Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 2 Microsorum 2018 21, June publication Taxonomists of Plant Date of Society American 10.1600/036364418X697166 the DOI by 2018 Copyright © Botany Systematic eie pmrhe,sc as clearly such with monophyletic flux. apomorphies, are in genera is I defined segregate ferns the Group microsoroid of the Phylogeny Some of 2004b; Pteridophyte taxonomy al. Generic 2014; et 2016). Sundue (Schneider dis- and paleotropics ferns the Testo epilithic in of clade mainly or spp.) tributed epiphytic, 183 ca. hemiepiphytic, genera; terrestrial, (12 diverse a subfamily (Polypodiaceae, Microsoroideae), ferns microsoroid of species tween 2013). al. et (Zhang data to of addition sources in Sigel other markers, 2012; genetic al. inherited should biparentally taxa et hybrid include putative Beck of investigations (e.g. detect Thus, 2014). origins al. et repeated to hybrid- as with insufficient such associated ization, variation. is processes evolutionary intraspecific in- alone complicated processes and morphology various plasticity Furthermore, through phenotypic arise morpho- cluding origin, may hybrid useful variation of a logical hypotheses provides in- establishing taxa morphology for While parent basis 2015). putative al. Moran between et 1997; termediate Testo al. 2004; et Watkins on Manickam solely and (e.g. based How- are 2014). evidence hybrids al. 1954; fern morphological of et Wagner cases Rothfels (e.g. reported 2010; many lineages al. ever, fern et docu- of Beck been 1990; range has Barrington wide Hybridization a 1989). relative in with al. mented sterility et hybrid (Barrington spores), overcome ease unreduced to and them via polyploidy allow reproduction both which of asexual rates 2016). (i.e. high apogamy Ebihara have taxa, varieties; ferns non-hybrid and 720 Furthermore, subspecies, vs. species, hybrid (380 including flora of third hybrid pteridophyte one ca. of for the account numbers hybrids Japan, high in the example, have For in taxa. which significant ferns, particularly of is diversification Hybridization lineages 1966). separated previously (Mayr between flow gene allowing by of diversity decrease generation also may the Burke hybridization and Alternatively, by (Chapman 2007). fertile or are hybrids 1985) if taxa Hewitt new completely and (Barton parents relative fitness diversity to reduced have species hybrids if increase reinforcement can through Hybridization 1959). Stebbins 1949; (Anderson diversification evolutionary in role important yboi soitoswt ns(afe ta.20;Kreier 2003; al. et (Haufler ants with associations symbiotic urn drs:Dprmn fBtn,Ntoa uemo aueadSine -- mkb,Tuua aa,305-0005 Japan, Tsukuba, Amakubo, 4-1-1 Science, and Nature of Museum National Botany, of Department address: Current 1 ee eivsiaeapttv aeo yrdzto be- hybridization of case putative a investigate we Here, an plays species, between interbreeding or Hybridization, eateto raimcadEouinr ilg n avr nvriyHrai,HradUiest,Cambridge, University, Harvard Herbaria, University Harvard and Biology Evolutionary and Organismic of Department en fteSceyIlnsi provided. is Islands Society the of ferns in ie hnltrlpna,adrudrioesae ihetr agn.Gntceiec sas rsne o h is iespotn the supporting time first the for presented also is evidence Genetic margins. entire endemic with scales of rhizome round origin and hybrid pinnae, lateral than wider analysis. pinna phylogenetic molecular and Polynesian morphology on French based described is Polynesia French Abstract Keywords irsrmspectrum Microsorum 21) 32:p.397 pp. 43(2): (2018), — — e yrdmcooodfern, microsoroid hybrid new A gapCp Microsorum 3 Microsorum Moorea, , tohieaense – 413 3 h mlctoso yrdzto o h aooyo irsri en r icse,adakyt h microsoroid the to key a and discussed, are ferns microsoroid of taxonomy the for hybridization of implications The . rbcL ytecmiaino oita r itiue oeo esi igeln ewe h oteadmris apical margins, and costae the between line single a in less or more distributed are that sori of combination the by maximum mlctosfrteTxnm of Taxonomy the for Implications oit sad,Tahiti, Islands, Society , olH Nitta, H. Joel Lecanopteris Plpdaee,aNwHbi enfo rnhPlnsa with Polynesia, French from Fern Hybrid New a (Polypodiaceae), ( irsrmgrossum Microsorum 3 uhrfrcrepnec ([email protected]) correspondence for Author Microsorum omnctn dtr ljnr Vasco Alejandra Editor: Communicating hc forms which , 1,2,3 ascuet 23,USA 02138, Massachusetts trnL 3 adAmer, Saad 3 tohieaense – irsrmpunctatum Microsorum F . 397 ( irsrmcommutatum Microsorum /tmsac/om,o rs aeil ebru brvain follow abbreviations Vascular Herbarium P material. the fresh from or area p/item/search/form), (https://science.mnhn.fr/institution/mnhn/collection/ downloaded at study database specimens specimens our herbarium herbarium Plants using of from 1) images species Fig. GH, Polynesia; microsoroid French each Islands, (Society of specimens multiple rnhPlnsa(it ta.21) hr r ee aaof Copel., Islands: taxa Islands, Society seven the Society are spicatus from There Tahiti, known 2017). and ferns al. microsoroid Moorea et (Nitta of Polynesia ferns French the of genetically. survey verified been have that described cases been any are we of have 1997), unaware (Nooteboom ferns taxa microsoroid the hybrid in previously various (Tejero-D help genera Although to and taxa 2009). species of may of distinctness hybridization delimitation genetic of inform the Studies for 2008). and al. evidence 2017), et provide al. Kreier et 2004b; Wei al. b; et 2010a, al. et b; Wang Microsorum 2004a, 2008; al. al. et et (Schneider Kreier other investigations within molecular nested be in to the genera shown in been have placed 1997) formerly Nooteboom species many However, genus 2008). al. et irsrmmembranifolium Microsorum ino hspata e hybrid, new a as between plant recogni- this a support of gross that be tion analyses in could our molecular of species and results suspected morphological present these we therefore Here, of taxon. we undescribed any previously that match and not morphology, did of that population a Moorea encountered our we During press). survey, in Florence field 2011; al. et Nitta 2003; Smith and grossum between M. hybrid putative (a Copel. (Brack.) punctatum sorum [ Copel. (Baker) eicuesmlso endemic of samples include we oe.fo aai n ics h aooi implications for taxonomic results the our discuss of and Hawaii, from Copel. 1 spr faohrsuy ercnl odce field a conducted recently we study, another of part As opooia Analysis Morphological n hre .Davis C. Charles and Microsorum irsrmgrossum Microsorum .commutatum M. Lf)L Wang, Li (L.f.) ,adpsil niaigahbi rgnfrteHawaiian the for origin hybrid a indicating possibly and ), Cpln 92 ye n ae19;Murdock 1996; Game and Sykes 1932; (Copeland ) Microsorum sbdyi edo aooi eiin(Schneider revision taxonomic of need in badly is Microsorum 5 aeil n Methods and Materials Microsorum irsrmparksii Microsorum L)Cpl,and Copel., (L.) ntebsso opooy(omn1991; (Bosman morphology of basis the on 3 3 irsrmmembranifolium Microsorum — tohieaense and eosre opooia hrcesin characters morphological observed We irsrmcommutatum Microsorum 1 RB. Ching, (R.Br.) Lns.&Fsh)S.B.Andrews, Fisch.) & (Langsd. .membranifolium M. . a edsigihdfo other from distinguished be can irsrmspectrum Microsorum Microsorum Microsorum Cpl)Copel.], (Copel.) irsrmpowellii Microsorum .punctatum M. rmMoorea, from ) Microsorum Furthermore, . 3 3 tohieaense maximum Lepisorus ´ ı ze al. et ez (Blume) (Kaulf.) Micro- and on , Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 398 esrda h ietpito h hzm.W xmndsoe fthe of diameter spores rhizome examined We and rhizome. specimen, the of per point mea- frond widest were the largest traits at dis- single frond measured lamina Quantitative the of pairs. on degree pinna sured length, of stipe number and sori, width, section, frond rhizome of scales, length, rhizome color, arrangement frond spore lamina, diameter, include: in sori of characters immersion of degree Morphological (2018). Thiers B niae yrcage .SceyIlns asaatdfo iiei omn ne raieCmosLicense. Commons Creative under Commons Wikimedia from adapted Maps Islands. Society B. rectangle. by indicated (B) Fig. .Mpo h td ra .Fec oyei,soigtelctoso ora(oit sad)adU o MrussIlns.Lcto finset of Location Islands). (Marquesas Pou Ua and Islands) (Society Moorea of locations the showing Polynesia, French A. area. study the of Map 1. YTMTCBTN Vlm 43 [Volume BOTANY SYSTEMATIC oce pcmn olce o hssuyaedpstdi CadGH. and UC in deposited are study 2004). this al. for et collected (Abramoff specimens ImageJ Voucher used width images spore digital and on perine, made Measurements length. the spore to perpendicular excluding approximately way similar spore measured a the in was measured was of length part Spore widest with the camera). taken across microscope (photographs digital microscope DP70 compound Olympus a using plant unknown Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. na12%aaoegli oiid1 modified in gel agarose 1.25% a on eetimd n h osnu eune eeepre nFASTA in exported were sequences consensus the format. sequences and primer PCR trimmed, containing were regions Flanking plastid specimen. each for per obtained marker sequence contigs, consensus into single assembled a 2012), and al. edited, et manually (Kearse 9.1.3 v. Geneious into imported akn nrn,i igePRrato Sheteze l 08.We 2008). al. et (Schuettpelz reaction 25 PCR two single the a separated in introns, lacking and bp) 900 n nsatmtclytimdwt nerrctf f40 e base. per 4.0% of cutoff error an Geneious, into with imported trimmed were automatically files ends we AB1 sequencing and case, M13F. first using the the from sequence not genotype to was known reaction of this sample if PCR single (M13F); reverse a primer selected the forward for need the used the obviated with and only orientation sequencing forward we the colony in one that obtained least was at allele except Typically, per costs. reduce regions, to sequencing plastid for (M13R) for primer M13R. as and the way in M13F performed same were primers sequencing and vector-based success amplification of using Verification amplified manufacturer and the specimen, following 12 Massachusetts) structions. Waltham, alleles Scientific, separated we present), were each peaks within multiple result (i.e. not did sequence sequencing clean purified direct a that case in excised, the In products. the plastid the as direct-sequence way to attempted gapCp manufacturer first the We following structions. Massachusetts) extraction gel Billerica, Montage the (Millipore, using the kit purified allowed clean were a bands This using Excised excised hr. blade. and 2 light razor UV for under current distinguished visually V be 60 to bands a to subjected and Massachusetts), lifed e esy.I diint h owr n ees PCR reverse to and used were forward (South ESRBCL654R the GENEWIZ and to ESRBCL628F sequence to addition primers sequencing internal In primers, Sanger Jersey). and New hr. Plainfield, cleaning 1 for enzymatic applied V for 80 of current a with gel, agarose 1% 3 including electrophoresis for gel by (2008) inspected al. for et (2007) Schuettpelz Pryer and Schuettpelz of rbcL those followed settings for thermocycler 2008) al. et (Schuettpelz ESGAPCP11R1 ShetezadPyr20)for 2007) Pryer and (Schuettpelz 2018] nrnadthe and intron oiidCA rtcl(ol n ol 97 rteDes kit DNeasy a the using or either 1987) gel manufacturer Doyle the silica following and in California) (Doyle Valencia, preserved (Qiagen, protocol tissue CTAB frond modified on performed was 91 n en I1(ie l 09 for 2009) al. et (Li 1Ir1 FernL and 1991) ( n asivc 92 oe ta.19a.W hrfr eune two sequenced therefore We 1998a). al. the (Gastony et markers, ferns nuclear Vogel in cases 1992; known Yatskievych all almost and in inherited maternally are 1). these (Appendix here used data GenBank all for specimen the or ula aa ugopseiswr eetdfo te aai Poly- in taxa other from selected were ( species podiaceae Outgroup data. nuclear var. var. of specimen one specimens and Hawaii, two as well spectrum as M. Islands, Society the of from ferns microsoroid all hereafter 2008; for markers of taken was two care that accessions Special ensure available GenBank. these to on all markers microsoroid selected same with the Islands for therefore data ferns Society microsoroid our We the complemented context. b; place and 2010a, global to sequencing, al. useful a et are Wang in and 2008; ferns 2017), al. et al. Kreier et 2004b; Wei taxonomic al. broad et a including (Schneider ferns sampling microsoroid of studies previous in used hsHwia nei pce oohrPacific other to species endemic Hawaiian this spectrum sorum aC short gapCp The ucsflpatdPRpout eesn ihu ute modification further without sent were products PCR plastid Successful h C mlfcto a efre sn RL- Tbre tal. et (Taberlet TRNLF-f using performed was amplification PCR The Plastid ti mosbet eethbiiaineet sn nypatdlc,as loci, plastid only using events hybridization detect to impossible is It N xrcin akrSlcin n Sequencing and Selection, Marker Extraction, DNA m .membranifolium M. L pothifolius pothifolius and trnL gapCp C rdcsuigEGPPF n SAC1R ntesame the in ESGAPCP11R1 and ESGAPCP8F1 using products PCR gapCp rbcL rbcL trnL nrnadthe and intron rbcL gapCp Kuf)Cpl var. Copel. (Kaulf.) only). short C rdc a ie ih5 with mixed was product PCR – rbcL 6clne eeslce e ein( region per selected were colonies 16 rmr sdhr yial mlf both amplify typically here used primers ShetezadPyr20) h eutn B ie were files AB1 resulting The 2007). Pryer and (Schuettpelz trnL – and Bc.Hm xDDn ..hn rmOkinawa. from X.C.Zhang D.Don) ex (Buch.