: Tue, 17 Sep 2013 14:03:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 5 YTMTCBTN Vlm 38 [Volume BOTANY SYSTEMATIC 558 n nlso of inclusion and of exclusion the following phylogeny molecular nrdcdb tpaii 94(e eei 93 Mizutani the 1953; Benedix for was (see Po it 1914 responsible 1966; in since Stephani likely taxon by and this was introduced of (Zhu changes variability morphology nomenclatural variable This numerous its 2002). of So because challenge a The 1995). Gradstein and of Reiner-Drehwald affinity 1975; Vital and from tinct taxa. related distinctive- of morphological morphology the on of consideration based without ness groups defining of lem angustiflora Fg.1 ) icmcito of Circumscription 2). 1, (Figs. angustiflora C. its unit to biological the adaptation minutissima of C. an recognition new considered A habitat. character epiphytic a growth, creep- and ing two), (rarely innovation single a only with gynoecia with clade robust a in myriocarpa al. outside et clade Wilson robust species, a by formed study species systematic molecular which in (2007b), a on based with oil-bodies morphology in similarities sporophyte of context and in questioned was genus of Po status and the in confirm puigarrii to M. evidence Myriocoleopsis found Myriocoleopsis we gates, 2002; Endress (e.g. morphology 2009). al. create et on Schneider can based natu- structures relationships of unusual interpretation ral of new the feature concerning of arguments key invention conflicting a the or is sals underleaves of absence the Cololejeunea the of evolution if the mine that suggested in we stylus Thus, 1). (Fig. clade as data, Po 95.Hwvr hr r ifrne between differences and are there However, 1995). presence Gradstein the and (Reiner-Drehwald and perianths 5-keeled anatomy, inflated similarities, stem of form, morphological lobule and by lobe supported as such also are taxa these and subspecies typical the both (including another this of accessions study, In evidence. phylogenetic and similarities morphological as such Po species, & (Herzog)Bernecker several in branching oee,tesau of spikes. androecial status long the and gynoecia gametophyte, cymes, However, the compound in in axes arranged leafy erect of occurrence rnfre to transferred treat- this rather However, styli. for underleaves called ment appendages to lobule homologous to than as appendages phytic ´ Recently, h eei ocp of concept generic The ncnrs otetopeiul icse uaiesegre- putative discussed previously two the to contrast In s20) hc rvddagmnsitrrtn gameto- interpreting arguments provided which 2001), cs Myricoleopsis Myriocoleopsis ´ ´ s2001). cs Chondriolejeunea s19;ZuadS 01.Tepyoeei relation- phylogenetic The 2001). So and Zhu 1994; cs .vuquangensis C. Chondriolejeunea ,wr eovdotieo h Ocaeadinstead and clade CO the of outside resolved were ), Chondriolejeunea Cololejeunea Myriocoleopsis Seh)Mzt h bv aeilsrtsteprob- the illustrates case above The Mizut. (Steph.) si urnl yohszd vltoayrever- Evolutionary hypothesized. currently is as Chondriolejeunea cif.Crety he pce r recognized are species three Currently, Schiffn. sa needn eu.Terepii genus rheophilic The genus. independent an as srequired. is a rgnlyeetdb cife 14)for (1944) Schiffner by erected originally was Myriocoleopsis a o eovdi Oi h Lanalysis ML the in CO in resolved not was Myriocoleopsis Rie-rhadadGasen19;Kis 1997; Gradstein and (Reiner-Drehwald Aphanolejeunea Myriocoleopsis stefre a hre nreilspikes, androecial shorter has former the as ae na ntmclsuy(i and (Kis study anatomical an on based Myricoleopsis pce eense nthe in nested were species ´ cs, Cololejeunea and eursadtoa td odeter- to study additional requires Myriocoleopsis Po a o upre ymolecular by supported not was Cololejeunea ´ .koponenii C. s&Nn,wssubsequently was Ninh, & cs a ergtda eu dis- genus a as segregated was (Po Cololejeunea .angustiflora C. a salse ae nthe on based established was ´ oehrwt one with together and s21)bsdo observed on based 2010) cs Chondriolejeunea Fg ) h fiiisof affinities The 1). (Fig. a elrsle nour in resolved well was Cololejeunea species, ooeenaerostrata Cololejeunea (Po sa independent an as a is proposed first was Cololejeunea ´ s Po cs) a considered was .minutissima C. .minutissima C. .m. C. Myriocoleopsis ´ Cololejeunea Cololejeunea (Gradstein s and cs, though , subsp. .This C. of of phylogeny and variability taxon. morphological this of the segregation of The context 2002). So and angustiflora (Zhu perianths keeled Aphanolejeunea nifaeei lsiiaino hsgns iecurrently Five genus. of this subgenera of accepted classification infrageneric on n a hnewe ute pce r de othe to added are species further when change sampling may taxonomic com- limited cells our and of reflect (number may size stem) stem the posing homo- and of type level branching 1991; low in relatively Wake plasy The 1966; 2005). (Hennig or Fischer group and convergent given Kleunen natu- a of the of level resolve relationships to high ral evidence (see a little 0.917) provide = with evolution = (HI characters parallel (HI cells Such papillae sigmoid hyaline 3). margin of and/or lobe Fig. position free 0.837) the the = and 0.900), (HI 0.750), = cells (HI hyaline cells with lobe pro- dorsal leaf 0.667), of = of (HI trusions vitta subgenera homoplastic: as circumscribe recovered were to used commonly tts uha h curneo both of occurrence character the of as combination such deviant a the states, this displays in to present which corresponds is species that study characters present morphological of the variability in recovered species pce-ihgnr flvrot,sc as and 2009), such al. et liverworts, other (Hentschel of in reported genera been also species-rich have results characters Comparable morphological 1). each of (Fig. suite although a study, by characterized present well the was in polyphyletic or phyletic fspecies. of hpbetween ship eovda oohltcgop iha neti number uncertain an with groups monophyletic the as are resolved study this in serrulata included delinea- Examples species tions. robust obtaining and underlying processes of evolutionary exploration for diversity 2009b, genetic mor- and high phological al. exhibiting exhaus- complexes out et species carry on to Heinrichs investigations need tive the 2004; support results al. Our 2011). et 2010, Feldberg and (e.g. variation of plasticity morphological underestimation intra-specific overesti- by an from caused or mation as diversity cryptic 2011). interpreted by al. caused was numbers et species Renner result 2009b; later al. The et Heinrichs required (e.g. species of redefinition fraction on considerable based a but monophyletic data as molecular resolved were morphologically of species number recognized vast a liverworts: leafy on studies morphological lanciloba C. with species as such several variation including study, our monophyletic as in recovered not were accessions multiple with formalize to morphologies. their necessary determine is and studies, entities sampling taxonomic these taxon Future enhanced 1). (Fig. an D5 with for Mizut. ex Benedix two subgenera while conserved: Chondriolejeunea be new clades, can detected names Several such newly subgeneric old for for 2004). introduced reason al. be main should et the (Ranker as parallel identified phenomena or been convergent has from evolution resulting homoplasy Rampant u eut ovdsvrlise bu h eei concept generic the about issues several solved results Our vlto fMrhlgclFeatures— Morphological of Evolution mlcto o pce Classification— Species for Implication Cololejeunea Scapania ope n the and complex tp.Ti eea rn scnitn ihother with consistent is trend general This Steph. rmohrlnae nC sitiun nthe in intriguing is CO in lineages other from oee,te aea vnbge impact bigger even an have they However, . .angustiflora C. tp rnhn n bvt ocylindrical, to obovate and branching -type Dmr. uot Hircse l 2012a). al. et (Heinrichs Dumort. (Dumort.) eei o 1and D1 for Benedix ooeenaplanissima Cololejeunea Cololejeunea Radula .lanciloba C. n h remaining the and uot Dvse l 2011), al. et (Devos Dumort. C. eersle spara- as resolved were ope htbt were both that complex subgen. Cololejeunea Mt. by.and Abeyw. (Mitt.) i ftit species thirty of Six eea characters Several Lejeunea Frullania Pedinolejeunea Cololejeunea Cololejeunea Cololejeunea tp and -type subgen. Raddi C. Delivered by Publishing Technology to: The Natural History Museum, LIS, IP: 157.140.122.156 on: Tue, 17 Sep 2013 14:03:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. aias(rdti ta.20,21) odeapefor example good e.g. A madothecoides liverworts, rheophilic C. 2011). from derived 2003, is selection rheophilic al. ecological or et epiphyllous (Gradstein as adap- such habitats conditions the extreme as such to constraints tion ecological and/or rather plants these small in options developmental limited 0.827 of in = combination characters) (HI homoplasy all rampant for towards trend The dataset. 