-Ham. a nlddt ettemnpyyof monophyly the test to included was F aC long gapCp a nlddt netgt h hlgntcafnte of affinities phylogenetic the investigate to included was , trnL ecp o neln eprtr e o5°)and 56°C) to set temperature annealing for (except and c.60b) nadto osotrbceilfragments bacterial shorter to addition in bp), 600 (ca. oyb lnn sn h OOT i Tem Fisher (Thermo kit TA TOPO the using cloning by copy – trnL gapCp trnF “ – long trnL F – rmteMrussIlns n pcmnof specimen one Islands, Marquesas the from , F negncsae) SBLFadESRBCL1361R and ESRBCL1F spacer), intergenic aC long gapCp trnL icuigthe (including ” oisb e lcrpoei sflos each follows: as electrophoresis gel by copies – and F and IT TA. E YRDMCOOU RMFEC OYEI 399 POLYNESIA FRENCH FROM MICROSORUM HYBRID NEW AL.: ET NITTA etciu ellipticus Leptochilus cesosoiiae rmtesm voucher same the from originated accessions gapCp – trnF aC short gapCp pentadactylum “ short mlfcto ucs a visually was success Amplification . negncsae oehr aebeen have together) spacer intergenic ,and ” 3 oisof copies trnL rbcL A ufr(ilpr,Billerica, (Millipore, buffer TAE trnL gapCp nmlil pcmn from specimens multiple in ) m aC short gapCp – C rdc e aeo a on lane per product PCR L m n SAC81and ESGAPCP8F1 and , – trnF Microsorum Hler)DDPle from D.D.Palmer (Hillebr.) F L6 Tub xMra)Noot. Murray) ex (Thunb. icuigbt the both (including h C rtcl and protocols PCR The . gapCp negncsae alone spacer intergenic aC short gapCp 3 odn y,loaded dye, loading — rDavalliaceae or ) Sheteze al. et (Schuettpelz Microsorum N extraction DNA ,and aC long gapCp ’ instructions. s or .ellipticus L. long Micro- ’ ’ sin- with )per sin- trnL (ca. lgmnso ahmre eaaey nso h eutn alignments resulting for the (50% of data missing generate Ends 90% to than separately. more settings with marker default under each 2002) of al. alignments et (Katoh MAFFT using lyzed otg n xotdi AT omt l el eeae sequences 1). generated (Appendix newly GenBank resulting All in the format. deposited from FASTA been obtained have in were exported alleles) then Con- and final contigs. were contigs into the alleles assembled (i.e. and putative sequences alignment, sensus representing the the to from sequences belong exported of to selected, considered Lists were consensus allele. less the same from with difference each bp sequences 5 containing than groups monophyletic and were occupying excluded, sequences or chimeric clade Obviously a together clades. within between fused sequences positions other intermediate became all artifacts that to sister (i.e. sequences either sequences PCR) distinct during chimeric within of 2 with fragments with implemented of sequences, tree consisting the phylogenetic as of a most 2006) in containing resulted (Stamatakis generally This was RAxML Geneious. analysis re- phylogenetic primer using a Geneious, and alignment, within performed the implemented from as trimmed were 2002) gions al. et (Katoh MAFFT eunigerr sn iia praht hto rs ta.(09:for (2009): al. et Grusz of that to each approach similar a using errors sequencing alleles 3818 Nitta punctatum pcmn ( specimens nxetdyhg ubrof were number high trees unexpectedly of 50% a 25% produce initial to summed The tree. trees less. rule) remaining (majority the or and 0.01 burn-in, We as reached 10,000,000 discarded generations. frequencies for 1000 deviation split run standard every average of the to once when converged allowed had chain chains were that cold confirmed Chains the priors. sampling flat hot, generations, (three using each chains four cold) of runs one Huelsenbeck independent and two (Ronquist with 3 CIPRES on MrBayes 2003) in treespace of searches out carried for used partitioning No partitions. was between branchlengths and topology for except upr hscnlso:ec ou leewsrcvrdol for only recovered was that allele evidence long rogue contami- of each lines of several conclusion: amounts are this There small support 2011). even al. et to (Ruecker sensitive DNA nating highly is which (cloning), epciey epriindtecnaeae lsi lgmn notwo into for alignment one plastid concatenated partitions, the partitioned We respectively. hs pcmn rmorfnlaayi,a ecudntb ofdn of excluded confident be therefore not We could pattern. we mor- as a homology. analysis, such the their final show our accessions about not from other specimens do nothing and these species origin; exactly; same hybrid the species indicate of specimens same these the of phology of others matched ayn eghdet ifrn rmr sdi rvossuis were studies) previous in used primers different the using to trimmed due length varying rtro sipeetdi M Lfr ta.21)o h hM web- se- PhyML the the GTR in on the of 2017) resulted lection al. This et (Lefort (http://www.atgc-montpellier.fr/sms/). SMS server in implemented as criterion RCr em21) rlmnr aiu ieiod(L nlssof analysis (ML) likelihood maximum Preliminary rbcL 2016). Team Core (R R iidtems prpit oe fDAsqec vlto for evolution sequence DNA of model trnL appropriate most the tified heuristic the as random settings 10 same with the replicates under search. BS replicate trees per 1000 two sequences using tree-bisection- addition evaluated than was more with support no was tistical replicates saving tree(s) score addition a parsimonious swapping, with most branch random the (TBR) 1000 for reconnection 2002) search for (Swofford 4.0b heuristic PAUP* conducted A in out CIPRES. carried were on ML a). by analyzed (-f those option as analysis rapid the to using analysis tree bootstrap the likelihood of best results platform the the 1000 wrote conducted computing tree, and likelihood replicates, best Gateway (BS) the bootstrap for searched Science We 2010). CIPRES al. et (Miller the on GTR a (GTRGAMMA) using with out carried 2006) were (Stamatakis separately RAxML gene nuclear each and alignment plastid all for alignment plastid analyses. single a subsequent into markers two these concatenated we so GapCp uigoriiilpyoeei nlss(Lol) eosre an observed we only), (ML analysis phylogenetic initial our During o hlgntcaayi sn aeinifrne(I,w is iden- first we (BI), inference Bayesian using analysis phylogenetic For datasets same the of analyses phylogenetic (MP) parsimony Maximum concatenated the of analyses phylogenetic (ML) likelihood Maximum hlgntcAnalysis Phylogenetic – and gapCp F or ” , .W eiv hs r riat u otepooo eused we protocol the to due artifacts are these believe We ). aC long gapCp short trnL lee eeietfe hl conigfrciea n other and chimeras for accounting while identified were alleles .mmrnflu it 573 Nitta membranifolium M. oyprseie,altimdA1flswr lge using aligned were files AB1 trimmed all specimen, per copy $ – F nvrfrbt) lsi eune fteespecimens these of sequences plastid both); for (never gapCp e elct,adsapn ntebs re ny Sta- only. trees best the on swapping and replicate, per 5 eaaeyddntrva n togyspotdconflicts, supported strongly any reveal not did separately ,and “ 1 rbcL trimEnds htmthdohr o-yrdseis(i.e. species non-hybrid other, matched that ) I lgmns hc eeaaye eaaey We separately. analyzed were which alignments, 1 aC short gapCp n n for one and ,GTR G, — N eune nFSAfra eeana- were format FASTA in sequences DNA ” gapCp ucino the of function 1 I eaaeyuigAkaike using separately lee ntrepttvl non-hybrid putatively three in alleles 1 trnL ,GTR G, 1 , – .pnttmNta1399 Nitta punctatum M. F oe fsqec evolution sequence of model G trnL n nikdalparameters all unlinked and , “ ips – 1 F hc a eune of sequences had which , ” ,adGTR and G, akg Hil21)in 2014) (Heibl package ’ information s 1 – models, I ,and clades 4 “ gapCp rogue rbcL M. , Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. Microsorum 400 icosin riaceous in grossum close M. is It 3). to dissection 2, lamina commutatum Figs. of 2; degree and 1, size fern (Tables overall microsoroid Islands described Society previously the any from of that match not irsri en eercvrda oohltc(desig- 2018). monophyletic al. as et recovered individual nated (Nitta were for analysis Dryad ferns each on Microsoroid dataset by our produced 4, to Fig. phylogenies reader in phylogeny the ML refer the on and based therefore tree summary We mostly a conflicts. were present supported BI strongly and no MP, with ML, consistent by obtained Topologies intron. Microsorum punctatum Microsorum powellii Microsorum membranifolium Microsorum grossum Microsorum commutatum Microsorum spicatus Lepisorus mesd lal iil dxal n omn asdbmst m Deep mm; 1 to bumps raised forming and adaxially visible clearly Slight immersed, adaxially; noticeable not superficial, Sori h ra iia eoioy(it ta.2018). al. et on (Nitta available Repository Digital are Dryad analyses the preliminary and trees, phylogenetic alignments, hzm cls(niemrisi h nnw ln,vs. plant, unknown the in in margins margins denticulate (entire in epiphytic scales vs. rhizome plant, unknown the in petric from in vs. guished irregularities, sori with but scattered plant, unknown many the in costae the of commutatum .0.A npeiu studies, previous in As 1.00). eg rire l 08 htol mlfe h negncspacer intergenic the the amplified between only that 2008) al. et Kreier (e.g. and mutatum between intermediate membranifolium ( be sori M. to arranged regularly appears with plant species occasionally unknown clear, the sori has arranged the in irregularly plant of than occurrence unknown pairs The spores. The pinna misshapen fewer 2). with (Table usually latter and smaller much with well matches h is a 0 po h lgmn u oueo the of use to due missing alignment the were of (50.2%) bp 600 total ca. 106 211 first the of 3). out 242 (Table GenBank from including taxa quences bp 149 2175 representing was individuals final alignment The plastid taxa. 10 concatenated representing individuals 28 from sequences Table measurements, morphological including study this with associated Data Morphology lsi Phylogeny Plastid “ f “ ” M .Qaiaiemrhlgcltat fmcooodfrsfo h oit sad,Fec oyei.Dge fsrlimrina olw:None follows: as immersion soral of Degree Polynesia. French Islands, Society the from ferns microsoroid of traits morphological Qualitative 1. rmrst(aelte l 91 ypeiu studies previous by 1991) al. et (Taberlet set primer , ” .punctatum M. 3 3 .grossum M. nFg ;MLBS 4; Fig. in and , tohieaense maximum mmrncosi h nnw ln,v.co- vs. plant, unknown the in (membranaceous oeo oeie w ie fsr nete side either on sori of lines two sometimes or (one trnL .powellii M. — h opooyo h nnw ln does plant unknown the of morphology The .powelli M. UA 3 (UAA) .membranifolium M. n hs ihsatrdsr ( sori scattered with those and ) — ,dfesi oa ragmn from arrangement soral in differs ), ). egnrtdattlo 0nwplastid new 50 of total a generated We .commutatum M. intfdMsl eua lgtCerRudCahae ih in light Clathrate, Round Clear Slight regular Mostly Pinnatifid intfdRglrSih elwOaelnelt togycahaeDenticulate Denticulate lustrous Non-clathrate, Ovate-lanceolate clathrate Strongly Denticulate Ovate-lanceolate Yellow clathrate, Weakly Non-clathrate, round Yellow Irregularly clathrate Strongly Yellow Ovate-lanceolate Round None Yellow Yellow Ovate-lanceolate- Slight Scattered Yellow Deep Moderate Regular Simple Pinnatifid Slight Regular Regular None Pinnatifid Pinnatifid Scattered Coenosorus Pinnatifid Simple Irregularly .powellii M. ygot ai tretilo epi- or (terrestrial habit growth by Results iscinSrlarnmn oa meso pr oo hzm cl hp hzm cl color scale Rhizome shape scale Rhizome color Spore immersion Soral arrangment Soral Dissection pinnatifid 0 u ifr nlmn etr from texture lamina in differs but , 5 xnand exon Microsorum 0% MPBS 100%; .Teukonplant unknown The ). ctee oet oeaeCerOaelnelt ltrt Denticulate Clathrate Ovate-lanceolate Clear moderate to None Scattered nlmn etr,btis but texture, lamina in .grossum M. trnF 5 ,adcnb distin- be can and ), sntmonophlyetic not is