559 (COLOLEJEUNEA) LIVERWORTS EPIPHYLLOUS IN HOMOPLASY AL: ET YU 2013] yln el;B=dra rtuin oiino yln papillae. hyaline of ( position regions = chloroplast C two protrusion; and dorsal (nrITS) = nuclear B one cells; combined hyaline the of analysis parsimony maximum Fig. .Eape fcaatr xiiighg ee fhmpay(I>08)mpe notesrc osnu rersligfo h heuristic the from resulting tree consensus strict the onto mapped 0.85) > (HI homoplasy of level high exhibiting characters of Examples 3. Seh)Benedix, (Steph.) Cololejeunea .stotleriana C. a etersl fthe of result the be may rdt ta.as al. et Gradst. aeo h tm nsm iewrslnae.Nme and Number lineages. liverworts some sur- ventral in the stem) on the found of leaves of face row third (a underleaves (Gradstein 2011). habitats al. to et Gradstein adaptions 1975; Vital as and fertility and rhizomes, stems, creeping thick of consisting morphology unusual rather of segretgate ela hs of those as well ruby h otitrsigfauei h vlto of evolution the is feature interesting most the Arguably, Colura trnL Myriocoleopsis – Hircse l 02) htotnehbta exhibit often that 2012c), al. et (Heinrichs F and rbcL xldn E R n Icae.A= A clades. DI and DR, LE, excluding ) and Myriocolea pueaformer Spruce–a Delivered by Publishing Technology to: The Natural History Museum, LIS, IP: 157.140.122.156 on: Tue, 17 Sep 2013 14:03:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. dpaini xetdgvntesotlvdntr fthis of nature Such short-lived habitats. the growth. given these expected colonize strat- epiphyllous is the successfully of adaptation for to part be required may preference capacity egy dispersal high the a Adopting with coincide sampling. sparse or distributions not pattern dioecious distributional in found a that ranges, study intercontinental current possess the in monoecious observation several the hypothesis This with 2006). consistent al. propa- et is (Pohjamo asexual out ruled of be significance can’t dis- gules the long in although role dispersal, important which an tance 1908; in play 2009), (Yeates to al. considered et studies were Hock spores 2009a; previous al. et in Heinrichs 2008; discussed Frahm been sys- has breeding and 2009).tems ranges distribution al. wide et of Hentschel association The 2009; 2007, Vanderpoorten al. and et Feldberg Devos 2012a; 2010; 2011, liver- 2005a, leafy al. of et genera (Heinrichs other Po worts in reported and been have Zanten results rable Mun 1978; 1988; Gradstein (Zanten and Zanten vicariance 1981; ancient rather gemmae by or the spores than of 2007a), dispersal distance al. long et by some caused Wilson of 2007; ranges al. in intercontinental et occurred (Heinrichs liverworts evidence Tertiary epiphyllous current the of the events biogeo- Given divergent the lineage. that of the analysis exhaustive of an history for graphical allow not did work of sampling global The and 2010). Vanderpoorten Shaw 2001; of (Shaw follow- events combination evolution vicariance morphological ancient a ing and molecular of rela- of result These rates slow 2012). the and/or al. dispersal be long-distance et either supporting may mechanisms Dong ranges 2011; al. large al. et tively et Heinrichs Fuselier (e.g. b; species liverwort 2009a, in with- ranges or with disjunct distributions out wide of occurrence frequent the of the in origin leaf of transloca- 1969). (Crandall collar the like the ontogeny, of shoot of tion translo- stage early as the such in processes evolutionary a cation simple of the involve invention the may in However, structure 2006). events new al. et rare (Igic alternative plants are inven- of structures The history because new intriguing 2008). of also Miglietta tions is law re-evolution and Dollo’s of rejecting Collin hypothesis example further 1970; a add (Gould would intrigu- it is reversal as evolutionary ing, of structure, hypothesis new The a stylus. of gain the a or reversal, underleaves, evolutionary to an homologous of thus result the be could and question of in structure underleaf-like the of pretation of ancestor common the Myriocoleopsis of state ancestral the be comprising clade ( the to or as more of a sampling with investigation comprehensive further requires this but 2012c), al. et Colura lejeunea 6 YTMTCBTN Vlm 38 [Volume BOTANY SYSTEMATIC whereas underleaves, of absence underleaf the one by have Lejeuneaceae pair. in leaf per genera 2003). of al. majority et important The (Gradstein Lejeuneaceae highly in considered characters were taxonomic underleaves of presence 560 PadM;Fg ) ec,oeudrefprpi efcould leaf pair per underleaf one Hence, 1). Fig. ML; and MP h bec fcerydfndgorpia agsmay ranges geographical defined clearly of absence The Biogeography— Drepanolejeunea a eovda itrete to either sister as resolved was oss n nela e ef ntecretstudy, current the In leaf. per underleaf one possess Cololejeunea ,and and eetsuisspotdtehypothesis the supported studies Recent Colura Cololejeunea Cololejeunea Diplasiolejeunea and (including Colura Myriocoleopsis Cololejeunea pce sarsl frestricted of result a as species pce,e.g. species, Cololejeunea and Cololejeunea Myriocolea Myriocoleopsis utemr,teinter- the Furthermore, . ´ ze l 04.Compa- 2004). al. et oz Lejeunea Siphonolejeunea Colura and Chondriolejeunea r characterized are aaaelikely are taxa tp branches -type .angustiflora, C. e Heinrichs see ; ntepresent the in Myriocoleopsis and Cololejeunea B;Fg 2) Fig. (BI; Diplasio- swell as ´ cs is , art,S .H n .W rhm 97 dpierdaini h aquatic the in radiation Adaptive 1997. Graham. W. S. and H. C. S. Indian Barrett, 2003. Srivastava. C. S. and G. Asthana, the of phylogeny the Inferring 2003. Piippo. S. and Muona, J. I., Ahonen, oln .adM .Mgita 08 eesn pnoso ol’ Law. Dollo’s on opinions Reversing 2008. Miglietta. P. M. and R. Collin, order liverwort the of phylogeny Molecular 2004. I. Ahonen, Project 211 the University. and Normal 30825004) China East (no. National the China the for R-LZ of professor- and Foundation Sciences, visiting Science of Academy senior Natural Chinese the student the by HS Chinese granted 2010614081), ship the No.: by (Fellowship support council financial S. acknowledges and YY of Mu Malaysia), donations collections. their Lumpur, for F. (Kuala Thailand) per- (Songkhla, Yong and the Chantanaorrapint K.-T. and France), DNA, specimens (Paris, of extract Gradstein loan to the for mission (E) Edinburgh Garden Botanical eei,E .15.Indomalayische 1953. H. E. Benedix, rmr etn n/rpooea etn curn in Chondriolejeunea occurring neoteny The of protonemal 2003). gametophytes and/or al. et neoteny (Gradstein primary events heterochronic involving eo,N n .Vnepotn 09 ag ijntos speciation, disjunctions, Range 2009. leafy Vanderpoorten. in A. branches and of N. Devos, development and Morphology 1969. B. Crandall, ae htteuiu hrceitco the of characteristic unique the that cated selection a in result re-colonization. may fast towards distributions Both temporal 2009). al. and et spatial (Hock distribution island-like its and habitat oso h vlto fteelvrot seilyi the com- in and especially restricted highly liverworts environment. a these petitive in divergence of species evolution of context fac- the developmental to and on studies ecological tors future of allow impact the will reconstruct phylogeny recovered infra- the of species. variable circumscription Thus, morphologically hampered several and has entities genus homo-generic morphological this rampant consistent in the is of plasy that hypothesis likelihood conclusion This our 2011). the with al. increase et inherited (Wake to Such homoplasy hypothesized habitats. ephemeral are to limitations adaption of probably is result and of the space ontogeny morphological the limits which in group, constraint development this a be to presumed ol,J .adJ .Dye 97 ai N slto rcdr for procedure isolation DNA rapid A 1987. Doyle. L. J. and J. J. Doyle, eo,N,M .Rne,S .Gasen .Sa,adA Vanderpoorten. A. and Shaw, J. Gradstein, R. S. Renner, A. M. N., Devos, Cololejeunea og .S,A Scha A. S.-S., Dong, Acknowledgments. Perspectives— ln aiyPneeica:Isgt rmpyoeei analysis. in phylogenetic from 225–258 Insights Pp. Pontederiaceae: family plant study) molecular of appraisal first data. A (Jungermanniopsida): Lejeuneaceae Garden. cal Garden Botanical Missouri Magill. the E. R. and Hollowell, perspectives C. V. & problems progress, bryophytes: of tematics in 87–118 Pp. Jungermanniopsida). (Marchantiophyta, rnsi clg Evolution & Ecology in Trends pceu oau en vegetabilis regni novarum specierum Press. University Cambridge Cambridge: Sytsma. J. K. and Givnish J. T. n opooia rnfrainrtsi h iewr genus liverwort the in rates transformation Leptoscyphus morphological and Hepaticae. ml uniiso rs eftissue. leaf fresh of quantities small Evolution spe- and liverwort netics cryptic epiphyllous morphologically the and in cies endemics, narrow Tramps, 2012. .Po T. in divisions subgeneric traditional liverwort leafy challenge the data Molecular 2011. ´ s .R cmd,J ete,H cnie,adJ Heinrichs. J. and Schneider, H. Reitner, J. Schmidt, R. A. cs, h Bryologist The . rohtrmBibliotheca Bryophytorum ope stegetvreyo etncfeatures neotenic of variety great the is complex ehfezrNv Hedwigia Nova zur Beihefte . Evolution ¨ ,and e-ewm,P ence .Flbr,A Bombosch, A. Feldberg, K. Meinecke, P. fer-Verwimp, rvosmrhlgclsuishv indi- have studies morphological Previous Cololejeunea oeua vlto n dpieradiation adaptive and evolution Molecular Radula 5 582–594. 65: 0:297–308. 106: Metzgeriopsis eaegaeu otehraimo h Royal the of herbarium the to grateful are We ieaueCited Literature 3 779–792. 63: . Taxon 3 602–609. 23: .Vol.98.St.Louis:MissouriBotani- ao (including taxon 0 1–155. 60: 0 1623–1632. 60: oorpsi ytmtcbtn from botany systematic in Monographs Diplaziolejeunea ¨ 3:1–88. 134: Cololejeuneen lr(rse,Gray,S R. S. Germany), (Dresden, ller Gasene l 06 was 2006) al. et (Gradstein ) htceia Bulletin Phytochemical 0 1–261. 30: ooeena( taxonomic (A Cololejeunea . . edsRepertorium Feddes oeua Phyloge- Molecular Aphanolejeunea d.B Goffinet, B. eds. Tuyamaella oeua sys- Molecular Cololejeunea 9 11–15. 