oiimre,aailyvsbebtntfrigrie up,o up eysih;Moderate slight; very bumps or bumps, raised forming not but visible adaxially immersed, Sori ihu the without , 5 .powellii M. .gosm M. grossum, M. 0% BIPP 100%; , .powellii M. trnL trnL YTMTCBTN Vlm 43 [Volume BOTANY SYSTEMATIC .com- M. – – and ) F F trnL “ se- M. e 5 ” , e aeoi,adAsrla ntelatter. the in Australia) and Caledonia, New h Lpyoeisi is n .A nteplastid the in As 6. and 5 Figs. in phylogenies on based ML trees summary the present strongly we taxa. any so lacked topologies, and 13 similar conflicting were the analyses representing of BI Results and specimens 3. MP, Table ML, in 20 summarized are from statistics Alignment total in quences omr n h eaotri ld (including named clade taxa other lecanopteroid and the the of and case the in former, clade lepisoroid the micros- within nested Islands and related, Society two other ferns, The oroid resolved. well not are BIPP 50%; s. s. spectrum grossum (MLBS other each membranifolium M. (MLBS Noot. oasrnl upre ld loincluding also clade lopendria supported 4). strongly (Fig. a to tree plastid punctatum, the detail. in in those discuss tions not do we so study, this the of not and sampled focus studies previously previous with other consistent 2010a, of broadly al. were Relationships taxa et 2017). Wang al. 2008; et al. Wei et b; Kreier b; 2004a, al. et (Schneider ot (MLBS Noot. and Hoshiz., & Sm. A.R. Parris, unwal upre ssse to sister as supported weakly turn ld of clade identified with matches Maupiti of those match sequences MPBS 53%; n aii,acliae pcmno nnw rgn an origin, unknown of rubidum specimen cultivated Moorea, unidentified (Huahine, a Islands Tahiti), Society and the Pou), (Ua Marquesas irsrmlucidum Microsorum containing clade BIPP 100%; 5 oit sad irsri en cuyarneo posi- of range a occupy ferns microsoroid Islands Society ula Phylogeny Nuclear oiimre,frigcnpcosyrie up o2 to bumps raised conspicuously forming immersed, Sori ” ld es rire l 20)(MLBS (2008) al. et Kreier sensu clade irsrmmusifolium Microsorum – Kne oe.fo aa (MLBS Japan from Copel. (Kunze) ld,and clade, punctatum .membranifolium M. Br.. Copel., (Burm.f.) Microsorum 5 and , 5 .8,btrltosisbtenteefu groups four these between relationships but 0.98), .spicatus L. Microsorum 5 0;BIPP 50%; 1.00). 5 linear M. 8;MPBS 98%; 0;MPBS 80%; irsrmcuspidatum Microsorum – 3 5 Leptochilus ld and clade Rx. oe. and Copel., (Roxb.) commutatum irsrmspectrum Microsorum maximum 8;MPBS 98%; ln rmMoe r etdwti a within nested are Moorea from plant — Microsorum .punctatum M. and epoue 9nwncerse- nuclear new 69 produced We irsrmthailandicum Microsorum 5 pce rmLo,and Laos, from species .grossum M. .4.Most 0.64). 5 .powellii M. irsrmpapuanum Microsorum nldn cesosfo the from accessions including ltrt Denticulate Clathrate .spectrum M. oehrfr the form together Copel., 3;BIPP 93%; r lsl eae n belong and related closely are 5 h center the light-brown center in dark clade, otyfo e Zealand, New from mostly 5 6;BIPP 86%; .commutatum M. irsrmgrossum Microsorum u n pcmnfrom specimen one but , 7;BIPP 87%; eune fteun- the of Sequences . irsrmwhiteheadii Microsorum M. irsrmhainanense Microsorum membranifolium r oedistantly more are , 5 3 snse ihna within nested is ld r itrto sister are clade DDn Tagawa, (D.Don) 5 .0,wihi in is which 1.00), 5 maximum irsrmsco- Microsorum 0% MPBS 100%; 4;MPBS 64%; 5 “ 5 microsoroid oked& Booknerd Lecanopteris Microsorum .0.The 1.00). – .0.The 1.00). Entire Entire Entire hzm scale Rhizome mm. 5 (MLBS margin (Baker) plastid clade, 5 , Sori M. , 5 5 5 Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 2018]

Table 2. Quantitative morphological traits of microsoroid ferns from the Society Islands, French Polynesia. Data shown as mean 6 SD (range, sample size).

Stipe length (cm) Frond length (cm) Frond width (cm) Rhizome diam (mm) No. pinna pairs L. spicatus 2.8 6 1.5 (0–6.4, n 5 23) 33.2 6 8.7 (19.6–51.4, n 5 23) 1.6 6 0.5 (0.8–2.6, n 5 23) 4.1 6 1.3 (2.5–8, n 5 23) 401 0 6 0(0–0, n 5 23) POLYNESIA FRENCH FROM MICROSORUM HYBRID NEW AL.: ET NITTA M. commutatum 36.7 6 14.8 (10.4–74, n 5 17) 97.7 6 31.3 (39.7–150, n 5 17) 24 6 5.9 (15.1–36, n 5 17) 7.1 6 3(5.1–17, n 5 17) 14 6 4.5 (6–20, n 5 16) M. grossum 27.8 6 13.5 (9.6–61.4, n 5 30) 63.4 6 21.5 (31.4–109.1, n 5 30) 20 6 6.2 (2.1–29.2, n 5 30) 5.3 6 1.4 (3.4–9.2, n 5 30) 6.3 6 3.5 (0–14, n 5 30) M. membranifolium 69.4 6 20.1 (50–105, n 5 7) 170.1 6 61.8 (111–278.8, n 5 7) 42.5 6 15.2 (25–64.8, n 5 7) 9 6 3.5 (6–15, n 5 7) 6.6 6 8.6 (0–20, n 5 7) M. powellii 19.8 6 10.6 (4.8–42, n 5 16) 47.6 6 19.7 (18.5–84.2, n 5 16) 18.6 6 5.5 (9.3–27.3, n 5 16) 5.9 6 1.6 (4–9.7, n 5 15) 7.2 6 3.4 (3–13, n 5 16) M. punctatum 0 6 0(0–0, n 5 25) 72.1 6 16.7 (40.5–100.2, n 5 25) 7.4 6 1.2 (5–10.7, n 5 25) 6 6 1.2 (3.4–9, n 5 25) 0 6 0(0–0, n 5 25) M. 3 maximum 10.9 6 9.8 (0.6–38, n 5 16) 92.8 6 33.6 (27.4–155, n 5 16) 17.1 6 8.7 (3.7–29, n 5 16) 6.8 6 1.1 (5.2–8.8, n 5 16) 2.6 6 2.9 (0–10, n 5 16) M. 3 tohieaense 25.6 6 6.8 (20.8–30.4, n 5 2) 55.5 6 2.4 (53.9–57.2, n 5 2) 16 6 0.5 (15.6–16.3, n 5 2) 4.8 6 0.4 (4.5–5, n 5 2) 6 6 0(6–6, n 5 2) .punctatum M. in hybrids, putative cluding fsqecsotie from obtained sequences of hybrid- of implications of the taxonomy discuss the and for ization below these of turn each progenitor in describe its cases We elucidating investigation. although further requires origin, taxa hybrid of be also grossum M. hs ohapa nteacsinof accession the in appear both these hc oehraesse to sister are together which of copies two the between varies nodes internal two and into species commutatum, non-hybrid M. including the one clades: the split in main These tree plastid trees. the in nuclear observed divergences deeper same BIPP nih notedtiso t rgn.W eetda exact an detected We origins. from its of set of one between details match the into insight that evidence genetic M. first the provide we Furthermore, ment). MPBS and that to alleles closely distinct multiple match has Moorea from plant unknown ora.Frhroe utpedvretallswr also were alleles in- for divergent recovered both multiple in Furthermore, origins Moorea). hybrid for (i.e. hypotheses stances the support data short membranifolium ecieas hybrid describe a as Moorea from plant between sup- unknown the strongly of status analyses the port phylogenetic and morphological our for Maupiti short gapCp oti w itntallsfor alleles distinct two contain to MPBS hc oehraesse to sister MPBS are together which BIPP a 100%, ( as ferns monophyletic microsoroid are the whole that show data nuclear phylogeny, ac xcl ihtoeof those with exactly match per ob suoeebsdo 3b eeini xn9. exon in deletion bp 53 a on based additional pseudogene a be or to appears error, of PCR-introduced duplications populations, (e.g. diploid well as species spicatus non-hybrid other some otecaecontaining clade the in to phylogenies: two the between varies oit sad,sgetn that suggesting Islands, Society the MPBS Microsorum Microsorum atrso yrdzto nMicrosorum in Hybridization of Patterns .commutatum M. 3 long punctatum .spicatus L. 5 M. maximum and 5 5 5 5 u ti nla fteerpeetallcdvriyin diversity allelic represent these if unclear is it but ) .0,but 1.00), rvr lsl ( closely very or ) 5;BIPP 85%; 9;BIPP 99%; 3 6;BIPP 96%; .membranifolium M. 2;BIPP 92%; long n htteHwia endemic Hawaiian the that and , aC short gapCp only). tohieaense 5 Microsorum M. .spectrum M. – 0.81; 3 (MLBS n n itrt h ld including clade the to sister one and ) commutatum side yrdbetween hybrid a indeed is gapCp Fg.5 ) oee,tesrcueo some of structure the However, 6). 5, (Figs. and 3 3 n h te including other the and .membranifolium M. maximum Microsorum irsrmgrossum Microsorum (MLBS tohieaense 5 aC short gapCp 5 maximum 5 5 .ellipticus L. .0,weesin whereas 1.00), One . ihtoeof those with .0.In 1.00). .0.Tepaeetof placement The 1.00). utpeallswr eoee within recovered were alleles Multiple . 5 0.94). aC short gapCp Discussion aC short gapCp 3 n itrto sister one , .membranifolium M. 5;MPBS 85%; 5 aC long gapCp .grossum M. – aC short gapCp a utpedsic lee that alleles distinct multiple has tohieaense aC short gapCp 9;MPBS 99%; Microsorum grossum .grossum M. and .commutatum M. M. snt edtce oeo the of some detected We not. is aC short gapCp n h nnw ln from plant unknown the and .punctatum M. aC long gapCp — MLBS .commutatum M. var. h hlgntcplacement phylogenetic The 3 .commutatum M. )to .commutatum M. tohieaense and n xcl ( exactly and MLBS ld (MLBS clade pothifolius seTxnmcTreat- Taxonomic (see 5 and aC long gapCp 5 .commutatum M. and , .commutatum M. aC short gapCp M. . leefrom allele .grossum M. 5 long , 9,MPBS 99%, , 0;BIPP 70%; and .punctatum M. .grossum M. .membranifolium M. 7;BIPP 97%; .punctatum M. .punctatum M. 3 .punctatum M. 5 .spectrum M. lee recovered alleles — (MLBS (MLBS aC long gapCp (MLBS maximum .powellii M. rvdsmore provides .powellii M. 1,MPBS 81%, h eut of results The osre in (observed rmnearby from ti itrto sister is it , hc we which , aC short gapCp gapCp ti sister is it , .spicatus L. ssse to sister is 5 rmthe from 5 5 5 5 5 5 ssister is These . ( gapCp 100%, 100%; ,and 0.98), 1.00), and , from The . Ex- . 99%; 98%; 87%; each may also and , M. 5 L. Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. inadsbeunl eaeetrae rmteSociety is possibility the Another Marquesas. that from the in extirpated survived but became Islands, subsequently and tion of haplotype plastid distributed membranifolium widely more a of been have haplotype plastid of single sequences branifolium a plastid indicating our identical, However, sample. membranifolium to failed from sequences of itnus ewe hs scenarios. these to needed between is Asia distinguish and region Pacific by South the throughout from islands Pacific of branifolium colonizations independent (an the to taxa tohieaense sister Asian Japan) two from of position unidentified phylogenetic The Islands. by dispersal spore location type the near Islands of Society the in of present that haplotypes currently possible plastid is additional it are First, there explanations. pattern disjunct conceivable This several 4). 1A, has (Figs. distant km 1380 ca. Islands), of accessions tohieaense of membranifolium close of a set other observed of the and we genes Furthermore, plastid between hybrid. match the of formation 402 h yepplto of population type the ro niae cclrprpyi.Nt la prs .Csa cl.Saeas A Scalebars: scale. Costal F. spores. clear Note paraphysis. acicular indicates Arrow G) D: (GH); Fig. M. M. .membranifolium M. M. M. 3 3 .Poorpsof Photographs 2. 3 3 it 1040 Nitta tohieaense tohieaense tohieaense. lsi eune r lal oecoeyrltdto related closely more clearly are sequences plastid tohieaense ld Fg ) niae h osblt fmultiple of possibility the indicates 4), (Fig. clade nteSceyIlns lentvl,teecould there Alternatively, Islands. Society the in rmAi.Adtoa eunigo specimens of sequencing Additional Asia. from Microsorum .membranifolium M. stemte,and mother, the is nteps htcnrbtdt yrdforma- hybrid to contributed that past the in M. rmMoe,Tht,adHaieaeall are Huahine and Tahiti, Moorea, from G) htgah yJ .Nitta. H. J. by Photographs (GH). htcnrbtdt yrdfrainta we that formation hybrid to contributed that with 3 oee,nn ftepatdo nuclear or plastid the of none However, ol etersl frcn long-distance recent of result the be could .membranifolium M. tohieaense rmteSceyIlns Rather, Islands. Society the from Microsorum M. .membranifolium M. ceso rmLo and Laos from accession 3 .membranifolium M. tohieaense ac xcl ihsequences with exactly match 3 .commutatum M. rmU o (Marquesas Pou Ua from tohieaense ietycnrbtdto contributed directly rmteMarquesas the from ugsigthat suggesting , .Woepati iu .Aailsraeo aia .Rioei iu .Rioesae .Sorus. E. scale. Rhizome D. situ. in Rhizome C. lamina. of surface Abaxial B. situ. in plant Whole A. . .membranifolium M. aPou Ua gapCp stefather, the is .rubidum M. YTMTCBTN Vlm 43 [Volume BOTANY SYSTEMATIC .mem- M. .mem- M. – alleles M. M. M. M. M. 3 3 osdrdt eahbi ntebsso t intermediate its of basis the on hybrid a be to considered rdc yrdsoohtsb rsig(bhr ta.2009). repeatedly al. and et two (Ebihara site the crossing this by of at sporophytes hybrid gametophytes together produce exist long-lived may that species possible parent also so is 1971), Kaur and it (Nayar ca- branching by gametophytes growth clonal have of pable ferns Microsoroid 2011). (Barrington short gapCp including branifolium erdcini siae ohv upre trl hybrid sterile a supported of vegetative have population con- to such estimated is rhizomes; is possibility long-creeping reproduction One of maintained. became growth is population tinuous it type of how the or when locality) established, unknown is type It