19: Porellales ,eds. - , Delivered by Publishing Technology to: The Natural History Museum, LIS, IP: 157.140.122.156 on: Tue, 17 Sep 2013 14:03:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. enih,J 06 oeua hlgn n igorpyof biogeography and phylogeny Molecular 2006. J. Heinrichs, 03 UE L OOLS NEIHLOSLVROT CLLJUE)561 (COLOLEJEUNEA) LIVERWORTS EPIPHYLLOUS IN HOMOPLASY AL: ET YU the in systematics angiosperm and Morphology 2002. K. P. Endress, 2013] atan .A,R isn .R rdti,H cnie,adJ. and Schneider, H. Gradstein, R. S. Wilson, R. A., F. Hartmann, algorithm accurate and fast, simple, A 2003. Gascuel. O. and S. Guindon, 2004. Gradstein. R. S. and Schneider, H. Heinrichs, J. M., Groth-Malonek, 2006. Heinrichs. J. and Ilkiu-Borges, A.-L. Wilson, R. R., S. Gradstein, phy- A 2003. Schneider. H. and Reiner-Drehwald, E. M. R., S. Gradstein, Habitat 2011. Vanderpoorten. A. and Ilkiu-Borges, A.-L. R., S. Gradstein, enih,J,A obsh .Flbr,H .Kee,J etce,J. Hentschel, J. Kreier, P. H. Feldberg, K. Bombosch, A. J., Heinrichs, vn,A 98 oe ntegenus the on Notes 1938. A. X. Evans, Rico. Puerto of Hepaticae 1911. A. Evans, rdti,S .adD .Vtl 95 On 1975. Vital. M. D. and R. status the S. and irreversibility and Gradstein, Law Dollo’s on Dollo 1970. J. S. Gould, Devos, N. Costa, P. D. Konrat, von M. Engel, J. J. Shaw, bryophytes B. of C., biogeography L. Fuselier, and dispersal Diversity, 2008. P. J. Frahm, enih,J,S og .Flbr,A Scha A. Feldberg, K. Dong, S. J., Heinrichs, edeg . .Va J. K., Feldberg, J. and Glenny, D. Rycroft, S. D. Wilson, R. Hentschel, J. K., Feldberg, Scha A. Wilson, R. Groth, H. K., Feldberg, enih,J,S og .Y,A Scha A. Yu, Y. Dong, S. J., Heinrichs, enih,J,S .Gasen .Wlo,adH cnie.2005b. Schneider. H. and Wilson, R. Gradstein, R. S. J., Heinrichs, enih,J,J etce,A obsh .Fei,J es,M Edelmann, M. Reise, J. Fiebig, A. Bombosch, A. Hentschel, J. J., Heinrichs, Jnemnide lgohlca) p 3–5 in evolution 433–458 and Biodiversity Pp. Plagiochilaceae). (Jungermanniidae: Lejeuneaceae). oeua era. molecular ijntosi h liverwort and the delimitations species in on hypotheses disjunctions Testing 2006. Heinrichs. likelihood. maximum by phylogenies Biology large estimate to DNA. Evolution ribosomal & nuclear of inferred sequences (Hepaticae) ITS Lejeuneaceae using the in relationships Phylogenetic morphol- and sequences DNA ogy. chloroplast of on evolution based neotenic (Lejeuneaceae) and relationships Phylogenetic Society Linnean (Hepaticae). the Lejeuneaceae of Journal of cal genera the of analysis logenetic the 9–22. morphologies: 114: unusual of of evolution case the triggers specialization hw .J hw n .Va J. and Shaw, J. Scha A. A. Shaw, Zhu, R.-L. Long, D. Eckstein, Aphanolejeunea feouinr laws. evolutionary of liver- the of 92–101. phylogeography 114: and status genus The 2011. wort Shaw. J. A. and (mosses). biogeogra- and speciation cryptic phy. markers, classification, DNA on chloroplast comments and (Jungermanniales, with nuclear Adelanthaceae on of based Marchantiophyta) phylogeny A 2010. Heinrichs. (Lejeuneaceae). y of nym genus 2012b. Schmidt. liverwort leafy Jungermanniales). temperate egahcldistribution. geographical and variation genetic between liverwort correlation data: sequence leafy DNA roplast the of biogeography Herbertus Phylogenetic 2007. Heinrichs. of sequences. morphology nrITS from and in range niopsida): speciation Cryptic 2004. rnfro h hohtcedmcliverwort endemic solved: rheophytic mystery year-old the 150 of A 2012c. transfer Schmidt. R. A. and Hentschel, 6 1105–1114. 56: nuclear markers. from DNA inferred chloroplast Porellales) and (Jungermanniopsida, at of or l. species distribution s. One Dumort. 2010. and Shaw. Delimitation J. A. eight? and least Shaw, B. Konrat, von M. 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JQ991259; JQ991144, JQ991029, 25955 25948 microscopica Verwimp & NIA. &Po Mu JQ991263. JQ991368. kulenensis 20100421–21 JQ991297. JQ991182, eBn ceso ubr r itdi h olwn re:nrITS, order: following the order. in trnL–F alphabetical listed are in numbers analyses accession phylogenetic GenBank for used numbers accession Q916 JQ991281. JQ991166, CHINA. JQ991280; CHINA, JQ991279. JQ991164, Q919 JQ991274. JQ991159, blepharophylla 1624 Davison biddlecomiae Cololejeunea CHINA. CHINA. JQ991271; JQ991156, eg&Y 20100719–43 Yu bhutanica Cololejeunea & Peng BOLIVIA. apiculata Q353 DQ238571. DQ238563, ZEALAND. NEW Q914 Q929 IIISLANDS. FIJI JQ991299; JQ991184, &Po (A BB25 cardiocarpa Cololejeunea SWITZERLAND. Scha Scha Q907 Q912 JQ991267. JQ991152, JQ991037, pr 91 Sporn 9960/T Streimann & JQ991300. JQ991185, JQ991070, GE) Q904 Q919 JQ991284. JQ991169, JQ991054, (GOET), cordifolia Po JQ991291. CHINA. Benedix. M2542 JQ991289. CALEDONIA. JQ991285. JQ991170, NEW JQ991286; CALEDONIA. NEW tLneb)Shfn MALAYSIA. Schiffn. Lindenb.) et JQ991293. JQ991178, JQ991063, 20100714–6B (HSNU), Yu gottschei Cololejeunea & Peng cif.FJ ISLANDS. FIJI JQ991288. Schiffn. JQ991173, JQ991058, (E), 37291 distalopapillata Shevock & Long S at)Mzt CHINA. Mizut. Hatt.) (S. CHINA. 