the apogamy. from only tohieaense known (currently this if complex. determine species to a needed is is taxon analysis level ploidy including of sampling intensive More cluding lal stepatddt,btnihrd hycnrdc it: contradict they the from do derived in not neither alleles but the considering data, only as plastid pattern the disjunct the as show clearly not do data nuclear The diploids. commutatum of those whereas n hybrids. and branifolium iaigete looyliyo nrseii variation, intraspecific in- or alleles, multiple allopolyploidy separate either to dicating cloning required Moorea Microsorum h la,ocsoal isae prsadlmtdrange limited and spores misshapen occasionally clear, The nte atrta hudb osdrdi that is considered be should that factor Another 5 5cm;B aC long gapCp .membranifolium M. ugs hti sicpbeo eulrpouto or reproduction sexual of incapable is it that suggest .membranifolium M. – tefmyb ope nldn utpespecies multiple including complex a be may itself rmU o,wihtgte etwti clade a within nest together which Pou, Ua from C tree, ol edrc-eune,cnitn ihsexual with consistent direct-sequenced, be could 5 Osmunda GapCp 2cm;D 3 .membranifolium M. M. tree, maximum 3 cesosof accessions – E tohieaense nWs igna S o a 10yr 1100 ca. for USA Virginia, West in M. 5 250 rmMoe n aPu(i.5). (Fig. Pou Ua and Moorea from .membranifolium M. 3 — m tohieaense rmMoe Fg ) nthe In 6). (Fig. Moorea from ;F m; hsseiswspreviously was species This snse ihnacaein- clade a within nested is 5 rmU o swl as well as Pou Ua from .membranifolium M. 100 m ssse to sister is .A m. .commutatum M. – cosisrange its across ,E C, – F: .mem- M. .mem- M. it 3929 Nitta M. from M. 3 , Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 2018] opooybetween morphology A Tbe1.Rcnl,Ntae l 21)dtce matching detected (2017) al. et sori ( sequences Nitta plastid scattered Recently, irregularly 1). and (Table lobes shaped irregularly have of laminae grossum 5 Fig. 1cm;B .Ilsrto of Illustration 3. del intfdlmne Cpln 92.The 1932). (Copeland laminae) pinnatifid (deeply 5 M. 5cm;C 3 maximum rbcL 5 Microsorum .punctatum M. 500 IT TA. E YRDMCOOU RMFEC OYEI 403 POLYNESIA FRENCH FROM MICROSORUM HYBRID NEW AL.: ET NITTA and m .A B: A, m. r oeo essml,btoften but simple, less or more are trnHpsbA 3 tohieaense it 3929 Nitta sml aia)and laminae) (simple ewe cesosof accessions between ) .Dti faailsraesoigarneeto oiadvis .Hbt .Rioesae Scalebars: scale. Rhizome C. Habit. B. veins. and sori of arrangement showing surface abaxial of Detail A. . G) C: (GH). it 1040 Nitta M. G) lutain yC .Nitta. C. C. by Illustrations (GH). h yohssthat hypothesis the neetnl,o h orsmlsicue noranalysis, our in included species. samples two four these between the hybrid of a Interestingly, as status its confirming nuclear maximum of analysis phylogenetic Our M. 3 maximum otisallsfo ohpttv aet,further parents, putative both from alleles contains and .grossum M. .grossum M. stemte of mother the is rmMoe,cnitn with consistent Moorea, from gapCp hw that shows M. 3 maximum M. 3 . Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. oee n additional one covered matching sequences plastid 404 ( he fo uhn,Moe,adTht)hv lsi se- plastid have Tahiti) and matching Moorea, quences Huahine, (from three rbcL trnL rbcL aC short gapCp long gapCp fti aito sdet t yrdoiis u nuclear Our origins. of hybrid specimen its one to pentadactylum includes due only is sampling variation some that this possible is of It 2013). Ranker and Vernon 2003; Palmer (var. varieties allopolyploid. two as an treated be may it spectrum that suggests data, to related closely membranifolium spectrum M. membranifolium M. mutatum gapCp endemic Hawaiian spectrum that M. suggestions previous any of unaware with and 2013). al. 2013), et al. et (Rothfels in observed been have ferns Adiantum in of duplications lineages apparent particular Similar within 6). 5, (Figs. these of and duplication, of result of copies tinct 2016). Balukjian punctatum and (Hembry insects, plants and including and organisms birds, Bora, diverse in observed Bora been in- (Moorea have Raiatea, splits is Windward (Huahine, size Phylogeographic and sample Maupiti) Leeward hypotheses. our the such but between test 2014), al. to et sufficient (Sigel bias pro- hybridization observed the ecological structuring we and/or in and involved (Fosberg Geographical be others could 1B). cesses the Fig. than 1987; flora Sachet diverse drier less m), 2016). (380 and altitude al. climate, maximum main lower et the a of with Yamada smallest Islands, and Society 2014; westernmost the al. is 2000; genotype, et plastid al. sole the Testo et of site Xiang 2013; the 1998b; Maupiti, al. al. et et (Vogel Zhang taxa fern hybrid of other parentage in bias a be maximum to appears there Thus, shown). matching sequences plastid with analysis taining r2 or 2 either of count chromosome parents. tetraploid putative these of either for ta.21) h omnacso of ancestor common the 2010), al. et n htfrsacaewith clade a forms that one eune of sequences var. iet 34 Vinette Table irsrmspectrum Microsorum – n 1 F pothifolius 5 trnL lee in alleles .cuspidatum M. .Ainetsaitc ygn ein ecn isn aacluae snme fclsi h lgmn htaentA ,T rGoto total. of out G or T, C, A, not are that alignment the in cells of number as calculated data missing Percent region. gene by statistics Alignment 3. 4 (L¨ 144 osre nboth in (observed hnmnnta a enosre nseveral in observed been has that phenomenon a , samrhlgclyvral ao,adhsbeen has and taxon, variable morphologically a is – Rtfl n cutpl 2014), Schuettpelz and (Rothfels F C o nlddi h urn phylogenetic current the in included not UC) , and a eo yrdoii.W eetdtodistinct two detected We origin. hybrid of be may ,btadtoa apigicuigmultiple including sampling additional but ), so yrdoii ihamte rmthe from mother a with origin hybrid of is .spectrum M. v ta.17) hc,i obnto ihour with combination in which, 1977), al. et ove osre in (observed – aC long gapCp .grossum M. .spectrum M. cuspidatum .commutatum, M. o cesosN.idvdasN.tx o hrces%msigdt o % asmn-nomtv characters parsimony-informative (%) No. data missing % characters No. taxa No. individuals No. accessions No. .grossum M. ld Fg ) ti hrfr osbethat possible therefore is It 4). (Fig. clade 4 4 4 1011 6 (22.8%) 269 1.1% 1180 149 240 240 3 3 4 9 63 1 (41.8%) 416 46.3% 995 148 237 237 4 4 4 152.%65(31.5%) 685 22.7% 2175 149 242 242 and 82 3772.%30(39.1%) 300 (29.6%) 337 21.8% 25.3% 767 1138 13 12 23 21 38 32 M. 3 .hainanense M. aC short gapCp M. — M. eense ihnacaecon- clade a within nested were .membranifolium M. – and aC short gapCp maximum hmevsec aetodis- two have each themselves n lsl eae to related closely one , spectrum Leptochilus Unlike .punctatum M. 3 hl n fo apt)has Maupiti) (from one while , 3 maximum maximum short nLnseca (Rothfels Lindsaeaceae in u elc xc matches exact lack we but irsrmspectrum Microsorum aii oit Islands Society Tahiti) , M. and Culcita n var. and pcmnfo Moorea from specimen htapa ob the be to appear that n hc ssse othe to sister is which , 3 5 ny i.5.Plastid 5). Fig. only; .grossum M. ld n father a and clade long ihan with a neie both inherited has 4 Wge 1963) (Wagner 148 .spectrum M. maximum Fg ) ere- We 4). (Fig. Astrolepis and and is ,6,and 6), 5, Figs. ; pentadactylum Microsorum Microsorum .grossum M. Plagiogyria .ellipticus L. YTMTCBTN Vlm 43 [Volume BOTANY SYSTEMATIC dt not (data eare we , .com- M. gapCp (Beck M. a a has (var. 3 ; hthbis( hybrids that species papuanum the for plied pce including species restrict punctatum to (e.g. is alternative genera An warranted. monophyletic in- would distinct it Lecanopteris name, multiple this in- sinking with be species volve all may If include here studies. to taxonomic observe expanded future genus, we the for hybridization of formative revision of taxonomic patterns a is for the sampling allow our Although to data. sufficient nuclear studies not with this time demonstrate first furthermore previous the and for 2008), al. with et Kreier b; 2004a, agree that analysis showing phylogenetic our en Sgle l 04 et ta.21)adgieappropriate guide and 2014) delimitation. al. et generic Testo 2014; al. microsoroid et between (Sigel be- flow ferns gene overlap to barriers into niche of insight maintenance of further the provide degree also and should the taxa sperm progenitor and tween antheridiogens, sizes, hy- neck of of archegonia dynamics effect the including into bridization, studies of distinctness Detailed of the genera. clarify Inclusion help group. proposed may hybrid this studies of future in in morphology species taxa of these or basis hybrids the putative on Nooteboom origin other ferns. nine microsoroid lists in (1997) hybridization of evidence hybrids. tochilus whiteheadii ofr hte hsseisi nedo yrdoii,adif occur. and parents origin, its hybrid where of indeed so, is species to this needed whether is confirm elsewhere and Hawaii in not sampling is Additional far. Pacific it other that the of indicate any analyses from derived phylogenetic our of and results 2013), Ranker the and Vernon 2003; (Palmer Hawaii in curring ude21;Rtfl ta.21;Shatbr ta.2018). al. et and Schwartsburd (Ranker 2015; rare al. Were et extremely are Rothfels they 2015; but Sundue of, unheard genera not fern the are between of we events affinity Hybridization genetic events involved. the species indicating hybridization again clade, other this The within (2008). ( al. detect et Kreier of of pre- species other are absolute and comprising species clade lack these a of within and all nested Furthermore, similar, these barriers. that genetically mating indicate zygotic father) are or clades mother the two as acting of capable grossum aito screae ihgntcdvriyi hstaxon. this in morphological diversity the genetic of with Furthermore, any correlated if is clarify variation help may morphotypes u td stefrtt u nweg oso genetic show to knowledge our to first the is study Our aooi mlctosfrMicrosorum for Implications Taxonomic Leptochilus Microsorum twudrsl na es w diinlintergeneric additional two least at in result would it , M. cu rqetyadrcpoal ie ihrseisis species either (i.e. reciprocally and frequently occur . Kee ta.20) oee,orrslsshowing results our However, 2008). al. et (Kreier tetp pce of species type (the , Phymatosorus 3 Lepisorus M. .spectrum M. orsodn othe to corresponding , Microsorum tohieaense 3 pi orecognize to split .musifolium M. maximum nosnnm,wihde o seem not does which synonymy, into ) .grossum M. ,possibly (type steol native only the is splpyei Shedre al. et (Schneider polyphyletic is between ) Microsorum .scolopendria M. , , .thailandicum M. .spectrum M. Microsorum .scolopendria M. “ Phymatosorus irsri .s clade s. s. Microsoroid Microsorum .punctatum M. n lsl related closely and ) Microsorum — Microsorum ol eap- be could ) h eut of results The ape thus sampled )alsooccur Microsorum oonly to and , and , and and were Lep- oc- M. M. M. M. ” Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 2018] aa aaocrigi oit sad n aaiclrdb ao;tx o curn nteeaesadotrusi black. in outgroups and areas these in occurring not taxa taxon; by colored Hawaii and Islands Society in occurring Taxa taxa. ldswti eeata ontcnanaySceyIlnsseisadhv enclasdfrpotn;nme fclasdseisidctdin indicated species collapsed of number plotting; for collapsed been have ( branchlengths and large short or species Extremely genera clade. Islands monophyletic collapsed indicate Society the Diamonds within any specimens. divergence contain multiple first with at not taxa placed do for Diamonds shown parenthesis. that code genera individual specimen within or clades / and information Locality bold. usiuin e ie missing * site. per substitutions LC clade; membranaceoid probability olpe ozr.Srnl upre oe mxmmlklho bootstrap likelihood (maximum nodes supported Strongly zero. to collapsed Fig. .Mxmmlklho hlgntcte nerduigpatdmres( markers plastid using inferred tree phylogenetic Maximum-likelihood 4. . .5 niae ytikndlns brvain tndsidct ao ldsfloigKee ta.(08:M (2008): al. et Kreier following clades major indicate nodes at Abbreviations lines. thickened by indicated 0.95) 5 IT TA. E YRDMCOOU RMFEC OYEI 405 POLYNESIA FRENCH FROM MICROSORUM HYBRID NEW AL.: ET NITTA eaotri ld;LP clade; lecanopteroid rbcL . † missing trnL – F . 5 eioodcae cM clade; lepisoroid . 5;mxmmprioybootstrap parsimony maximum 95%; 5 rbcL oemcoood;MI microsoroids; core and trnL – F nldn oit sad irsri en n related and ferns microsoroid Islands Society including ) 5 , irsri s . ld.Saea indicates Scalebar clade. s.) (s. microsoroid 1 3 10 . 