59H JQ991294; JQ991179, JQ991064, (HSNU), CHINA. JQ991296; JQ991181, JQ991066, (GOET), ˚ ´ ¨ s&Po & cs gt. tp.FJ ISLANDS. FIJI Steph. ngstr.) paoeenamadeirensis Aphanolejeunea Appendix lrNC281 ller ¨ ¨ ´ ´ e-ewm ewm 29744 Verwimp & fer-Verwimp 31568 fer-Verwimp ´ s03279/AV cs HN) Q905 Q910 JQ991295. JQ991180, JQ991065, (HSNU), s03283/O cs s08037/Q cs Mu GE) Q908 Q913 JQ991278. JQ991163, JQ991048, (GOET), Txe)G i tPo et Kis G. (Tixier) GE) Q908 Q913 JQ991258. JQ991143, JQ991028, (GOET), eune nbl eeotie rmGenBank. from obtained were bold in Sequences . E,J916,J917,JQ991290. JQ991175, JQ991060, (E), ¨ ´ lrNC18A ller GE) Q902 Q917 JQ991302. JQ991187, JQ991072, (GOET), h ta.20100826–23B al. et Zhu s03267/C cs ´ ooeenaaequabilis Cololejeunea E .Jns .M cut TANZANIA. Schust. M. R. Jones) W. (E. age l 20090923–17D al. et Wang Seh)R .Shs.PPANWGUINEA. NEW PAPUA Schust. M. R. (Steph.) ooeenaangustiflora Cololejeunea rdti 7235 Gradstein s THAILAND. cs. ooeenafloccosa Cololejeunea iirFJ ISLANDS. FIJI Tixier ooeenaduvignaudii Cololejeunea Po .Nms rgn,vuhr hraim,adGenBank and (herbarium), vouchers origins, Names, 1. ´ HN) Q906 Q911 Q926 IIISLANDS. FIJI JQ991266; JQ991151, JQ991036, (HSNU), Ty. .Eas MADEIRA. Evans. A. (Tayl.) HN) Q908 Q913 JQ991298. JQ991183, JQ991068, (HSNU), Po s TANZANIA. cs. GE) Q903 Q918 JQ991273. JQ991158, JQ991043, (GOET), ´ HN) Q902 Q917 Q922 IIISLANDS. FIJI JQ991282; JQ991167, JQ991052, (HSNU), GE) Q901 Q916 JQ991261. JQ991146, JQ991031, (GOET), HN) Q904 Q919 Q924 E CALEDO- NEW JQ991264; JQ991149, JQ991034, (HSNU), E .Jns .M cut MALAWI. Schust. M. R. Jones) W. (E. s IIISLANDS. FIJI cs. HN) Q902 Q917 JQ991262. JQ991147, JQ991032, (HSNU), eg&Y 20100720–75B Yu & Peng oge l 38620 al. et Long GE) Q903 Q918 Q933 INDONESIA. JQ991303; JQ991188, JQ991073, (GOET), J) Q907 Q912 JQ991257. JQ991142, JQ991027, (JE), HN) Q903 Q918 JQ991283. JQ991168, JQ991053, (HSNU), HN) Q905 Q910 JQ991265. JQ991150, JQ991035, (HSNU), ooeenacocoscola Cololejeunea ooeenajaponica Cololejeunea ooeenaceratilobula Cololejeunea o ort8 eag 503 Herangi 81 Konrat von ooeenadenticulata Cololejeunea eg&Wi20100920–8B Wei & Peng ooeenacalcarea Cololejeunea GE) Q907 Q912 Q927 GREECE. JQ991277; JQ991162, JQ991047, (GOET), E,J915,J917,JQ991287. JQ991172, JQ991057, (E), ooeenalanciloba Cololejeunea Seh)Pande (Steph.) rlee iu.NEPAL. Mizut. et Grolle Mu ´ GE) Q908 Q913 Q928 CHINA. JQ991268; JQ991153, JQ991038, (GOET), s THAILAND. cs. ¨ (AustinexPearson)A.Evans.U.S.A.Tennessee: hn ta.20091031–67C al. et Zhang rhad&Dewl 960325 Drehwald & Drehwald Mn. .Eas BRAZIL. Evans. A. (Mont.) og33920 Long lrNC17A ller og34790 Long nta 736 Inuthai HN) Q902 Q917 JQ991292. JQ991177, JQ991062, (HSNU), HN) Q909 Q914 JQ991269. JQ991154, JQ991039, (HSNU), Shfn)Gol.MADEIRA. Grolle. (Schiffn.) ooeenaappressa Cololejeunea hnroeenaschimizui Chondriolejeunea Scha Po Po Po HN) Q902 Q917 JQ991272. JQ991157, JQ991042, (HSNU), ooeenalaevigata Cololejeunea ´ GE) Q906 Q911 JQ991276; JQ991161, JQ991046, (GOET), ´ SneLc)Shfn IIISLANDS. FIJI Schiffn. Lac.) (Sande Lh.e idn. cif.CHINA. Schiffn. Lindenb.) et (Lehm. Po ´ s&Po & cs s&Po & cs ¨ s86203/W cs ´ e-ewm ewm 25854 Verwimp & fer-Verwimp Scha Mu s&Po & cs E,J914,J916,JQ991275. JQ991160, JQ991045, (E), ooeenadozyana Cololejeunea Seh)Mzt E CALEDONIA. NEW Mizut. (Steph.) ´ CHINA. . ¨ .W oe.MALAWI. Jones. W. E. lrNC1D ller ¨ E,J914,J915,JQ991270; JQ991155, JQ991040, (E), Po ooeenahaskarliana Cololejeunea e-ewm ewm 18866/A Verwimp & fer-Verwimp Scha ´ HN) Q909 JQ991174, JQ991059, (HSNU), HN) Q906 JQ991171, JQ991056, (HSNU), ooeenadecliviloba Cololejeunea HN) Q900 JQ991165, JQ991050, (HSNU), ´ s03309/BM cs ´ hnaaraitKL1/1 Chantanaorrapint HN) Q918 JQ991253, JQ991138, (HSNU), s&Po & cs ´ s03284/L cs s03279/AY cs ooeenafalcata Cololejeunea HN) Q901 JQ991176, JQ991061, (HSNU), Shfn)Mzt CHINA. Mizut. (Schiffn.) ¨ iir() IIISLANDS. FIJI (I). Tixier e-ewm Verwimp & fer-Verwimp Scha .W oe GERMANY. Jones W. E. ooeenahorikawana Cololejeunea paoeenasintenisii Aphanolejeunea tp.AUSTRALIA. Steph. P .Ce)R .Schust. M. R. Chen) C. (P. Hrk)S at CHINA. Hatt. S. (Horik.) E,J913,JQ991148, JQ991033, (E), ooeenaceatocarpa Cololejeunea eg&Y 20100720–102 Yu & Peng og17551 Long ¨ ooeenalatilobula Cololejeunea eg&Y 20100714–18 Yu & Peng e-ewm Verwimp & fer-Verwimp ´ eg&Y 20100720– Yu & Peng Po s03308/H cs GE) DQ987349, (GOET), HN) JQ991055, (HSNU), HN) JQ991051, (HSNU), ´ s6966/AA cs HN) JQ991049, (HSNU), A vn)Benedix. Evans) (A. HN) JQ991067, (HSNU), HN) JQ991069, (HSNU), Scha GE) JQ991026, (GOET), HN) JQ991044, (HSNU), Chondriolejeunea ogtsM2403C Hodgetts ohd Pereira & Borhidi temn 41383 Streimann Aphanolejeunea Scha ¨ Mt. Tilden. (Mitt.) e-ewm & fer-Verwimp Cololejeunea E,JQ991041, (E), N ia. G. Kitag.) (N. Cololejeunea Cololejeunea Cololejeunea Cololejeunea Cololejeunea SneLac.) (Sande ¨ fer-Verwimp (HSNU), (HSNU), Hodgetts (GOET), (GOET), (Horik.) (Lehm. Steph. 