2 5 5;Bysa neec posterior inference Bayesian 95%; usiuin e ie aebeen have site) per substitutions 5 Microsorum irsri en;ME ferns; microsoroid 3 tohieaense in 5 Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 406 otecereiec ercvrfrhbi rgn of origins hybrid for recover we maximum evidence clear the to .grossum topologies two M. the conflicting between species supported, non-hybrid for statistically observed group. this We in evolution genome nuclear of nature complicated Fig. vlto fNcerGnmsi Microsorum in Genomes Nuclear of Evolution .(Continued) 4. and and M. .punctatum M. 3 tohieaense u aae lorvasthe reveals also dataset our , omacae(MLBS clade a form gapCp — phylogenies: naddition In YTMTCBTN Vlm 43 [Volume BOTANY SYSTEMATIC 5 M. 85%; 3 MPBS mutatum aC short gapCp commutatum .0,wihi ntr itrto sister turn MPBS in is which 1.00), Fg ) utemr,tepsto of position the Furthermore, 6). (Fig. 5 5 (MLBS 0;BIPP 70%; 6;BIPP 96%; hlgn Fg ) whereas 5), (Fig. phylogeny omacae(MLBS clade a form 5 9;MPBS 99%; 5 5 .8,wihi hnsse to sister then is which 0.98), .0 nthe in 1.00) .punctatum M. 5 5 9;MPBS 99%; 9;BIPP 99%; aC long gapCp .powellii M. .grossum M. (MLBS 5 5 7;BIPP 97%; .0 nthe in 1.00) phylogeny lovaries also .com- M. 5 and 98%; M. 5 Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. eaae ysahs ausls hn5%(LS PS r05(IP niae ihds.Eteeysotbaclnts( branchlengths short Extremely dash. with indicated (BIPP) nod 0.5 at or shown MPBS) values (MLBS, (BIPP) 50% probability than posterior less inference values Bayesian slashes; and by (MPBS), separated bootstrap parsimony maximum (MLBS), bootstrap likelihood Maximum 2018] oeis(is ,5,btsse to sister but 5), and 4, plastid (Figs. the logenies in groups more related other distantly amongst appearing phylogenies, nuclear between punctatum curn nSceyIlnsadHwi ooe ytxn aantocrigi hs ra n ugop nblack. in outgroups and areas these in occurring not taxa taxon; by colored Hawaii and Islands Society in occurring aebe olpe ozr.Saea niae usiuin e site. per substitutions indicates Scalebar zero. to collapsed been have Fig. .Mxmmlklho hlgntcte nerduignuclear using inferred tree phylogenetic Maximum-likelihood 5. , .grossum M. and , IT TA. E YRDMCOOU RMFEC OYEI 407 POLYNESIA FRENCH FROM MICROSORUM HYBRID NEW AL.: ET NITTA .commutatum M. “ core Microsorum aC short gapCp MLBS ; ” 5 (i.e. 100%; phy- M. aC short gapCp hrfr osbeta yrdzto a lyda important an is played It has hybridization introgression. that or possible sorting therefore lineage indicate incomplete may nodes of internal processes at genes nuclear between conflict MPBS nldn oit sad irsri en n eae aa Taxa taxa. related and ferns microsoroid Islands Society including 5 2;BIPP 92%; 5 .4 nthe in 0.94) aC long gapCp Microsorum , 1 3 10 2 5 usiuin e site) per substitutions 3 re(i.6.Such 6). (Fig. tree tohieaense nbold. in es Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 408 comdt ag ubro ape oi plcto of Application cannot loci. for / methods and sequencing samples DNA next-generation labor-intensive of developed recently is number scales. large cloning, time a long here, accommodate over used ferns microsoroid method of The evolution the in role curn nSceyIlnsadHwi ooe ytxn aantocrigi hs ra n ugop nblack. in outgroups ( and branchlengths short areas Extremely dash. these with indicated in (BIPP) nod 0.5 at or occurring shown MPBS) values (MLBS, not (BIPP) 50% probability than taxa posterior less inference values taxon; Bayesian slashes; and by by (MPBS), separated bootstrap colored parsimony maximum Hawaii (MLBS), bootstrap and likelihood Maximum Islands Society in occurring aebe olpe ozr.Saea niae usiuin e site. per substitutions indicates Scalebar zero. to collapsed been have Fig. .Mxmmlklho hlgntcte nerduignuclear using inferred tree phylogenetic Maximum-likelihood 6. YTMTCBTN Vlm 43 [Volume BOTANY SYSTEMATIC aC long gapCp nih notecmlxeouinr itr fti group. this of history evolutionary combined complex loci, the better into unlinked provide insight should of 2009), number (Edwards methods larger species-tree with much a for produce can datasets that 2017) al. et (Rothfels complexes species polyploid nldn oit sad irsri en n eae aa Taxa taxa. related and ferns microsoroid Islands Society including Microsorum , 1 3 10 2 5 usiuin e site) per substitutions 3 tohieaense nbold. in es Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. lcd lgtyimre,apaigsihl asdadax- paraphyses. raised uniseriate with slightly ially, appearing immersed, slightly irregularly placed, occasionally costa; ro margins, pinna the single and costae a to between in parallel anastomosing. located areolae variously round, large veins of smaller row one 5 forming veins acuminate. lateral apex ovate, clathrate, cainlymissh occasionally scat- 0.5 few scales with costal costae stramineous; and slightly scales, rachises evident, tered base; more the veins at primary narrowing with pinnae, 13 lateral pinnae apical entire; cending, ru,wt lgtyset(omrn rgac;pinnae fragrance; (coumarin) sweet slightly 9 a gla- surfaces with membranaceous, brous, subconform, apically natifid, h ae taieu;sae iia otoeo rhizome. on those to similar scales Laminae stramineous; base, the 20 oad h agn,mrisetr;pylpdadistinct. phyllopodia lighter entire; margins, margins and Fronds margins, center between the brown towards dark center, in green; pale 1 scaly, scales densely rhizome to moderately creeping, long wide, in ie hnltrloe v.mr rls ofr apical conform less or in more (vs. pinnae ones lateral than in wider m 2 pinna to fronds vs. m 1 than less in membranifolium co- scattered (vs. laminae (vs. membranaceous with in but riaceous habit, and size in te n ok ufc,s hyaecniee ohterrestrial both epipetric. considered are and they so surface, rocky a and on steep French but Moorea, terrestrially, Tohiea, growing observed Mt. were of Plants slope Polynesia. the on m 400 ca. at located Microsorum 2018] nw rmasnl ouainls hnc.1 m 10 ca. than less population single a from known iue ,3. 2, Figures .Soagaarne ndsic oi o etitdt h pclsget pclsgetntarpl arwd hzm ogcepn.2 long-creeping. rhizome narrowed; abruptly not segment apical segment; apical the to restricted not sori, distinct in arranged Sporangia 1. .Soagacvrn niefriesraeo rn ie omn onsrs,cnetae taia emn;aia emn butynarrowe abruptly segment apical segment; apical at concentrated coenosorus), a forming (i.e. frond of surface fertile entire covering Sporangia 1. – – lnstretilo lithophytic. or terrestrial Plants iia to Similar 11 clg n Distribution and Ecology .Lmnesml,bs rnaet bue tp beto ral lt ln eryetr egh ...... 3 . alate. . narrowly . only . . or . not . present, . stipe . cuneate; . . base length. pinnatifid, entire deeply nearly to along lobed alate irregularly broadly least or at absent usually stipe simple, obtuse; rarely to Laminae truncate base simple, 2. Laminae 2. 0c og lbosecp o e ctee clsnear scales scattered few a for except glabrous long, cm 30 UC!). 2012, Jul P hzm short-creeping. rhizome 3 OLYNESIA .Sr oeo esi eua ie ewe oteadmris 5 margins. and costae between lines regular in less or more Sori 4 scattered. 3. Sori 3. 50 . m 5 cm, 2.5 .Lmneirglrylbd oi1 sori lobed; irregularly Laminae 4. .Lmnedel intfd sori pinnatifid; deeply Laminae 4. – 32 .membranifolium M. 0c og e 2 set long, cm 60 .grossum M. – irsrmgrossum Microsorum 3 .Lmnemmrncost ucraeu;scnayvisa es atyvsbet eydsic.6 distinct. very to visible partly least at veins secondary subcoriaceous; to membranaceous obscured. Laminae veins secondary coriaceous; 5. Laminae 5. ..Nta1040 Nitta J.H. 37 oit sad:Moe,M.The,33m 12 m, 393 Tohiea, Mt. Moorea, Islands: Society . ntxue u ifr nsalrsz (fronds size smaller in differs but texture, in tohieaense – – .Pat ersra;lmnemmrncos 7 membranaceous. laminae terrestrial; Plants 6. .Pat ppyi;lmnesboicos . subcoriaceous. laminae epiphytic; Plants 6. 3 maat itnt oncigveins connecting distinct; apart, mm 6 . mwd,rud ett,cahae light clathrate, peltate, round, wide, mm 1.5 – aooi Treatment Taxonomic .commutatum M. ()pis ierlnelt,sihl as- slightly linear-lanceolate, pairs, 6(7) .Fod o2( to Fronds 7. .Fronds 7. 15 pn la,peual o fertile. not presumably clear, apen, – n oimr rls nrglrrows regular in less or more sori and ) 7c,oaelnelt,del pin- deeply ovate-lanceolate, cm, 17 IT TA. E YRDMCOOU RMFEC OYEI 409 POLYNESIA FRENCH FROM MICROSORUM HYBRID NEW AL.: ET NITTA e oteMcooodFrso h oit sad,Fec Polynesia French Islands, Society the of Ferns Microsoroid the to Key 2( dxal...... adaxially...... – – ..it,hb o.T nov. hyb. J.H.Nitta, 5)mmhighadaxially...... hltp:G! stps ! PAP!, P!, isotypes: GH!; (holotype: maat monomorphic. apart, cm 5 ). , — o oeie w rows) two sometimes (or w ;aia in ie hnltrlpna;sr lgtyimre,ol omn lgtyrie bumps raised slightly forming only immersed, slightly sori pinnae; lateral than wider pinna apical m; 1 .membranifolium M. Microsorum and – – Spores )m pclpnamr rls aesz sltrlpna;sr epyimre,frigrie up ca. bumps raised forming immersed, deeply sori pinnae; lateral as size same less or more pinna apical m; 3) 19 ;smlrto similar ); Rhizomes Veins irsrmcommutatum Microsorum 3 , – 3 mda,smtmsfsn together. fusing sometimes diam, mm 2 29 – 1mmdiam,notfusing...... 4cm,largerthan Sori – eiuae main reticulate; 3 44 3 4.5 3 0.1 tohieaense YPE 1.5 Microsorum n apical and ) 10 2 – – ()mm 5(7) nextent in :F . mm, 0.2 – – mm, 2 31 Stipes ...... RENCH ...... m m, is sahbi ti xlddfo h UNrdls IC 2017). (IUCN list red IUCN the from however, excluded is rare; it extremely hybrid is a and as locality type the from known locality. Bue oe.and Copel. (Blume) arntn .S 90 yrdzto n looyliyi Central in allopolyploidy and Hybridization 1990. S. D. Barrington, processing Image 2004. 1949. Ram. E. J. Anderson, S. and Magalhaes, J. P. D., M. Abramoff, and space lab provided SA manuscript. the manuscript. CCD revised the traits. and of measured materials author and primary specimens the collected was and analyses, conducted Harvard JHN). to the Fund and Research JHN), JHN), (Systematics to Association to Award Society Systematics Research the Botany the Student JHN), (Graduate Tropical to Biologists Systematic Fellowship in of Fieldwork (Fernald (Award Herbaria Garden Plant University America the of JHN), Society of to Students American Club Graduate the for CCD, Grant Setup to (Research CCD), Taxonomists University and Harvard JHN from to Funds DEB-1311169 (Doctoral Foundation Grant Science Improvement National us the Dissertation by lending part for in provided PTBG was lo- of project provided staff of the Station specimens thank Research We their fieldwork. Pacific for South support Gump gistical California, and of B. Theophilus, University the Richard Tohei at Berkeley Staff Vinette, fieldwork. Suzanne with assisted illustrations. Colin Wang spores. Tristan the and drew scales provided rhizome Nitta of kindly observations C. Ebihara for microscope Atsushi the manuscript. of use the of improved origins greatly possible that the on discussions tohieaense helpful for Butaud François 2014, Jul 9 m, 410 Tohiea, Mt. Moorea. arntn .S 01 hudhbisb rtce ylisting: by protected be hybrids Should 2011. S. D. Barrington, arntn .S,C .Hulr n .R et.18.Hybridization, 1989. Werth. R. C. and Haufler, H. C. S., D. Barrington, atn .adG eit 95 nlsso yrdzones. hybrid of Analysis 1985. Hewitt. G. and N. Barton, ek .B,J .Alsn .M re,adM .Wnhm 02 Identifying 2012. Windham. D. M. and Pryer, M. K. Allison, R. J. B., J. Beck, uhrContributions. Author Acknowledgments. osrainStatus Conservation Notes diinlSeie Examined Specimen Additional Etymology American Inc. Sons, ImageJ. with fteMsor oaia Garden Botanical Missouri the of sandbergii eiuain n pce ocpsi h ferns. the 55 in 79: concepts species and reticulation, Society Botanical Torrey clg n Systematics and Ecology utpeoiiso oyli aa utlcssuyo h hybrid the of study multilocus A taxa: polyploid of origins multiple etnTsoada nnmu eiwrpoie comments provided reviewer anonymous an and Testo Weston . — – 64...... hsi yrdbetween hybrid a is This Polystichum and — ipooisInternational Biophotonics h e yrdi ae fe h type the after named is hybrid new The .membranifolium M. nrgesv Hybridization Introgressive orcimminganense Botrychium irsrmmembranifolium Microsorum eaegaeu oAa .SihadJean- and Smith R. Alan to grateful are We yooyadioyedocumentation. isozyme and Cytology : ieaueCited Literature 3:465 138: 6 113 16: — H ocie h td,gtee h data, the gathered study, the conceived JHN Microsorum — – – 7 297 77: 148. rnhPolynesia. French 471. and ..Nta3929 Nitta J.H. nVermont. in – .spectrum M. 1 36 11: irsrmcommutatum Microsorum 305. irsrmmembranifolium Microsorum e ok onWlyand Wiley John York: New . 