29522 rbcL, Po Po Po Po Zhu ´ ´ ´ ´ cs cs cs cs Delivered by Publishing Technology to: The Natural History Museum, LIS, IP: 157.140.122.156 on: Tue, 17 Sep 2013 14:03:19 Copyright (c) American Society for Plant Taxonomists. All rights reserved. cif.ITALY. Schiffn. JQ991335. JQ991220, CHINA. JQ991105, JQ991334. JQ991219, Q901 Q916 JQ991311. CALEDONIA. JQ991196, KENYA. JQ991081, JQ991310; JQ991195, JQ991080, (E), schaeferi 20100720–51 Yu rotundilobula Cololejeunea Scha CHINA. JQ991336; CALEDONIA. CHINA. NEW Steph. JQ991332; JQ991217, JQ991102, 20100720–91 (HSNU), Yu pseudoserrata Cololejeunea & Peng JQ991330. CHINA, JQ991329; CHINA. JQ991301. Benedix JQ991186, (Horik.) JQ991071, (HSNU), CHINA.7C JQ991325; JQ991210, JQ991095, (HSNU), JQ991309. ooeenaminutissima Cololejeunea AY144483. AY125942, AY125346, Q905 Q920 Q935 AAYILNS aPalma: La ISLANDS. CANARY JQ991315; JQ991200, JQ991085, Q910 Q925 JQ991340. JQ991225, JQ991110, planissima Cololejeunea CHINA. CHINA. JQ991322. JQ991207, JQ991092, NC6E ornata Cololejeunea 03 UE L OOLS NEIHLOSLVROT CLLJUE)563 CHINA. Benedix. macounii 20100423–16 (COLOLEJEUNEA) LIVERWORTS EPIPHYLLOUS 20111012–72 Peng IN & HOMOPLASY AL: ET longifolia YU Cololejeunea USJ46100 s.n. Carranza-Morse CHINA. JQ991366. JQ991251, JQ991305; JQ991190, CHINA. JQ991304; BANGLADESH. Tixier (Herzog) 2013] ewm ewm 24798 Verwimp ECUADOR. & Verwimp serrata 20090221–41A Wei JQ991314. CHINA. CHINA. JQ991319. JQ991204, JQ991089, cut BRAZIL. Schust. Q923 Q938 BRAZIL. JQ991318; JQ991203, JQ991317. JQ991202, JQ991087, ninguana Cololejeunea cif.BRAZIL. Schiffn. ¨ HN) Q908 Q923 Q938 CHINA. JQ991328; JQ991213, JQ991098, (HSNU), e-ewm ewm 15936 Verwimp & fer-Verwimp HN) Q913 Q928 JQ991333. JQ991218, JQ991103, (HSNU), Seh)Bndx E CALEDONIA. NEW Benedix. (Steph.) eg&Y 20091031–106A Yu & Peng hn ta.20091101–48F al. et Zhang eg&Y 20100720–75A Yu & Peng eg&Y 20100713–30 Yu & Peng rle MADEIRA. Grolle. Src xUdr. .Eas CHINA. Evans. A. Underw.) ex (Spruce ooeenaminutissima Cololejeunea ooeenapseudoplagiophyllaCololejeunea ooeenamalanjae Cololejeunea Scha HN) Q908 Q913 JQ991308. JQ991193, JQ991078, (HSNU), Mu eg&Wi20100912–40A Wei & Peng og35468 Long ¨ e-ewm ta.24473 al. et fer-Verwimp rdti 9800 Gradstein HN) Q919 Q924 JQ991339. JQ991224, JQ991109, (HSNU), ¨ Scha lrNC13I ller HN) Q911 Q926 JQ991341. JQ991226, JQ991111, (HSNU), .Eas CHINA. Evans. A. eg20100522–9A Peng HN) Q907 Q912 Q937 CHINA. JQ991307; JQ991192, JQ991077, (HSNU), eg20100711–5A Peng ooeenalinopteroides Cololejeunea eg&Wi20100921–33C Wei & Peng ¨ eg&Wi20100921–39 Wei & Peng e-ewm ewm 11238 Verwimp & fer-Verwimp iir E CALEDONIA. NEW Tixier. Mt. eei xMzt HN.Xizang: CHINA. Mizut. ex Benedix (Mitt.) Mt. by.CHINA. Abeyw. (Mitt.) S. cif.CHINA. Schiffn. (Sm.) u20100922–6 Yu P .W tP .Ln ipo CHINA. Piippo. Lin) J. P. et Wu C. (P. E,J910,J912,J913;GREECE. JQ991337; JQ991222, JQ991107, (E), HN) Q911 Q926 JQ991331. JQ991216, JQ991101, (HSNU), iir E CALEDONIA. NEW Tixier. Scha GE) Q903 Q918 JQ991313; JQ991198, JQ991083, (GOET), Scha HN) Q902 Q917 JQ991312. JQ991197, JQ991082, (HSNU), GE) Q906 Q911 JQ991306. JQ991191, JQ991076, (GOET), ooeenaperaffinis Cololejeunea ooeenaocelloides Cololejeunea ooeenaschmidtii Cololejeunea eg&Wi20100920–3A-4 Wei & Peng GE) Q906 Q921 JQ991316. JQ991201, JQ991086, (GOET), ¨ ooeenaobliqua Cololejeunea HN) Q900 Q925 JQ991320; JQ991205, JQ991090, (HSNU), ooeenarossettiana Cololejeunea ¨ GE) Q918 Q923 JQ991338. JQ991223, JQ991108, (GOET), e-ewm ewm 25642 Verwimp & fer-Verwimp HN) Q901 Q926 JQ991321. JQ991206, JQ991091, (HSNU), ooeenamaritima Cololejeunea HN) Q904 Q929 JQ991324; JQ991209, JQ991094, (HSNU), u20100921–1 Yu HN) Q903 Q928 JQ991323. JQ991208, JQ991093, (HSNU), og281211 Long e-ewm ewm 12006 Verwimp & fer-Verwimp ooeenamadothecoides Cololejeunea ssp. tp.MALAWI. Steph. eg&Wi20100913–12A Wei & Peng myriocarpa HN) Q909 JQ991194, JQ991079, (HSNU), HN) Q910 JQ991215, JQ991100, (HSNU), HN) Q909 JQ991214, JQ991099, (HSNU), GE) Q904 JQ991199, JQ991084, (GOET), HN) Q916 JQ991221, JQ991106, (HSNU), .C ue .X u.CHINA. Luo. X. J. et Wu C. P. ooeenapseudofloccosa Cololejeunea Po ´ E,J917,JQ991189, JQ991074, (E), u20100922–11 Yu s&Po & cs Mu ooeenaraduliloba Cololejeunea .Rb OT RICA. COSTA Rob. H. ooe ta.56319 al. et Koponen eg&Wi20100912– Wei & Peng h ta.20100822–55 al. et Zhu ¨ Mu lrNC11D ller Ne tMn. .M. R. Mont.) et (Nees HN) JQ991136, (HSNU), HN) JQ991075, (HSNU), HN) JQ991104, (HSNU), ¨ GE) JQ991088, (GOET), e 20100210–62A Wei lrNC2B ller ´ Shfn)Schiffn. (Schiffn.) s04027/BD cs Ne tMont.) et (Nees Hrk)S Hatt. S. (Horik.) tp.CHINA. Steph. olyM280H Porley iir NEW Tixier. Cololejeunea Mu Cololejeunea Cololejeunea C Massal.) (C. ¨ lrNC9I ller (HSNU), (HSNU), (HSNU), (HSNU), (HSNU), (GOET), (GOET), (Steph.) Scha eg& Peng Mu Wang (JE), (H), ¨ ¨ Zhu fer- ller tenuicornis rdti ta.10033 al. et Gradstein JQ991369. JQ991254, AZORES. Dumort. VIETNAM. DQ238583. ECUADOR. DQ987441. ECUADOR. cerina Lejeunea DQ987411. GERMANY. DQ983688, DQ987307, catinulifera DQ987422. MALAYSIA. sp. 04–06 al. et 22538/A 20111018–86C CHINA. Peng & Wang Q828 Y406 DQ238582. AY548096, ssp. DQ987288, Winkl. S. 34521 isne l 04–02 al. et Wilson CHINA. CHINA. Mizut. iir IIISLANDS. FIJI Tixier. 