3 irsrmcommutatum Microsorum – Microsorum Microsorum 42. tohieaense irsrmpunctatum Microsorum mrcnFr Journal Fern American irsrmgrossum Microsorum irsrmpowellii Microsorum G,P A,UC). PAP, P, (GH, — udn o this for Funding . h ora fthe of Journal The eiou spicatus Lepisorus RB. Ching. (R.Br.) nulRve of Review Annual S OCIETY , 3 3 mhigh mm 1 maximum tohieaense sonly is Betula I SLANDS Annals M. d; 3 3 : Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ek .B,M .Wnhm .Ytkeyh n .M re.21.A 2010. Pryer. M. K. and Yatskievych, G. Windham, D. M. B., J. Beck, 410 ol,J .adJ .Dye 97 ai N slto rcdr o small for procedure isolation DNA rapid A 1987. Doyle. L. J. and J. J. Doyle, hybrid Islands. 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B. and H. D. Hembry, aikm .S,V rdyrj n .D akmr 1997. Rajkumar. D. S. and Irudayaraj, V. S., V. Manickam, L¨ obr,F .adM-.Sce.18.Foao apt,SceyIslands. Society Maupiti, of Flora 1987. Sachet. M.-H. and R. F. Fosberg, oa,R n .E akn r 2004. Jr. Watkins E. J. and R. Moran, CIPRES the Creating 2010. Schwartz. T. and Pfeiffer, W. A., M. Miller, 1966. E. Mayr, afe,C . .A rme,E enpa,T .Rne,A .Smith, R. A. Ranker, A. T. Hennipman, E. Grammer, A. W. H., C. Haufler, chlo- the of inheritance Maternal 1992. Yatskievych. G. and G. Gastony, rs,A . .D ida,adK .Pyr 09 eihrn the Deciphering 2009. Pryer. M. K. and Windham, D. M. L., A. Grusz, ove, Botany uemBulletin Museum interpreting and genus fern delimitation Botany the tematic species in evolution to polyploid approach diploids-first la en( fern cloak uniiso rs eftissue. leaf fresh of quantities speciation. microsoroids. phy- the the of of reconstruction history logenetic a towards attempt an including podiaceae) needn aeohtsi the in gametophytes independent Plus. Gakken emerging? CRAN.R-project.org/package isedrc C,arpdadetato remto o acdn of barcoding for method free ferns. extraction and rapid a PCR, direct Tissue Poly- PhyML. (Polypodiaceae, in selection ferns diverse epiphytic highly a paleotropical of podiopsida). relationships of the lineage Inferring ferns: microsoroid and organization data. the sequence for of platform analysis software integrated desktop Ashton, An Basic: extendable B. Geneious Thierer, and 2012. Drummond. T. J. Duran, A. C. and Mentjies, Markowitz, P. S. Cooper, A. Buxton, S. 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August (accessed ips Pteridaceae). ; 0 3059 30: 0 92 10: – org/documents/RedListGuidelines. – 227. 4911. adnoci radicans Vandenboschia ednBtnclSeries Botanical Leiden hiate yavapensis Cheilanthes – Astrolepis abig:HradUni- Harvard Cambridge: . 8 1647 28: 6 1636 96: 95. – 3066. Lmrosdca)from (Lomariopsidaceae) Taxon 3 1372 43: 3 yoaooia ta of atlas Cytotaxonomical farrarii mrcnJunlof Journal American – – 6 265 46: (Pteridaceae). 1649. 1645. Microsorum 8 1155 48: rse alli´ et eres – :461 6: 9 11 19: enc P Bernice e hybrid new A : 1387. o .Tokyo: 1. vol , YTMTCBTN Vlm 43 [Volume BOTANY SYSTEMATIC – Lecanopteris – – 268. 4 1 14: – 464. 15. American 1167. es complex complex . Paris: . Bishop (Poly- – – 161. Sys- 8in . ie,E . .D ida,adK .Pyr 04 vdnefrreciprocal for Evidence 2014. Pryer. M. K. and Windham, D. M. M., E. Sigel, from inferred phylogeny Fern 2007. Pryer. M. K. and E. The Schuettpelz, 2008. Pryer. M. K. and Windham, D. M. Grusz, L. A. E., Schuettpelz, Haufler, H. C. 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Rothfels, Botany mrcnJunlo Botany genomes. of allopolyploid Journal shape American origins multiple how investigating for in origins Brazil. 1037 speci leptosporangiate 400 nuclear of utility plants. lution epiphytic the of of aspects diversification Exploring Grammitidaceae): polygrammoid and of (Polypodiaceae phylogeny the ferns Unraveling 2004b. Ranker. A. T. and fern of Malesian podiidae). relationships Phylogenetic enigmatic 2004a. Ranker. A. the T. and Haufler, H. C. oyei,wt e species, new a with Polynesia, Science .R mt.21.Asecond A 2018. Smith. R. A. oaia Review Botanical 36 records. new Additional Polynesia: French 4 563 54: ferns. and lycophytes extant for tion Press. Hawaii atr:DAbroigrvascnrsigcmuiystructure 278 gametophytes. 87: community and sporophytes contrasting fern reveals between barcoding DNA matters: 261 of taxonomy https://doi.org/10.5061/dryad.hm813. the Repository. for implications with tohieaense ferns? models. mixed under inference Computing. Statistical for Foundation R http://www.R-project.org. Austria: Vienna, puting. te prxmtl 0mlinyasago. years million 433 each 60 from approximately diverged Pryer. that other M. genera K. between and Windham, hybridization D. Natural M. 2015. Fraser-Jenkins, R. C. Roskam, C. H. hlgntc:Rpdrslto fhbi oyli complexes polyploid hybrid sequencing. of single-molecule PacBio resolution using Rapid phylogenetics: nuclear ferns. single-copy for in Transcriptome-mining Korall, markers P. 2013. Wong, Pryer. K.-S. M. G. K. Stevenson, Cys- W. and D. Graham, the W. S. Ruhsam, in M. allopolyploidy rampant confirm (Polypodiales). data topteridaceae nuclear copy oyopimaayi,drc eunig n cloning. Microbiology length and Environmental sequencing, fragment direct PCR-restriction analysis, nested polymorphism genus-specific of by resolution mixtures and selection. 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Microsorum mrcnFr Journal Fern American oouu nvriyof University Honolulu: . h e Phytologist New The mrcnNaturalist American – Dypeiaee from (Dryopteridaceae) 1036. Plpdaee Poly- (Polypodiaceae, 9 1572 19: clgclMonographs Ecological 6:1077 165: pce n species and species (Psilotaceae). ra Digital Dryad . – Microsorum 1574. – ple and Applied 1087. Blumea Taxon Systematic Pacific ’ orea, 213: 101: 185: 56: 42: 3 Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ye,W .adJ .Gm.1996. Game. C. J. and R. (*and W. parsimony Sykes, using analysis Phylogenetic PAUP* 2002. D. Swofford, 2018] ubrs favailable, individual one if known, than number(s) if locality during more (herbarium), used if specimen rbcL voucher code colon taxon, that individual by for taxon, sampled followed follows: analysis as phylogenetic presented Data study. et,W .adM ude 04 rmr eippyimin Hemiepiphytism Primary 2014. Sundue. M. and L. W. Testo, Tejero-D for primers Universal 1991. Bouvet. J. and Pautou, G. Gielly, L. P., Taberlet, et,W . .E akn r,J itran n .Mmn 2015. Momin. R. and Pittermann, J. Jr., Watkins E. J. L., W. of Testo, Dynamics 2014. Barrington. S. D. and Jr., Watkins E. J. L., W. Testo, evolution. in hybridization of role The 1959. G. Stebbins, phy- likelihood-based Maximum RAxML-VI-HPC: 2006. A. Stamatakis, ubrcrepnst lee1 h eodt lee2 t..Newly etc.). 2, allele to accession second first MG452009 numbers the the accession 1, are (e.g., sequences allele number generated to allele corresponds assigned number arbitrarily of order Missing enn .L n .A akr 03 urn ttso h en and ferns the of status Current 2013. Ranker. A. T. and L. A. Vernon, herbaria public of directory global A Herbariorum: Index 2018. B. Thiers, hn,R,T i,W u .L,L ho .Hag .Hag .Si and Shi, S. Huang, Y. Huang, L. Chao, L. Li, Y. Wu, W. Liu, T. R., Zhang, ag . .P i .P in,J enih,H cnie,adX-.Zhang. X.-C. and Schneider, H. Heinrichs, J. Xiang, Q.-P. Qi, X.-P. L., Wang, anr .H .15.Rtclt vlto nteAplcinaspleniums. Appalachian the in evolution Reticulate 1954. J. H. W. Wagner, On 1998b. Gibby. M. 1998a. and Barrett, Gibby. A. J. Rumsey, M. J. F. and Russell, J. S. Barrett, C., J. J. Vogel, Rumsey, J. F. Russell, S. C., J. Vogel, aaa . .Ytb-auaa .Hr,adN uaai 2016. Murakami. N. and Hori, K. Yatabe-Kakugawa, Y. K., Yamada, ag . .W,Q-.Xag .Hircs .Shedr n .C Zhang. X.-C. and Schneider, H. Heinrichs, J. Xiang, Q.-P. Wu, Z. L., Wang, anr .H .16.Aboytmtcsre fUie ttsferns States United of survey biosystematic A 1963. J. H. W. Wagner, in,L,C .Wrh .N mr,adD .MCue.20.Population- 2000. McCauley. E. D. and Emery, N. S. Werth, R. C. L., Xiang, 2017. Zhang. X.-C. and Zhang, H.-R. Zhao, C.-F. Wei, R. X.-P., Wei, A PPENDIX te ehd) .40bt 0 udrad iae Associates. Sinauer Sunderland: 10. beta 4.0 v. methods), other okIslands. Cook ampla circum- the on genus. influence the of its scription and Polypodiophyta) (Polypodiaceae, DNA. chloroplast of Biology regions Molecular non-coding three of amplification Brittonia Pteris Phytologist New biology. Recon- reproductive The gametophyte ferns: with wood inheritance of American patterns ciling North in hybridization asymmetrical ferns. 526 in habit growth Society Philosophical American models. mixed and taxa of thousands informatics with analyses logenetic ica,Pteridophyta). niaceae, Islands. Hawaiian the of lycophytes Garden Botanical http://sweetgum.nybg.org/science/ih. York ium. New staff. associated and .Zo.21.Mlclreiec o aua yrdzto nthe in hybridization natural for genus evidence fern mangrove Molecular 2013. Zhou. R. Japan. lands, oica,Plpdosd)ifre rmfu hools N re- DNA genus chloroplast four fern gions. from paleotropical inferred Polypodiopsida) the podiaceae, of Phylogeny 2010a. abstract. Preliminary Evolution fern rock the in formation hybrid genus the in DNA chloroplast Asplenium of transmission maternal for Evidence smerchbi omto eeldb riiilcosn experi- crossing artificial between by ments revealed formation hybrid Asymmetric Botany of 00.Amlclrpyoeyadarvsdcasfcto ftribe DNA of plastid classification four of revised regions. analysis an a on based and (Polypodiaceae) phylogeny Lepisoreae molecular A 2010b. pcfcgne-isdhbiiainbetween and hybridization gender-biased specific gene nuclear 450 and 178: data. chloroplast morphological and from regions Evidence genus fern revisited: enigmatic podiaceae) the of position Phylogenetic ceso number, accession – ´ ı rbcL .carthusiana D. z .D,J .Mce,adA .Sih 09 hybrid A 2009. Smith. R. A. and Mickel, T. J. D., J. ez, 536. 3 Plpdaee rvdsnwisgtit h vlto of evolution the into insight new provides (Polypodiaceae) oeua hlgntc n Evolution and Phylogenetics Molecular .Gnakacsinnmesfrpatmtra sdi this in used material plant for numbers accession GenBank 1. caridadiae – oaia ora fteLnenSociety Linnean the of Journal Botanical 7 1 67: 464. or 7 1175 87: :103 8: Apeica,Pteridophyta). (Aspleniaceae, 2 2688 22: trnL – caPyoaooiae Geobotanica et Phytotaxonomica Acta e eln ora fBotany of Journal Zealand New 6. – slnu setoi Asplenium 118. – 7 1105 17: – F vdnefo hools DNA. chloroplast from Evidence : Peiaee,anwhbi enfo ot Rica. Costa from fern hybrid new a (Pteridaceae), 1180. – trnL niae by indicated 0:785 206: 2690. aC short gapCp mrcnFr Journal Fern American Acrostichum mrcnFr Journal Fern American IT TA. E YRDMCOOU RMFEC OYEI 411 POLYNESIA FRENCH FROM MICROSORUM HYBRID NEW AL.: ET NITTA oaiaActa Botanica – nentoa ora fPatSciences Plant of Journal International F – 1109. ceso number, accession – 795. 0:231 103: rmteOaaaaadRuy is- Ryukyu and Ogasawara the from Phymatosorus nentoa ora fPatSciences Plant of Journal International ceso ubrs favailable. if number(s) accession . — Asplenium M ln Biology Plant BMC mrcnFr Journal Fern American cesosof Accessions . 1:241 111: – 251. oaiaActa Botanica 3 1 53: 4 211 54: 4 143 34: 9 109 99: – Plpdaee nthe in (Polypodiaceae) 3 246. 6:28 162: aC long gapCp rotrsintermedia Dryopteris – alternifolium 7 147 67: 16. – ’ – Weatherbya ita Herbar- Virtual s G502( MG452032 – 225. rceig fthe of Proceedings mrcnJournal American – Lepisorus 146. gapCp 116. – 3 74. 13: 1:247 111: 38. – 0:59 103: 158. Phlebodium accession ie in given (Asple- Colysis (Poly- (Poly- – rbcL – Plant 249. 175: 111. Bio- — ), MG427052 912010) lomarioides 87GR00136) Lecanopteris (UBG 7865 AY083632. Hennipman AF470329, ippines, AY083626. aasa F731 AY083634. AF470331, Malaysia, AY083628. AF470325, Hennipman nipman s.n. Klein AY083631. AF470328, wesi, AY083645. nesia, Garden. Bo- Botanical Berkeley Tropical California, Xishuangbanna XTBG, of Garden; University tanical KBG, UCBG, Garden; Garden; Botanical ¨ RBGE, T Utrecht Botanical Garden; TBG, Edinburgh; Botanical Leiden Garden York Botanic New Royal LBG, NYBG, Garden; Garden; Botanical USA); Kunming Botanical (Florida, Alford Duke Charles of DBG, Garden of CAG, Zurich; University Garden the Botanical of Garden Botanical Botanical BGB, G¨ Old follows: BGG, as Berlin-Dahlem; abbreviations Garden garden Botanical (2018). Thiers MG550279 pyriforme auriculatum 239-36-90-30 BGB cult. 90-33 EU483026. Parris (C.Chr.) Ho. Copel. (Hook.) Copel. CL)Tia,E423,G162;2: 5162 Zhang GU126729; EU483034. EU482937, EU482939, Taiwan, (COLO) GU126728. BGZ) 50.0326) (UCBG carnosum Lemmaphyllum adnascens 92GR01365) (UBG s.n. AY083624. AF470321, blechnoides unn hn,G169,GU126727. GU126697, China, Yunnan, GU126725. m ..hn)L Wang Li X.C.Zhang) & Sm. hn,G265,GQ256177. GQ256259, China, asterolepis Lepisorus annuifrons 05-277 GQ256173. affinis Lepisorus 3: KX891354; 2: EU483031; EU482936, 1: Hosok. (Blume) J17 Lu SG GU126723. GU126693, hemionitideum 80894 Shui GU126721. — solida Davallia long gapCp 5157 ey,G266,GQ256178. GQ256260, Kenya, 1: Wang GQ256214. GQ256290, China, Tibet, (PE) GQ256174. GQ256257, China, GE)Tbt hn,D625,D623;2: DQ642236; DQ642154, China, Tibet, (GOET) GQ256180. Ching (Christ) u2440 1: Wu Hook. ex 2: DQ164505; J.Sm. Hook.) ex tign G,BtnclGre fteUiest fHiebr;BGZ, Heidelberg; of University the of Garden Botanical BGH, ottingen; aymdnorientalis Calymmodon , — , GE) U898 U803 3: EU483033; EU482938, (GOET), P)Tbt hn,G266,GQ256179. GQ256261, China, Tibet, (PE) B,E423,EU483028. EU482933, (B), — aC short gapCp L atKlmna,Idnsa U891 EU483025. EU482931, Indonesia, Kalimantan, East (L) ut BE171 RBGE cult. , i873 Li D L aasa F736 AY083629. AF470326, Malaysia, (L) enpa ..(B 85GR00170) (UBG s.n. Hennipman L uaei F732 AY083625. AF470322, Sulawesi, (L) — PU unn hn,A755,AY725049. AY725055, China, Yunnan, (PYU) KN as X079 JX103793. JX103709, Laos, (KUN) – – Ching oiplbu persicifolium Goniophlebium eaotrscrustacea Lecanopteris Cig Ching (Ching) Mkn)Ching (Makino) eaotrsbalgooyi Lecanopteris eioirsru buergerianum Lepidomicrosorium G207( MG427077 G535( MG550315 empylmdiversum Lemmaphyllum empylmrostratum Lemmaphyllum P)Cia Q535 GQ256242. GQ256315, China, (PE) eiou albertii Lepisorus , Ge. Parris (Grev.) G596 MG451947, MG451946, eiou clathratus Lepisorus P)Scun hn,G169,GU126726. GU126696, China, Sichuan, (PE) aC long gapCp cnie FJ11-129 H Schneider Bue C.Presl (Blume) enpa 80(B 87GR00084) (UBG 7820 Hennipman i 029(ut KBG) (cult. 2012-9 Kim oekm 05-278 Hovenkamp ut B 54022 TBG cult. GFrt)Sw. (G.Forst.) Crs)P.S.Wang (Christ) U)Jpn F732 Y865 2: AY083635; AF470332, Japan, (UC) P)Hia,Cia Q522 Q516 2: GQ256166; GQ256252, China, Hainan, (PE) enpa 81(B 87GR00087) (UBG 7821 Hennipman hn 4237 Zhang Ching G522 MG550283. MG550282, oekm 05-298 Hovenkamp U uaei F737 AY083630. AF470327, Sulawesi, (U) Bkr hn xS.X.Xu ex Ching (Baker) eaotrsspinosa Lecanopteris eioirsru superficiale Lepidomicrosorium empylmintermedium Lemmaphyllum eaotrspumila Lecanopteris trnL E,E423,EU483030. EU482935, (E), MG451945, B,E423,EU483029. EU482934, (B), aC short gapCp hn 29(ut SZBG) (cult. 4219 Zhang it 658 Nitta u Ykuo2072 LY Kuo Knee et)Copel. Mett.) ex (Kunze Copel. BX 07087 Xu WB Wl.e .m)CPel1: C.Presl J.Sm.) ex (Wall. – ouos.n. Kokubo hn 33(ut TBG) (cult. 4363 Zhang eaotrscarnosa Lecanopteris F P)Scun hn,G169,GU126724. GU126694, China, Sichuan, (PE) akr1935 Ranker eiou bampsii Lepisorus rirsn cl.BGG) (cult. s.n. Kreier ,MG451945 ), rnilTW075 Cranfill eiou angustus Lepisorus Rgl Ching (Regel) eiou bicolor Lepisorus hraimukon aa,GQ256262, Japan, unknown) (herbarium L atKlmna,Idnsa EU482932, Indonesia, Kalimantan, East (L) oiplbu pseudoconnatum Goniophlebium eaotrssarcopus Lecanopteris eaotrsdeparioides Lecanopteris B)Piipns X938 KX891353. KX891368, Philippines, (BM) it 682 Nitta G)Moe,Fec Polynesia, French Moorea, (GH) Hennipman CBCak)Cig1: Ching (C.B.Clarke) Copel. aC short gapCp U aasa F734 AY083627. AF470324, Malaysia, (U) KN hn,J132,J130;3: JX103805; JX103721, China, (KUN) .Hraimabeitosfollow abbreviations Herbarium ). hn 4111 Zhang aC short gapCp Rsnt)Tgw 1: Tagawa (Rosenst.) hn 4364 Zhang empylmsquamatum Lemmaphyllum T)Jpn Q528 GQ256176. GQ256258, Japan, (TI) Bd. Tagawa (Bedd.) L atKlmna,Indonesia, Kalimantan, East (L) eiou annamensis Lepisorus P)Gagi hn,GU126692, China, Guangxi, (PE) eignBtncGre;UBG, Garden; Botanic uebingen Ds. Bedd. (Desv.) U)Moe,Fec Polynesia, French Moorea, (UC) oiplbu argutum Goniophlebium rne n os341 Roos and Franken – TI hlpie,KX891369, Philippines, (TAI) G596( MG451976 em T.G.Walker & Jermy oiplbu mehibitense Goniophlebium G)Moe,Fec Poly- French Moorea, (GH) eaotrscelebica Lecanopteris hn 4325 Zhang Blume Mq)Cig&K.H.Shing & Ching (Miq.) hn 1854 Zhang U)Tia,DQ164442, Taiwan, (UC) L uaei AF470323, Sulawesi, (L) hn 5171 Zhang MG550279. hn 4515 Zhang ai li s.n. Klein David eaotrsluzonensis Lecanopteris eioirsru sub- Lepidomicrosorium Zink G520 MG550281. MG550280, P)Cia GQ256256, China, (PE) P)Gagi China, Guangxi, (PE) Tkd)Ching (Takeda) P)Jpn GU126698, Japan, (PE) Riw)Blume (Reinw.) eiou boninensis Lepisorus eaotrssinuosa Lecanopteris Ching oiplbu sub- Goniophlebium enpa ..(LBG s.n. Hennipman U )Philippines, L) (U, GE) AF470342, (GOET), .Vae11233 Viane R. odas551 Woodhams eiou accedens Lepisorus cnie ..(cult. s.n. Schneider Cig iWang Li (Ching) Bue iWang Li (Blume) P) GU126695, (PE), olco unknown collector ut G 239-12- BGB cult. Tis.&Binn.) & (Teijsm. U Philippines, (U) aC long gapCp hi80676 Shui Lemmaphyllum Lemmaphyllum ..Se S25 Shen Z.H. P)Xinjiang, (PE) ikr 12430 Dickore Ctenopterella P)Sichuan, (PE) P)Yunnan, (PE) Cs)Baker (Ces.) P)China, (PE) akr2079 Ranker Hovenkamp CCr)Li (C.Chr.) Lepisorus L Sula- (L) L Phil- (L) Jaarsma Copel. — Zhang (Wall. ,and ), David (A.R. Hen- (PE) (PE) , (L) — , Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. hn,G269,GQ256220. GQ256295, China, P)Yna,Cia Q531 Q526 2: GQ256226; GQ256225. GQ256300, China, GQ256301, China, Yunnan, (PE) GQ256224. subconfluens isorus 2295 Zhang 2: JX103782; P)Yna,Cia Q526 GQ256221. 1: Ching GQ256296, China, Yunnan, (PE) thunbergianus 2005-045A Rao Y.X.Lin GQ256170. EF463244, Polynesia, Tahiti: French MG550286; MG550285, MG550284, MG427052, MG452009, Moorea: DQ642234; DQ642153, spicatus isorus EU483037. EU482942, Taiwan, 20050117 Zhang 2: DQ179639. GQ256172; AY362563, GQ256255, China, Sichuan, ianus TW069 Cranfill GQ256205. GQ256282, China, Tibet, scolopendrium GQ256204. 2: JX103781; JX103697, China, 412 Crs)Cig&S.K.Wu GQ256249. & Ching (Christ) 3218 HQ712015. HQ711996, Tanzania, GQ256217. 4736 Zhang RV8253 marginatus isorus 3: EU483036; macrosphaerus Lepisorus 3: GQ256181. GQ256197; GQ256263, GQ256275, China, ooo,D625,G268;2: GQ256189; GQ256188. 1: DQ642155, DQ642156, Ching Willd.) Comoros, ex (Bory 4444 3: GQ256183. GQ256265, GQ256184; China, GQ256266, China, Sichuan, 1: GQ256182. GQ256264, China, Yunnan, hn 4437 Zhang pumilus Tagawa GQ256215. GQ256291, China, Sichuan, (PE) HQ712017. HQ711995, Kenya, perrierianus Lepisorus S.K.Wu & Ching 4352 Zhang GQ256211. GQ256287, China, GQ256210. venulosus GQ256208. GQ256284, China, Tibet, GQ256191. heterolepis Wang GQ256190. GQ256269, Japan, luchunensis Ching GQ256198. GQ256276, China, Tibet, GQ256230. GQ256303, China, 4488 1: Wu aa 5891 Jaman ee,Cia Q521 GQ256192. GQ256271, China, Hebei, nu,Cia Q523 GQ256195. GQ256273, China, Anhui, GQ256194. GQ256272, 4615 eiou kuchenensis Lepisorus Tagawa (Hayata) hn 4699 Zhang P)Yna,Cia Q524 Q516 3: GQ256196; GQ256274, China, Yunnan, (PE) P)Yna,Cia Q527 GQ256222. GQ256297, China, Yunnan, (PE) P)Yna,Cia Q528 GQ256187. GQ256268, China, Yunnan, (PE) P)Tbt hn,G262,GQ256248. GQ256320, China, Tibet, (PE) Mkn)Fae-ek uhCada1: Subh.Chandra & Fraser-Jenk. (Makino) hi80679 Shui hn 4440 Zhang hn 4468 Zhang ..Fn XZ-412 Fang Z.D. hraimukon eno,H719,HQ712016. HQ711998, Reunion, unknown) (herbarium hn S.K.Wu & Ching rnilTW093 Cranfill hn 0612 Zhang Hyt)Ching (Hayata) eiou medogensis Lepisorus eiou hsiaowutaiensis Lepisorus P)Yna,Cia Q527 Q519 2: GQ256199; GQ256277, China, Yunnan, (PE) eiou sublinearis Lepisorus eiou oligolepidus Lepisorus eiou sp Lepisorus Rsnt)Ching (Rosenst.) P)Tbt hn,G268,G260;2: GQ256209; GQ256285, China, Tibet, (PE) eiou rotundus Lepisorus P)Ida Q523 Q528 4: GQ256218; GQ256293, India, (PE) ..Tn YD-076 Tang Y.D. Y.X.Lin U)Mlyi,A326,GQ256168. AY362562, Malaysia, (UC) P)Jpn Q528 GQ256212. GQ256288, Japan, (PE) hn 4794 Zhang P)Tbt hn,G260,G263;2: GQ256235; GQ256308, China, Tibet, (PE) Kuf)Cig1: Ching (Kaulf.) Cig er i 1: Bir & Mehra (Ching) U)Tia,D625,GQ256206. DQ642158, Taiwan, (UC) P)Tia,G268,G260;2: GQ256207; GQ256283, Taiwan, (PE) Lf)L agCult: Wang Li (L.f.) P)Sax,Cia Q532 GQ256229. GQ256302, China, Shanxi, (PE) hn 3413 Zhang P)Yna,Cia Q523 GQ256167. GQ256253, China, Yunnan, (PE) P)Yna,Cia Q528 GQ256201. GQ256278, China, Yunnan, (PE) akr2051 Ranker P)Yna,Cia Q527 Q516 2: GQ256186; GQ256267, China, Yunnan, (PE) Ching P)Scun hn,G269,G262;2: GQ256223; GQ256298, China, Sichuan, (PE) Ching i097 Qi YCW)Ching (Y.C.Wu) eiou hachijoensis Lepisorus eiou lewisii Lepisorus eiou monilisorus Lepisorus eiou confluens Lepisorus CCr)Ching (C.Chr.) aC long gapCp U)Tia,E424,GQ256216. EU482943, Taiwan, (UC) ..Wn 60667 Wang M.Z. i 2012-10 Kim Bkr hn 1: Ching (Baker) P)Tbt hn,G268,GQ256203. GQ256280, China, Tibet, (PE) P)Tbt hn,G267,GQ256202. GQ256279, China, Tibet, (PE) hn 5168 Zhang hn 4151 Zhang hn 3360 Zhang hn 4518 Zhang P)Yna,Cia Q100 HQ712019. HQ712000, China, Yunnan, (PE) aoodanb 6785 Rakotondrainibe eiou thaipaiensis Lepisorus eiou sinensis Lepisorus eiou elegans Lepisorus hn 5064 Zhang eiou lineariformis Lepisorus P)Hbi hn,G268,GQ256213. GQ256289, China, Hubei, (PE) P)Tbt hn,G269,G261;3: GQ256219; GQ256294, China, Tibet, (PE) eiou pseudonudus Lepisorus eiou morrisonensis Lepisorus eiou henryi Lepisorus eiou onoei Lepisorus it 323 Nitta akrTW018 Ranker CL)Tia,E424,GQ256193. EU482940, Taiwan, (COLO) hr 2005042404 Ohora Ching hn Y.X.Lin & Ching Bkre aea hn 1: Ching Takeda) ex (Baker Bkr Ching (Baker) eiou mucronatus Lepisorus eiou schraderi Lepisorus eiou loriformis Lepisorus eiou megasorus Lepisorus hn 4533 Zhang G598 MG451949, MG451948, eiou likiangensis Lepisorus eiou inaequibasis Lepisorus cnie ..(ut BGG) (cult. s.n. Schneider eiou contortus Lepisorus hn S.K.Wu & Ching ..X 08188 Xi J.M. KN,J132,JX103806. JX103722, (KUN), P)Gagi hn,GQ256286, China, Guangxi, (PE) P)Scun hn,GQ256321, China, Sichuan, (PE) u2441 Wu P)Yna,Cia GQ256299, China, Yunnan, (PE) eiou stenistus Lepisorus P)Hbi hn,GQ256281, China, Hubei, (PE) ep3561 Hemp RV7675 G)Moe,Fec Polynesia, French Moorea, (GH) B245 EB Bkr Ching (Baker) P)Tbt hn,GQ256270, China, Tibet, (PE) P)Gnu hn,GQ256292, China, Gansu, (PE) i 023(ut KBG) (cult. 2012-3 Kim rnilTW087 Cranfill hn 5204 Zhang eiou pseudo-ussuriensis Lepisorus akr1915 Ranker W.M.Chu eiou sordidus Lepisorus Sa.Kurata Hyt)Tgw 1: Tagawa (Hayata) U)Tia,EU482941, Taiwan, (UC) hn xW.M.Chu ex Ching hi81060 Shui Crs)Ching (Christ) Fac.&Sv)Ching Sav.) & (Franch. KN as JX103698, Laos, (KUN) Heo.e .h. Li C.Chr.) ex (Hieron. ..LuDB06104 Liu C.C. hn 4659 Zhang eiou patungensis Lepisorus hraimunknown) (herbarium hraimunknown) (herbarium hn 5082 Zhang P)Yna,China, Yunnan, (PE) eiou kawakamii Lepisorus ..Fn XZ-266 Fang Z.D. T)Jpn GQ256305, Japan, (TI) hn 5117 Zhang P rneComore, Grande (P) eiou soulieanus Lepisorus hn S.K.Wu & Ching eiou excavatus Lepisorus P)Gagi China, Guangxi, (PE) eiou obscure- Lepisorus akrTW012 Ranker hn ..u1: S.K.Wu & Ching eiou miyosh- Lepisorus DM Tanzania, (DSM) Ching ..X s.n. Xu C.D. Hyt)H.It (Hayata) P)Chongqing, (PE) ..Lus.n. Liu Q.R. i 05620 Liu Y hn 5113 Zhang hn 4358 Zhang hn 5187 Zhang hn 4771 Zhang Wl.e Mett.) ex (Wall. CL)Tahiti, (COLO) CCr)Ching (C.Chr.) Mt. Ching (Mett.) Crs)Ching (Christ) (F´ U)Taiwan, (UC) P)Yunnan, (PE) hn S.K. & Ching (C.B.Clarke) e iWang Li ee) YTMTCBTN Vlm 43 [Volume BOTANY SYSTEMATIC ined. aC short gapCp hn 4249 Zhang hi80595 Shui P)Tibet, (PE) P)Tibet, (PE) hi81069 Shui P)Tibet, (PE) Lepisorus Lepisorus Lepisorus Lepisorus Lepisorus (GOET), (C.Chr.) (KUN) Zhang Zhang Zhang Zhang H.M. (UC) Lep- Lep- Lep- G.Y. (PE) (PE) (PE) (PE) (PE) (PE) (PE) (PE) (PE) o1: ˆ GQ256234. CL)Moe,Fec oyei,EF463253, Polynesia, French Moorea, (COLO) MG427062, MG451956, KY099829, Polynesia, French Moorea, aa,G260,GQ256236. GQ256309, Japan, tibeticus 026(ut KBG) (cult. 2012-6 short gapCp 2901 Smith MG451954, 1: Moorea MG427059; MG452017, Polynesia, Huahine: MG427057; MG452015, nesia, Bora: Bora S.B.Andrews Fisch.) 3560) (LBG hn 5205 Zhang 2: GQ256232; KN ita,J503,J503;2: JX520937; JX520933, Vietnam, (KUN) heterophyllus Leptochilus hemitomus aii rnhPlnsa G502 MG427055, MG452012, Polynesia, French Tahiti, MG452013, Polynesia, short gapCp 1: Moorea MG427054; MG452011, Polynesia, 55.0092) (UCBG 3481 Grether commutatum Microsorum 2: JX103811; ora rnhPlnsa G509 MG427061, MG452019, Polynesia, French Moorea, 1: X.C.Zhang Baker) X298 3: JX520938; 2: EU483046; hemionitideus 2: MG550287; ora rnhPlnsa G504 MG427056, MG452014, Polynesia, French Moorea, Noot. aC long gapCp GL J17 Lu SG 2: JX103788; JX103704, membranaceum aa,M421,MG427053, MG452010, Japan, folius EU483045. ellipticus 3: 2: JX103779; EU483044; EU482948, Vietnam, (UC) JX103808. JX103724, decurrens Leptochilus 2: EU483042; EU482946, JX103785. i 214 Liu U895 EU483061. EU482965, pteropus Leptochilus 4: JX103787; JX103703, 1863 Moorea: MG427064, MG451959, MG452021, nesia, Maupiti: MG427065; MG452022, Huahine: Copel. (Brack.) KBG) (cult. AY083637. AF470334, linguiforme Microsorum lastii MG427060. Microsorum MG452018, Polynesia, Pou: French hainanense Ua Pou, MG427058; Ua MG452016, (PTBG) Polynesia, French Tahiti, KN as X070 X074 3: JX103784; JX103700, Laos, (KUN) 1: Kaulf. (Cav.) A0303 insignis GQ256250. 