7725 Yong JQ991342. JQ991227, JQ991112, THAILAND. GE) Q912 Q927 JQ991352. JQ991237, JQ991122, (GOET), 0039/N JQ991350. JQ991235, BRAZIL. 20100720– Yu Tixier. & JQ991349. JQ991234, Peng JQ991119, (HSNU), CHINA. 77H JQ991348; JQ991233, JQ991118, (HSNU), stylosa Cololejeunea ECUADOR. JQ991346. 20100720–100 Yu CHINA. & Peng JQ991344. Q911 Q926 JQ991361. JQ991246, JQ991131, CHINA. JQ991362; JQ991247, JQ991132, DQ238573. Q928 Q933 THAILAND. JQ991353, JQ991238, THAILAND. JQ991354. JQ991239, JQ991124, (GOET), ooeenatrichomanis Cololejeunea iir OT RICA. ISLANDS. COSTA Tixier. FIJI JQ991359; 03282/BB JQ991244, JQ991129, (HSNU), trichomanis 20100719–16 Yu 20100719–18A 02105/E E,J919,J911,JQ991327. JQ991212, JQ991097, (E), GE) Q911 Q926 JQ991351; JQ991236, JQ991121, (GOET), J) Q910 Q925 JQ991370. JQ991255, JQ991140, (JE), og34518A Long J) Q917 Q922 Q937 CHINA. JQ991357; JQ991242, JQ991127, (JE), og32613 Long HN) Q910 Q925 JQ991360. JQ991245, JQ991130, (HSNU), ooeenastotlerianaCololejeunea eg&Y 20100714–7A Yu & Peng ooeenaspinosa Cololejeunea B) Q918 Q923 JQ991358. JQ991243, JQ991128, (BM), yiclossgymnocolea Myriocoleopsis A vn)Seh ETINDIES. WEST Steph. Evans) (A. ooeenawightii Cololejeunea Po eenacancellata Lejeunea yiclossvuquangensis Myriocoleopsis GE) Q833 Q866 DQ987437. DQ983676, DQ987333, (GOET), ssp. pue ECUADOR. Spruce. ´ enih 3695 Heinrichs rdti ta.10020 al. et Gradstein Po rdti 12073 Gradstein isne l 04–13 al. et Wilson s02102/N cs Scha HN) Q916 Q921 Q936 CHINA. JQ991356; JQ991241, JQ991126, (HSNU), ´ s&Po & cs Andicola u20100921–4 Yu li-ogse l 3024 al. et Ilkiu-Borges HN) Q913 Q928 JQ991343. JQ991228, JQ991113, (HSNU), Scha cordiflora ¨ Lh.e idn. otce idn.e Nees. et Lindenb. Gottsche, Lindenb.) et (Lehm. e-ewm ewm 16233 Verwimp & fer-Verwimp GE) Q839 Q866 DQ987433; DQ983686, DQ987329, (GOET), ouairrorata Colura ¨ Scha ooeenatenella Cololejeunea tp.e iu.CHINA. Mizut. ex Steph. Po e-ewm ewm 9212 Verwimp & fer-Verwimp HN) Q914 Q929 Q934 CHINA. JQ991364; JQ991249, JQ991134, (HSNU), GE) Q829 Y403 DQ238584. AY548073, DQ987279, (GOET), Po E,J919,J911,J912,CHINA. JQ991326, JQ991211, JQ991096, (E), ´ ´ s07014/AL cs s&Po & cs ´ ¨ sSV/H-0473/A cs GE) Q837 Q860 DQ987449. DQ983670, DQ987347, (GOET), .ECUADOR. e-ewm&Vrip29196 Verwimp fer-Verwimp& Gtsh)Seh VIETNAM. Steph. (Gottsche) Po GE) Q829 Y412 DQ238581. AY548102, DQ987259, (GOET), HN) Q915 Q920 JQ991345; JQ991230, JQ991115, (HSNU), ´ ´ ooeenaserrulata Cololejeunea HN) Q915 Q920 JQ991365; JQ991250, JQ991135, (HSNU), sFJ ISLANDS. FIJI cs s03289/BO cs B) Q917 Q922 JQ991347. JQ991232, JQ991117, (BM), HN) Q913 Q928 JQ991363; JQ991248, JQ991133, (HSNU), Hrk)Pande (Horik.) Diplasiolejeunea ese ot xMn.ECUADOR. Mont. ex Mont. et Nees ooeenayakusimensis Cololejeunea tp.MALAYSIA. Steph. rdt,IkuBre &Vanderpoorten. Ilkiu-Borges Gradst., rdti ad 10141 Mandl & Gradstein Scha Src)Hircse l ECUADOR. al. et Heinrichs (Spruce) J) Q915 Q920 JQ991355. JQ991240, JQ991125, (JE), eenacavifolia Lejeunea GE) Q839 DQ983689, DQ987339, (GOET), ¨ ooeenathailandensis Cololejeunea GE) Q827 DQ238568, DQ987277, (GOET), ooeenatranninhiana Cololejeunea e-ewm ewm 16234/A Verwimp & fer-Verwimp Src)M .Rie tGradst.. et Reiner E. M. (Spruce) eg&Y 20100719–2 Yu & Peng Scha HN) Q916 JQ991231, JQ991116, (HSNU), GE) Q838 DQ983678, DQ987318, (GOET), GE) Q838 DQ238564, DQ987348, (GOET), eei.Austraria. Benedix. (Po HN) Q914 JQ991229, JQ991114, (HSNU), ouacalyptrifolia Colura ¨ ooeenasubcardiocarpa Cololejeunea e-ewm 23508 fer-Verwimp Scha ´ ilsoeenainvoluta Diplasiolejeunea ´ s&T .Nn)Po Ninh) N. T. & cs eg&Y 20100720–47B Yu & Peng tR .Msa CHINA. Misra. N. R. et ¨ ooeenasocietatis Cololejeunea e-ewm Verwimp & fer-Verwimp Po p,ECUADOR sp., ooeenavitalana Cololejeunea ´ s&Po & cs Po GE) JQ991123, (GOET), tp.MALAYSIA. Steph. GE) JQ991120, (GOET), ´ se l 0028/M al. et cs og7721 Yong Drepanolejeunea J) JQ991139, (JE), Er. Lindb. (Ehrh.) Po ´ Po Cololejeunea eg&Yu & Peng s03288/CU cs ´ ´ s&Ninh & cs s&Po & cs Po S Hatt.) (S. Lejeunea (HSNU), ´ (GOET), (GOET), se al. et cs (Hook.) eg& Peng Colura Wilson Tixier. Tixier. (BM), Long ´ ´ cs. cs