2: EU483039; EU482944, China, iensis 2005042902 Fujimoto GQ256239. distans GQ256238. GQ256311, G)Tht,Fec oyei,M422,MG427063. MG452020, Polynesia, French Tahiti, (GH) hn 3800 Zhang Bc.Hm xD o)XC hn 1: Zhang X.C. Don) D. ex (Buch.-Ham. Rgl&Mak Ching Maack) & (Regel P)Yna,Cia Q533 GQ256240. GQ256313, China, Yunnan, (PE) Mkn)Tagawa (Makino) Bue rsrJn.1: Fraser-Jenk. (Blume) ike106 Hinkle P)Cia U898 U805 3: EU483055; EU482958, China, (PE) hn S.K.Wu & Ching Tub xMra)Noot. Murray) ex (Thunb. cantoniensis Leptochilus (UC), eiou tosaensis Lepisorus etciu ellipticus Leptochilus U) F735 AY083638. AF470335, (UC), aC short gapCp KN hn,J132,JX103810. JX103726, China, (KUN) eiou xiphiopteris Lepisorus eiou waltonii Lepisorus MG451960, Hne Noot. (Hance) MG550288. PU unn hn,A755,AY725051. AY725053, China, Yunnan, (PYU) G529 ora2: Moorea MG550289; aC short gapCp aC short gapCp Noot. P)Cogig hn,G260,GQ256231. GQ256304, China, Chongqing, (PE) stui1067 Tsutsumi hn 3509 Zhang WP-135 hn 4544 Zhang u2437 Wu CPel ot 1: Noot. (C.Presl) u2712 Wu DDn hn 1: Ching (D.Don) u2344 Wu — P) U893 EU483049. EU482953, (PE), KN hn,J130,J139;2: JX103791; JX103707, China, (KUN) U801 Huahine: EU483051; , Bkr Tardieu (Baker) U)Moe,Fec oyei,K093,MG427066, KY099833, Polynesia, French Moorea, (UC) ag14 cl.SCIB) (cult. 1348 Wang etciu digitatus Leptochilus T)Jpn Q530 GQ256237. GQ256310, Japan, (TI) Bue Fraser-Jenk. (Blume) KN ita,J138,JX103772. JX103688, Vietnam, (KUN) i 021 cl.KBG) (cult. 2012-15 Kim irsrmlucidum Microsorum eiou ussuriensis Lepisorus G521 G522 ora2: Moorea MG550292; MG550291, — etciu shintenensis Leptochilus i9 cl.XTBG) (cult. 95 Li Blume irsrmcuspidatum Microsorum aC short gapCp hn 3510 Zhang G523 G524 ora3: Moorea MG550294; MG550293, G525 G526 G527 MG550298; MG550297, MG550296, MG550295, KN as X064 JX103778. JX103694, Laos, (KUN) , KN as X066 JX103780. JX103696, Laos, (KUN) P)Cia U897 EU483043. EU482947, China, (PE) P)Yna,Cia Q536 Q523 3: GQ256233; GQ256306, China, Yunnan, (PE) og743 Dong KN as X069 X073 2: JX103783; JX103699, Laos, (KUN) it 3972 Nitta .Fjmt SF05051602 Fujimoto S. hn 3302 Zhang Copel. aC short gapCp Bue oe.AmrlyIslands: Admiralty Copel. (Blume) SKW ..Pa)Crseh 1: Christenh. K.L..Phan) & (S.K.Wu C)Oiaa aa,E425,EU483050. EU482954, Japan, Okinawa, (CT) U)AmrlyIlns AY362571, Islands, Admiralty (UC) hn 4694 Zhang i 204 Liu Mkn)H.It (Makino) eiou uchiyamae Lepisorus i 021 cl.KBG) (cult. 2012-12 Kim Cig ..u1: S.L.Yu (Ching) hn 4639 Zhang it 3837 Nitta oa ..(ut NYBG) (cult. s.n. Moran ..Lus.n. Liu B.D. it 3674 Nitta aC long gapCp Tub xMra)No.var. Noot. Murray) ex (Thunb. aC long gapCp .Rne 1776 Ranker T. i 022(ut KBG) (cult. 2012-2 Kim Bkr ..h xY.X.Lin ex W.M.Chu (Baker) uahsen2 Gulamhussein Bkr hn 1: Ching (Baker) eir7937 Perier G529 G530 Tahiti: MG550300; MG550299, P)Cia U895 EU483041. EU482945, China, (PE) P)Cia U899 EU483056. EU482959, China, (PE) hn 1923 Zhang P)Cia U897 U804 2: EU483054; EU482957, China, (PE) G)Haie rnhPolynesia, French Huahine, (GH) WP-201 u2462 Wu P)Cia U891 EU483047. EU482951, China, (PE) un504 Dunn G520 Tahiti: MG550290; irsrmgrossum Microsorum u2515 Wu it 4046 Nitta P)Tbt hn,GQ256307, China, Tibet, (PE) rirsn cl.BGG) (cult. s.n. Kreier P) U892 U809 3: EU483059; EU482962, (PE), G)Br oa rnhPoly- French Bora, Bora (GH) Bkr ot 1: Noot. (Baker) etciu wrightii Leptochilus P)Tbt hn,GQ256322, China, Tibet, (PE) G)Muii rnhPoly- French Maupiti, (GH) P) U890 EU483057. EU482960, (PE), o Rgl&Mak hn var. Ching Maack) & (Regel Rx. Copel. (Roxb.) ˆ ace-aaad 170 Sanchez-Baracaldo ace-aaad 175 Sanchez-Baracaldo .Fjmt 2005042904 Fujimoto S. P)Hiogin,China, Heilongjiang, (PE) KN ita,JX520934, Vietnam, (KUN) P,E426,EU483058. EU482961, (P), Microsorum it 377 Nitta MG451950, — KN hn,JX103727, China, (KUN) G597 MG451958, MG451957, PB)Haie French Huahine, (PTBG) KN as JX103701, Laos, (KUN) u2435 Wu DDn aaa1: Tagawa (D.Don) Hyt)XCZag& X.C.Zhang (Hayata) KN as JX103695, Laos, (KUN) Tahiti: ; P)Cia EU482949, China, (PE) rnil94-266-29 Cranfill aC long gapCp aC long gapCp aC long gapCp aC long gapCp T)Jpn GQ256312, Japan, (TI) G)Haie French Huahine, (GH) U)Moe,French Moorea, (UC) etciu axillaris Leptochilus og172 Dong U)NwGuinea, New (UC) Mkn)H.It (Makino) olco unknown collector eiou ussur- Lepisorus N) EU482950, (NY), iet 33.3 Vinette G)Okinawa, (GH) KN China, (KUN) mr13 Amer mr14 Amer KN China, (KUN) KN Laos, (KUN) mt 00-036 Smith 3 oec 9155 Lorence i 2012-14 Kim aC short gapCp Microsorum Microsorum Microsorum akr1941 Ranker Leptochilus Leptochilus Leptochilus Leptochilus P)China, (PE) MG451955, MG451951, MG451953, MG451952, Lns.& (Langsd. maximum anr& Wagner Lepisorus Ho.& (Hook. u2439 Wu — (GOET), ..Xu C.D. WP-136 pothi- Cult: ; (GH) (GH) (UC) (UC) Nitta (UC) (UC) o ˆ Kim (TI) S. Delivered by Ingenta to IP: 5.10.31.151 on: Tue, 11 Sep 2018 15:16:03 Copyright (c) American Society for Plant Taxonomists. All rights reserved. irsrmpunctatum Microsorum MG451968, MG451967, MG451966, Q536 Q524 Cult: GQ256244; GQ256316, atuhss.n. Baltrushes long iet 32.2 Vinette MG427072, MG452028, Polynesia, French Moorea, Q658 Huahine: DQ164508; isae(B 24091) (LBG Ridsdale MG451970, MG451969, G203 Laos: MG427073; DQ642246. DQ642162, papuanum Microsorum 2: aa noei,A403,AY083636. 1: AF470333, Copel. Indonesia, Java, musifolium Microsorum MG427071, MG452027, MG427070, short MG452026, Polynesia, French oyei,M422,M476;U o 1: Pou Ua MG427069; MG452025, Polynesia, short gapCp oyei,M422,MG427068, MG452024, Moorea: Polynesia, MG427067; MG452023, nesia, Huahine: DQ642245; DQ642161, branifolium J17 DQ401122. 2018] latum 2096 Ranker membranaceum 2: 1: DQ401125. Copel. 3: (Burm.f.) DQ401119, DQ401123; Zealand, New DQ401118, lington, Zealand, New gina, BGG) (cult. AB575280, Japan, nawa, oe.var. Copel. 2367 660 Rakotondrainibe aC short gapCp MG427075, MG452030, Hawaii, Kauai, U804 Oahu: EU483064; GE) U899 EU483066. EU482969, (GOET), thailandicum Microsorum var. ora1: Moorea akr1941 Ranker R 3584 LRP PU unn hn,A755,AY725050. AY725054, China, Yunnan, (PYU) G591 MG451972, MG451971, spectrum G533 aPu2: Pou Ua MG550303; KN as X078 JX103792. JX103708, Laos, (KUN) Copel. R 3558 LRP irsrmrubidum Microsorum GE) Q107 Q761 2: DQ179641; DQ212057, (GOET), U)Moe,Fec oyei,K093,MG427074, KY099832, Polynesia, French Moorea, (UC) G531 MG550312. MG550311, pentadactylum G531 G532 Tahiti: MG550302; MG550301, it 3929 Nitta CL)Tia,E426,EU483063. EU482966, Taiwan, (COLO) WL)Wligo,NwZaad Q010 DQ401124. DQ401120, Zealand, New Wellington, (WELT) RB. hn Cult: Ching (R.Br.) CL)Moe,Fec oyei,D193,D194;3: DQ179642; DQ179633, Polynesia, French Moorea, (COLO) U)Moe,Fec oyei,MG452029, Polynesia, French Moorea, (UC) R 3575 LRP Kauai: DDn hn var. Ching (D.Don) cnie ..(ut BGG) (cult. s.n. Schneider u2506 Wu G1350 HG WL)Tae,NwZaad Q016 DQ401121; DQ401116, Zealand, New Thames, (WELT) P aot,G261,GQ256245. GQ256317, Mayotte, (P) it 4045 Nitta U)Piipns F737 Y860 Taiwan: AY083640; AF470337, Philippines, (UC) irsrmpowellii Microsorum od10936 Wood aC short gapCp — aC long gapCp WL)Wligo,NwZaad DQ401117, Zealand, New Wellington, (WELT) G)Moe,Fec oyei,MG452032, Polynesia, French Moorea, (GH) Copel. L)Cpl China: Copel. (L.) Parris . oned&Noot. & Boonkerd irsrmscandens Microsorum KN as X075 X079 ora1: Moorea JX103789; JX103705, Laos, (KUN) aC short gapCp un458 Dunn IT TA. E YRDMCOOU RMFEC OYEI 413 POLYNESIA FRENCH FROM MICROSORUM HYBRID NEW AL.: ET NITTA Hler)D .Palmer D. D. (Hillebr.) cnie ..(ut BGH) (cult. s.n. Schneider U)Oh,Hwi,E426,EU483065. EU482968, Hawaii, Oahu, (UC) olco nnw UB 58.0649) (UCBG unknown collector it 4073 Nitta cutpl 0 cl.BGB) (cult. 603 Schuettpelz Kne Copel. (Kunze) G)Haie rnhPolynesia, French Huahine, (GH) Microsorum G537 G538 Philippines: MG550308; MG550307, carinatum irsrmspectrum Microsorum cnie ..(ut BGG) (cult. s.n. Schneider aC long gapCp MG451964, aC short gapCp PB)Kui aai EU482967, Hawaii, Kauai, (PTBG) PB)U o,Fec Polynesia, French Pou, Ua (PTBG) aC long gapCp G539 G530 ora2: Moorea MG550310; MG550309, it 1145 Nitta irsrmspectrum Microsorum GE) Q413 DQ642247; DQ642163, (GOET), irsrmnovaezealandiae Microsorum aC long gapCp mr15 Amer Bkr Copel. (Baker) G)Haie rnhPoly- French Huahine, (GH) cwrfgrsn cl.BGG) (cult. s.n. Schwertfeger un250 Dunn ..Cu&ZR He Z.R. & Chu W.M. R 3534 LRP hn 4194 Zhang irsrmscolopendria Microsorum 3 R 3573 LRP aC short gapCp G591 MG451962, MG451961, G535 MG550306. MG550305, TNS759256 idl 1: Tindale G593 MG451974, MG451973, aC long gapCp G)Moe,French Moorea, (GH) tohieaense od15756 Wood G)Tht,French Tahiti, (GH) GE) DQ164444, (GOET), irsrmpustu- Microsorum irsrmmem- Microsorum irsrmsp Microsorum MG451963, PB)U Pou, Ua (PTBG) WL)Pohan- (WELT) Kuf)Copel. (Kaulf.) it 654 Nitta — WL)Wel- (WELT) cnie s.n. Schneider P)China, (PE) , MG451965, MG550304. TS Oki- (TNS) aC long gapCp J.H.Nitta (GOET), (GOET), (Kaulf.) (PTBG) gapCp gapCp GLu SG (GH) (UC) Wu — C. , 9168 illosa KY931349. KY931051, KBG) AY459183. cnie ..(ut BGG) (cult. s.n. Schneider EU483068. EU482972, Q521 4: GQ256251; 3: EU483071; EU482975, KY931409. KY931130, Ching (Hook.) Vietnam, Thien-Hue, hn,H570,H571;4: HQ597018; HQ597009, China, KBG) 3: HQ597020; HQ597011, China, Yunnan, (KUN) Q900 HQ597019. HQ597010, RGBE) triglossa (cult. s.n. Nicholson GQ256246. GQ256318, 1: Hett. vieillardii noei,E427,EU483067. EU482970, Indonesia, whiteheadii Microsorum 2: DQ179644; hn 3100 Zhang CB,M,VM)HaBn,Venm Y316 KY931398. KY931116, Vietnam, piloselloides Binh, Hoa VNMN) MO, (CDBI, cambricum Polypodium BGG) wandae (cult. GG0411 Kreier DQ164516. hillii GQ256200. Platycerium DQ642157, Schweinf. Peninsula, Malay 4: EU483052. DQ642242; DQ642159, Taiwan, J17 Lu 1: Wang Li GQ256247. GQ256319, Korea, South (PE) ei,M423,MG427076. MG452031, nesia, 2: Moorea MG550315; MG550314, MG427077, GU126731. AY083639. AF470336, zippelii Madagascar, unknown) (herbarium aC short gapCp i 028(ut KBG) (cult. 2012-8 Kim cnie ..(ut BGG) (cult. s.n. Schneider CB)Gihu hn,K917,KY931449. KY931175, China, Guizhou, (CDBI) KN hn,J130,J139;3: JX103790; JX103706, China, (KUN) PU unn hn,A755,A755;3: AY725052; AY725056, China, Yunnan, (PYU) KN hn,J131,J139;2: JX103794; JX103710, China, (KUN) Bue J.Sm. (Blume) Bue iWang Li (Blume) Racib. Bkr Ching (Baker) rirG00 cl.BGG) (cult. GG0405 Kreier i 025(ut KBG) (cult. 2012-5 Kim Mt. oe.1: Copel. (Mett.) aarmalongifolia Paragramma rcoeiimnormale Tricholepidium elpsrspappei Neolepisorus ltcru stemaria Platycerium P)Cia U894 EU483070. EU482974, China, (PE) KJ748235. aC long gapCp L)M.G.Price (L.) ..D 04(ut KBG) (cult. 1004 Du X.Y. cnie ..(ut DBG) (cult. s.n. Schneider hn ta.6372 al. et Zhang rirG01 cl.BGG) (cult. GG0414 Kreier T.Moore yrsatonkinensis Pyrrosia yrsanuda Pyrrosia ecerpei palmatopedata Neocheiropteris rvnel559 Gravendeel elpsrsensatus Neolepisorus GE) Y657 Q109 2: DQ212059; AY362567, (GOET), KN hn,J132,J130;2: JX103804; JX103720, China, (KUN) G595 MG451976, MG451975, L. ..m Hoshiz. & A.R.Sm. ecerpei phyllomanes Neocheiropteris IN112 hi80596 Shui rirG00 cl.BGG) (cult. GG0407 Kreier ..Zu57112 Zhu W.M. GE) Q648 DQ164522. DQ164458, (GOET), ...Wos8-61 Woods P.J.B. hn ta.7667 al. et Zhang E,E427,EU483069. EU482973, (E), GE) Y656 DQ179643. AY362566, (GOET), irsrmvarians Microsorum mt ..(ut CAG) (cult. s.n. Smith CB,M,VM)HaBn,Vietnam, Binh, Hoa VNMN) MO, (CDBI, T)Gnn ee noei,AB232411, Indonesia, Gede, Gunung (TI) ecerpei ovata Neocheiropteris KN hn,J130,J138;2: JX103786; JX103702, China, (KUN) iWang Li Bav)Ds.var. Desv. (Beauv.) it 1040 Nitta Bue T.Moore (Blume) hn 3446 Zhang hn 4482 Zhang DDn hn 1: Ching (D.Don) Geeh)Ching (Giesenh.) P)Yna,Cia GQ256323, China, Yunnan, (PE) KN,H571,H572;5: HQ597022; HQ597012, (KUN), L aa noei,AY459175, Indonesia, Java, (L) GE) Q649 DQ164513. DQ164449, (GOET), elpsrsfortunei Neolepisorus GE) Q765 DQ179645. DQ179635, (GOET), GE) Q642 DQ164526. DQ164462, (GOET), ..D 0961 Du X.Y. cep 4/9(B 901312) (LBG 441/79 Schelpe Geeh)Ching (Giesenh.) htha s.n. Whitehead G)Moe,Fec Poly- French Moorea, (GH) Tub)Ching (Thunb.) E aeca Spain, Valencia, (E) CB,M,VM)Thua VNMN) MO, (CDBI, ltcru elephantotis Platycerium KN unn China, Yunnan, (KUN) yrsasubfurfuracea Pyrrosia hnS4-1 Shen aC short gapCp hn 728/1 Zhang P)Cia EU482955, China, (PE) P)Yna,China, Yunnan, (PE) Mt. enpa & Hennipman (Mett.) akr2087 Ranker GE) DQ164452, (GOET), rnilBF012 Cranfill hlcpei pap- Thylacopteris laurentii U) DQ179634, (UC), (F´ i 024(cult. 2012-4 Kim Bkr hit1: Christ (Baker) hn ta.6295 al. et Zhang i 021(cult. 2012-1 Kim KN Yunnan, (KUN) Neocheiropteris Crs)Ching (Christ) e Fraser-Jenk. ee) U)Sumatra, (UC) Neolepisorus ..D 0936 Du X.Y. Platycerium P)China, (PE) Microsorum P)China, (PE) hn 3611 Zhang hn tal. et Zhang MG550313, (T.Moore) eWild. De Pyrrosia (COLO) — (UC) , SG — ,